SURICATA 6

Conservation assessment of Scarabaeine

DUNG BEETLES
in South Africa, Botswana and Namibia:
IUCN Red List categories,
atlas and ecological notes

by:
Adrian L.V. Davis
Christian M. Deschodt
&
Clarke H. Scholtz

Pretoria
2020
 SURICATA
Suricata is the genus name of the suricate (meerkat), which is near-endemic to the arid western parts of southern Africa
(occurring in Namibia, South Africa, Botswana; and just entering into a small area in the extreme south of Angola).
Behaviourally, suricates are socially inclusive and innately inquisitive, symbolising the commitment of the South African
National Biodiversity Institute (SANBI) to include all biodiversity and serve all of Africa, and the scientific curiosity that
precedes and drives research and publication of research results. Sister journal to SANBI’s Strelitzia, Suricata is a peer-
reviewed journal and publishes original research such as monographs, revisions, checklists, Red Lists, Atlases and Faunas
of any taxa belonging to Regnum Animalia (the Animal Kingdom).

By Adrian L.V. Davis, Christian M. Deschodt & Clarke H. Scholtz

Editor: Yolande Steenkamp

Proofreader: Alicia Grobler
Design & layout: Elizma Fouché
Cover photographs: Christian M. Deschodt

Recommended citation: Davis, A.L.V., Deschodt, C.M. & Scholtz, C.H. 2020. Conservation assessment of
Scarabaeine dung beetles in South Africa, Botswana and Namibia: IUCN Red List categories, atlas and ecological
notes. Suricata 6. South African National Biodiversity Institute, Pretoria.

ISBN: 978-1-928224-39-6

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Copyright © 2020 by South African National Biodiversity Institute (SANBI)

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The views and opinions expressed do not necessarily reflect those of SANBI or the editors. The authors and publisher have
made their best efforts to prepare this book and make no representation or warranties of any kind with regard to the com-
pleteness or accuracy of the contents herein. All images in this book have been reproduced with the knowledge and prior
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of copyright or otherwise arising from the contents of this publication. Every effort has been made to ensure that the
credits accurately comply with the information supplied by the authors.
SURICATA 6 (2020) iii

CONTENTS
Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv
Preamble . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
A. Importance of group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
B. Data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
C. Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
D. Distribution data and methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
E. Ecological associations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
F. Temporal associations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
G. Conservation status and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
H. Potential threats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
I. Use as indicators. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
J. Format of species accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Subfamily SCARABAEINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Tribe ATEUCHINI: . . . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 20

Tribe CANTHONINI (DELTOCHILINI): . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 56

Tribe COPRINI:. . . . . . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 164

Tribe GYMNOPLEURINI: . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 262

Tribe ONITICELLINI: . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 288

Tribe ONITINI: . . . . . . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 332

Tribe ONTHOPHAGINI: . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 394

Tribe SCARABAEINI: . . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 556

Tribe SISYPHINI:. . . . . . . . . . . . . . . . . . . . . . . tribal, generic and species accounts . . . . . . . . . . . . . . . . . . 660

Poorly known species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690

SYNTHESIS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 698
A. Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 698
B. Cladistic groups. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 700
C. Major spatial patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 710
D. Climate, sampling and geographical bias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 712
E. Climate, endemism and conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 715
Appendix 1: Summary Table 1. South Africa: species distribution across biomes, conservation status and
ecological notes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 718
Appendix 2: Summary Table 2. Botswana and Namibia: species distribution across ecoregions and conser-
vation status. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 738
Appendix 3: Summary of average climatic, topographical and distributional data . . . . . . . . . . . . . . . . . . . . . . . . 747
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 765
Glossary of scientific terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 780
Taxonomic index of African taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 789
iv SURICATA 6 (2020)

Acknowledgements
We are grateful to the South African National Biodi- validated material into the dung beetle database, partic-
versity Institute (SANBI) for funding this book. We ularly Dr Werner Strümpher, Carmen Jacobs, Angelika
also thank the JRS Biodiversity Foundation for funding Loots and Susana das Neves. Many, difficult to obtain,
the dung beetle database of museum reference material papers with original species decriptions and published
from which much of the geographical distribution data type localities were kindly provided by Paul School-
was drawn. Prof. Mervyn Mansell and Brian Kenyon are meesters. We thank Antoine Mantilleri of the Muséum
thanked for maintaining the database. For organising National d’Histoire Naturelle, Paris, for the photographs
loans of reference material entered into the database, we of Xinidium howdeni [(c) MNHN/A. Mantilleri.]. All
thank museum scientific staff and curators, particularly, remaining species photographs were taken by Christian
Ruth Müller (Ditsong Museum), Elizabeth Grobbelaar Deschodt using professional hardware and software. Dr
and Riaan Stals (National Collection of Insects). In addi- Dawid Jacobs and Lukas Niemand are thanked for loan-
tion, we thank Philippe Moretto for loaning a specimen ing specialised photographic equipment. Elizma Fouché
of the rare species, Kolbeellus ateuchoides. Validation of is thanked for the design and layout of this publication.
species identity for museum reference material was most- Finally, we thank Prof. Catherine Sole for logistic support
ly conducted by one of the authors (Dr Adrian Davis) after taking over the leadership of the SRG upon the re-
except for the genus, Odontoloma, for which we thank Dr tirement of Prof. Clarke Scholtz. Subsequently, the SRG
Werner Strümpher. We are grateful to staff of the Scarab was dissolved by the University of Pretoria at the end of
Research Group (SRG) at the Department of Zoology & 2018 and replaced by the Invertebrate Systematics and
Entomology, University of Pretoria, for entering data of Conservation Group.
SURICATA 6 (2020) v

Preamble
Around three years elapsed between completion (November 2017), review and publication of the book. As taxonomic
revision is an ongoing process, a number of new tribes, genera, species and synonymies were described over those three
years. For the most part, accounts for these new taxa were not included in the book as extensive changes would have been
required to sections that summarise the composition and distribution of the southern African dung beetle fauna. However,
the new taxa and changes in status are listed below together with citations to the publications in which they were described
(see References).

New tribes
1. Some of the clades defined as ‘Basal Scarabaeinae’ within the most recent global phylogeny for the subfamily
Scarabaeinae (Tarasov & Dimitrov 2016) have subsequently been described as new tribes (Davis et al. 2019) al-
though, here, they remain listed within the tribe Canthonini (by common usage) or Deltochilini (by precedence)
pending a full revision of tribal classification; see, also, introduction to the tribe Canthonini: Clades 1A, 1B, 1C,
as well as notes in accounts for affected genera.
1A: Tribe Odontolomini Davis, Deschodt & Scholtz, 2019 (one genus: centred in southern Africa).
1B: Tribe Byrrhidiini Davis, Deschodt & Scholtz, 2019 (seven arid SW African genera: South Africa, Namibia).
1C: Tribe Endroedyolini Davis, Deschodt & Scholtz, 2019 (seven SE African forest genera: South Africa).
2. See recent revisions of tribal, generic and subgeneric composition (Tarasov & Dimitrov 2016; Daniel et al. 2020)
updated in the introductions to the genus Epirinus Dejean, 1833, and the tribe, Sisyphini Mulsant, 1842; also
noted in generic and species accounts for the Sisyphini.
3. New tribe pending (Catharsiini Takano) for the genera, Catharsius Hope, 1838, and Metacatharsius Montreuil,
1998 (see Takano 2018, unpublished).
vi SURICATA 6 (2020)

New genera, species and synonyms in

southern Africa plus missed species
1. Catharsius laticeps Boheman, 1857: missed sand and dune forest species of NE KwaZulu-Natal and SE Mozam-
bique coast (see notes in species account).
2. Proagoderus gangloffi Josso & Prévost, 2001: missed elephant dung specialist species of E Kaokoveld, N Namibia.
3. New genus in the tribe Onthophagini Burmeister, 1846 (Dierkens et al. 2017); Jossonthophagus Dierkens, Moretto
& Génier, 2017, accommodates the species of Onthophagus Group 14; no species accounts for two species recorded
in the target area of southern Africa.
4. New genus and species from arid SW Namibia (Deschodt & Davis 2018) in the new flightless tribe, Byrrhidiini
Davis, Deschodt & Scholtz, 2019 (formerly incorporated into tribe, Canthonini).
a. Ausmontins Deschodt & Davis, 2018, and Ausmontins jacobsi Deschodt & Davis, 2018, from near Aus.
b. Namakwanus kamfferi Deschodt & Davis, 2018, from Bloedkoppe, Namib Desert.
c. Namakwanus minutus Deschodt & Davis, 2018, from the Nubib Mountains.
d. Namaphilus nubibmontanus Deschodt & Davis, 2018, from the Nubib Mountains.
e. Namaphilus tirasmontanus Deschodt & Davis, 2018, from the Tiras Mountains.
5. New taxa in the Garreta lugens (Fairmaire, 1891) species group (Davis & Deschodt 2018).
a. Garreta namalugens Davis & Deschodt, 2018, from the rocky W mountain escarpment of Namibia.
b. See species account for Garreta australugens Davis & Deschodt, 2018, from SE Africa, recently separated from
Garreta lugens (Fairmaire, 1891), from East Africa.
6. New species of the flightless genus, Macroderes Westwood, 1842, from SW South Africa (Abdalla et al. 2018);
includes additional distribution records for previously described species.
a. Macroderes cederbergensis Abdalla & Deschodt, 2018, from the Cederberg, SW Western Cape.
b. Macroderes gifboomi Abdalla & Deschodt, 2018, from Gifberg near Vanrhynsdorp, SW Western Cape.
c. Macroderes leipoldti Abdalla & Deschodt, 2018, from the Hantamsberg, SW Western Cape.
d. Macroderes oreatus Abdalla & Deschodt, 2018, from Richtersveld National Park, Namaqualand.
e. Macroderes porselinus Abdalla, 2018, from Porseleinberg, SW Western Cape.
f. Macroderes soleiana Abdalla & Deschodt, 2018, from the Cederberg, SW Western Cape.
g. Macroderes tortuosus Abdalla & Deschodt, 2018, from the Gifberg near Vanrhynsdorp, SW Western Cape.
h. Validity confirmed for Macroderes pristinus Sharp 1880, from Diamondfields, Namaqualand.
7. New species of southern African Sisyphus Latreille, 1807 (Daniel et al. 2018).
a. Sisyphus swazi Daniel & Davis, 2018, from forest in Eswatini (formerly Swaziland).
b. Sisyphus inconspicuus Daniel & Davis, 2018, from shaded vegetation in SE Africa: South Africa, Mozambique.
c. Sisyphus australis Daniel & Davis, 2018, from shaded vegetation on the south coast of South Africa.
8. Review of Scarabaeolus (Balthasar, 1965a), by Zídek and Pokorný (2018) including new southern African species
with Scarabaeolus treated as a subgenus of Scarabaeus Linnaeus, 1758.
a. Scarabaeolus krugeri (Zídek & Pokorný, 2018) from Kruger National Park, South Africa (=unpublished synonym
of Scarabaeolus planipennis (Davis & Deschodt, 2015)).
b. Scarabaeolus lizleri (Zídek & Pokorný, 2018), which separates NW South African material from the Namib Des-
ert dune field species, Scarabaeolus rubripennis (Boheman, 1860); S. lizleri material included, here, in the species
account for S. rubripennis.
c. Scarabaelous namibensis (Zídek & Pokorný, 2018) from the central Namib Desert and Kaokoveld, Namibia (see
species account).
d. Scarabaeolus fragilis (Zídek & Pokorný, 2018) from Springbok, Namaqualand, South Africa.
e. Scarabaeolus rugosipennis (Zídek & Pokorný, 2018) from Makgaberg Plateau, Limpopo, South Africa; needs to be
re-examined as a probable synonym of Scarabaeolus niemandi (Deschodt & Davis, 2015).
f. Scarabaeolus cunene (Zídek & Pokorný, 2018) possibly from E Hartmann dune field, N Namibia.
g. Scarabaeolus orientalis (Zídek & Pokorný, 2018) from farm Rhenosterpoort, Gauteng, South Africa (=unpublished
synonym of Scarabaeus karae Davis & Deschodt, 2017).
SURICATA 6 (2020) vii

h. Scarabaeolus similis (Zídek & Pokorný, 2018) from Pienaars River, Gauteng, South Africa.
i. Again, incorrectly synonymises Scarabaeolus reichei (Waterhouse, 1890) with Scarabaeolus canaliculatus (Fairmaire,
1888); in this book, the two species are treated as valid, S. canaliculatus from the southern Namib Desert and
S. reichei from the W Coast of South Africa.
9. Review of African Garreta Janssens, 1940a (Pokorný & Zídek 2018); notes of relevance to southern Africa:
a. Overturns the Moretto and Génier (2015) revalidation of SE African Garreta wahlbergi Boheman, 1857, by syn-
onymising it, once more, with widespread African species, Garreta nitens (Olivier, 1789).
b. Cites Garreta fastiditus (Fahraeus, 1857) as a valid South African species although we consider its type locality
suspect and provide no species account.
c. Extends range of Garreta caffer (Fahraeus, 1857) across that occupied by subspecies Garreta laetus olivaceus
(Quedenfeldt, 1883), which is synonymised with Garreta laetus (Hope, 1842).
10. New genus in the tribe, Onthophagini Burmeister, 1846 (Roggero et al. 2019); transfer of Onthophagus species
to the new genus, Tiaronthophagus Roggero, Moretto, Palestrini & Barbero, 2019 (see notes under Group 24 in
introduction to the genus, Onthophagus Latreille, 1802); species in South Africa and Botswana only:
a. Tiaronthophagus ebenus (Péringuey, 1888) (see species account under Onthophagus ebenus).
b. No species account for Tiaronthophagus aequatus (Onthophagus aequatus Péringuey, 1900) recorded in Botswana.
11. New species and synonyms of Epirinus Dejean, 1833, from South Africa and a new record from Namibia (Deschodt
et al. 2019).
a. Epirinus inparrugosus Deschodt & Davis, 2019, from Blesberg-W, Swartberg, Eastern Cape.
b. Epirinus jacobsae Deschodt & Davis, 2019, from Mapelane, NE coastal KwaZulu-Natal.
c. Epirinus muellerae Deschodt & Davis, 2019, from central Drakensberg and E central Lesotho.
d. Epirinus pseudorelictus Deschodt & Davis, 2019, from coastal NE Eastern Cape.
e. Epirinus schoolmeestersi Deschodt & Davis, 2019, from Baviaanskloofberg, Eastern Cape.
f. See species accounts for Epirinus hluhluwensis Medina & Scholtz, 2005, and Epirinus ngomae Medina & Scholtz,
2005, which are retained even though they have, now, been synonymised with Epirinus davisi Scholtz & Howden,
1987, whose distribution records have been expanded to Eswatini.
g. See species account for Epirinus flagellatus (Fabricius, 1775) for which a new distribution record expands the ge-
neric distibution to N Namibia (Okonjima Nature Reserve).
12. New species and a revalidation from synonymy in South African Gyronotus van Lansberge, 1874a (Deschodt &
Davis 2019).
a. Gyronotus dracomontanus Deschodt & Davis, 2019, from the central Drakensberg grasslands near Bergville.
b. Gyronotus ovalis Deschodt & Davis, 2019, from grassland at the edge of Nkandla Forest, Zululand.
c. Gyronutus kearneyorum Deschodt & Davis, 2019, from grassland on Mount Sheba, Mpumalanga.
d. Revalidation of Gyronotus marginatus Péringuey, 1888, from coastal forest near East London, Eastern Cape, which
is removed from synonymy with Gyronotus pumilus (Boheman, 1857), but remains included in the species account
for that species in this book.
13. Subgenus Hyalonthophagus Palestrini & Giacone, 1988, formally raised to genus status with description of a new
species (Deschodt et al. 2019).
a. Hyalonthophagus pulcher Deschodt & Davis, 2019, from the Northern Cape, South Africa.
14. New species of Versicorpus Deschodt, Davis & Scholtz, 2011, from Namibia (Deschodt & Sole 2019).
a. Versicorpus daures Deschodt, 2019, from Brandberg (mountain) in W central Namibia.

Exceptions
The taxonomic updates to current nomenclature effected during the review of this atlas (see above) are also noted in the
generic and species accounts where applicable. However, in a few further species accounts, taxonomic treatment also di-
verges from the currently accepted nomenclature. These species are listed below.
1. Euonthophagus vicarius (Péringuey, 1901) is treated here as a valid species although it is currently still listed as a
synonym of Euonthophagus carbonarius (Klug, 1855).
viii SURICATA 6 (2020)

2. Gymnopleurus pumilus Reiche, 1850 is treated here as a tentatively valid species although it is currently still listed
as a synonym of Gymnopleurus virens Erichson, 1843.
3. Kheper vethi (van Lansberge, 1886) is treated here as a species of Kheper Janssens, 1940b although it currently
remains classified as Scarabaeus vethi van Lansberge, 1886.
4. Onthophagus cretus Péringuey, 1901 is treated here as a valid species although it is currently still listed as a synonym
of a poorly known species, Onthophagus setosus Fahraeus, 1857.
5. Onthophagus peringueyi Shipp, 1895 is currently classified as a synonym of Onthophagus suturalis Péringuey, 1888,
which we consider a synonym of Onthophagus pallidipennis Fahraeus, 1857 along with another currently valid
species, Onthophagus politissimus d’Orbigny, 1908.
6. Onthophagus venustulus Erichson, 1843 is treated here as a valid species although it is currently still listed as a syn-
onym of Onthophagus variegatus (Fabricius, 1798).
SURICATA 6 (2020) 1

INTRODUCTION
A. IMPORTANCE OF GROUP
Scarabaeine dung beetles (Order: Coleoptera; Family: of habitat modification, habitat fragmentation and loss of
Scarabaeidae; Subfamily: Scarabaeinae) have received mammals producing preferred dung types. Therefore, an
a great deal of attention in recent decades. Much of the atlas of the dung beetle species found in South Africa, Bot­
attention stems from programmes to augment control of swana and Namibia, is long overdue. This atlas provides
dung and dung-breeding flies in farm rangeland by intro- photographs of over 540 species together with notes on
ducing European and African dung beetle taxa into Aus- their taxonomy, distribution, ecology and conservation
tralia, the Americas and, lately, New Zealand. Research on status. As the alpha taxonomy is incomplete, the atlas is
dung beetle ecology to assist selection of species for intro- also incomplete, although it does comprise accounts for
duction has, subsequently, developed in several directions. most species in eight out of nine tribes of dung beetles
Dung beetles have been widely used to bioassay non-target found in the target region (Onthophagini excepted). Al-
effects of livestock pesticides in farm rangeland. They have though numbers of species and genera are quoted for the
also been widely used as biological indicators in ecologi- nine tribes, it should be noted that these are frequently
cal impact assessments (EIA) and studies of farm health, changing due to revisions of pre-existing nomenclature or
as well as in conservation studies investigating the effects addition of newly described genera and species.

B. DATA

Data sources Ditsong (formerly Transvaal) Museum (DM), (B) the

National Collection of Insects (NCI), and (C) the Scarab
Dung beetle data used in the atlas were drawn from (1) Research Group at the University of Pretoria (UP – now
a web-based catalogue of species, (2) museum records, part of the reference collection at the Iziko South African
and (3) both published and unpublished field records Museum in Cape Town). (3) Most qualitative field data
for distribution and ecological associations. (1) Web- were drawn from unpublished observations of soil, veg-
based species data were drawn from the Catalogue of Life etation and food type made between 1970 and 1986 by
(Schoolmeesters 2017). (2) Museum records were drawn personnel of the Australian CSIRO Dung Beetle Research
from three extensive, local, reference collections during Unit (DBRU) based in Pretoria. Most quantitative field
digital databasing (see pg. 2). These were those of (A) the data were drawn from unpublished or published studies
2 SURICATA 6 (2020)

made on regional and local dung beetle faunas of South Unpublished data (mostly
Africa and Botswana between 1983 and 2016 (see list un-
South Africa)
der Field surveys).
1. ‘DBRU collection records’: Qualitative data derived
from observations of soil, vegetation and dung types
Digital database at > 2 330 dated collection localities contained in
the dung beetle reference collection of the DBRU
Identities of dung beetles from the reference collections
(see pg. 1 in Data sources). This collection is now
of the three museums (DM, NCI, UP: see Data sources
part of the NCI and the locality records form part
from pg. 1) were validated to species level and the infor-
of the digital database (1970 to 1986).
mation on attached labels was digitised. The species in-
cluded in the database were restricted to those with val- 2. ‘In Western Cape’ or ‘in SW Cape’: Quantitative
idated published names. Information was derived from data recorded to cattle dung over one year on dif-
examination of 91 225 museum specimens in total. The ferent soil and vegetation types at 11 survey sites on
database has been uploaded onto the website of the An- farms and reserves near Cape Town (‘In Cape of
imal Demography Unit at the University of Cape Town Good Hope Nature Reserve’; Farms: ‘At Bonne At-
(click on dung beetle icon at: http://vmus.adu.org.za/) tente, Modderrivier, Geelbek, Waylands, Oranjefon-
and will also be available on the website of the South Af- tein, Pampoenvlei or Groote Post’) (1987 to 1988).
rican National Biodiversity Institute (SANBI). These re- 3. ‘In Ithala Game Reserve’: Quantitative data re-
cords for 555 species (two synonymised post 2017) form corded on sandy loam at different altitudes (450–
the core distribution data used for 542 species accounts 1 320 m) in different vegetation types (forest, shrub/
in this dung beetle atlas and assessment of conservation woodland, grassland) using three different dung
status. types (cattle, horse, pig) (KwaZulu-Natal) (1999).
4. ‘Near Carolina’: Quantitative data recorded to cat-
Data usage tle dung on the farm Foggy Valley near Carolina
(Mpumalanga) from finer-grained soils in natural,
The data were used in various ways. The web-based cat- Themeda-dominated grassland, regenerating fallow
alogue was used to support the cited species names and crop fields, and improved Kikuyu pasture (2001 to
their overall geographical range across countries of Africa. 2002).
Museum locality records were used to plot geographical 5. ‘At Viljoenskroon’: Quantitative data recorded to
distributions across the target countries of South Africa, cattle dung on sandy soils of Farm Huntersvlei near
Botswana and Namibia as well as most of Zimbabwe and Viljoenskroon (N Free State) in a crop field (sor-
the extreme south of Mozambique. Dates of collection ghum), exotic grassland pastures (Eragrostis, Smuts
for museum specimens and field data were used to assess finger) and natural grassland (2002).
months of activity by adults. Quantitative field survey
6. ‘On Mariepskop’: Quantitative data recorded from
data were used to augment plots of geographical distri-
two dung types (cattle, pig) on finer-grained soils
bution and to assess habitat, trophic and temporal associ-
in natural grassland or forest on the E escarpment
ations in studied species. Qualitative field survey records
of Mpumalanga (2004).
(DBRU) were used for comparison or support of quanti-
tative data, or as the sole assessment of associations shown 7. ‘At Wolkberg’: Quantitative data recorded from
by poorly studied species. two dung types (cattle, pig) on finer-grained soils
at different altitudes in natural grassland or forest of
Lekgalameetse Nature Reserve (Limpopo) (2005).
Field surveys 8. ‘In Maputo Special Reserve’: Quantitative data re-
corded to pig dung on deep sands in Maputo Ele-
Field surveys provide locality records for plotting geo- phant Reserve (S Mozambique) from grasslands of
graphical distributions and environmental records for fossil lagoons, coastal dune forest, and small, medi-
assessing ecological and temporal associations of dung um or large sand forest patches (2006).
beetle species within southern Africa south of 15°S. The
records comprise both unpublished and published data
and are cited liberally in many species accounts using the Published data (mostly South Africa)
initial expressions quoted in the brief descriptions below.
In these citations, whole numbers represent totals, deci- 1. ‘At Bushlands Halt’: Quantitative data from coastal
mal fractions represent averages per sample. KwaZulu-Natal for succession on dung over time
SURICATA 6 (2020) 3

(Doube et al. 1988) and associations with two soil the mesic NE (Chobe National Park) to the arid
and vegetation types (Giller & Doube 1994). SW Botswana Kalahari summarised as total num-
bers for six localities or five food types in supple-
2. ‘Measured...flight activity’: Experimental measure-
mentary tables stored on journal websites.
ment of flight periodicity in some Onitis species
(Caveney et al. 1989). 9. ‘At Phalaborwa’: Quantitative data for habitat (sev-
en categories), trophic associations (three dung
3. ‘In uMkhuze Game Reserve’: Quantitative habitat
types) and short-term temporal responses to weath-
data for four soil types and two categories of veg-
er conditions (three different) (Davis et al. 2014)
etative cover in uMkhuze Game Reserve (Doube
cited as totals in a supplementary table stored on
1991) summarised as means per sample in Appen-
the journal website.
dix B.8. of Dung Beetle Ecology.
4. ‘Along a gradsect (500 m to 2 800 m on latitude
29–30°S)’: Quantitative data recorded at 400–500 Mixed data (South Africa)
m intervals from coastal hills across the Highveld
to the top of Sani Pass in KwaZulu-Natal/Lesotho, 1. ‘In Gauteng’ or ‘in Gauteng bushveld’: Quantitative
primarily in grassland on sandy clay loam. Data re- data for diel periodicity (Davis 1996a), seasonal ac-
ported as means per sample by Davis et al. (1999a). tivity (unpublished; 1983 to 1984), or associations
with six habitat types near Pretoria (Davis 1996b).
5. ‘At Richards Bay’: Quantitative data recorded across Habitat and seasonal data recorded over one year.
a post-mining chronosequence of regenerating veg- Habitat summarised as total numbers across two
etation (grassland, to dense woodland to open un- soil and three vegetation types.
derstorey woodland to natural coastal dune forest)
reported as totals per vegetation type (Davis et al. 2. ‘In Abe Bailey NR’, ‘Leeuwfontein NR’, ‘Roode-
2002) or means per sample in supplementary tables plaat NR’, ‘Suikerbosrand NR’, ‘Telperion NR’ or
stored on the journal website (Davis et al. 2013). ‘Tswaing NR’: Quantitative data derived from dif-
ferent altitudes on different soil and vegetation types
6. Genus Pachysoma: Field observations reported by
forming part of a quantitative survey of six Gauteng
Harrison et al. (2003).
nature reserves. Data derived from raw unpublished
7. ‘At SA Wildlife College’: Quantitative data for two records or means per reserve in Davis et al. (2005).
soil types in the Kruger National Park and nearby
3. ‘In Northern Cape’: Quantitative data from obser-
farm rangeland summarised as means per sample
vations of soil type across five defined regions (Da-
(Davis et al. 2012).
vis et al. 2016) with mean numbers per soil type
8. ‘In Botswana’: Quantitative habitat (Tshikae et derived from unpublished raw data or mean data
al. 2013a) and trophic data (Tshikae et al. 2013b) per region cited in a supplementary table stored on
from deep sands along a 1 100 km gradsect from the journal website.

C. TAXONOMY
Nomenclature has been thoroughly checked. Subfamily, During databasing, it has become clear that a number of
tribal and generic names are mostly based on recent reviews uncorrected errors persist after over two centuries of work
of their validity (Smith 2006, 2009; Branco et al. 2007, on the systematics and taxonomy of the southern African
2011; Bouchard et al. 2011). Species names are mostly dung beetle fauna. The greatest number of errors occurs
those listed by the Catalogue of Life (Schoolmeesters 2017) in the Onthophagini, which is the largest tribe. It, thus,
as ‘Accepted species’ or as published synonyms. They are, includes the greatest number of species omitted from the
therefore, considered relatively well-validated on the as- book because of doubtful taxonomic identity. A total of
sumption that we have correctly matched the species names 49 currently valid species described from South Africa,
with reference material used to map distributions. Fur- Botswana and/or Namibia could not be reliably matched
thermore, the names listed by Schoolmeesters are regularly to reference material. Available information on these
updated to remove errors, accommodate revisions and add species is provided in a separate section entitled Poorly
newly described taxa. Nevertheless, some of the included known species. A further eight Onthophagus species were
species and/or their synonyms require further investigation. not considered as their recorded occurrence in South
4 SURICATA 6 (2020)

Africa, Botswana or Namibia could not be validated. 4. Synonyms containing more than one species in
Additionally, no consideration was made of many other the type series. Such names are marked as ‘pars’,
primarily onthophagine species as they probably remain e.g. Phalops adspersipennis Boheman, 1857, sensu
undescribed and thus lack valid names. Péringuey, 1901 (pars), of which only part of the
type series is a synonym of Phalops rufosignatus van
Although most included species are considered reasonably Lansberge, 1885a.
well-validated, possible taxonomic or nomenclatural prob-
5. Dung beetle species showing different irridescent
lems are recorded in about 10% of the species accounts.
colours (cupreous, green or blue) produced by multi-
These possible problems are variously dealt with as either
layer interference within the exoskeleton. In the past,
single species or putative complexes of two or more spe-
such intraspecific variation was often recognised
cies on the same page. Unresolved problems include: valid
with published names, e.g. Onthophagus (Phalops)
species names synonymised in error; synonymous species
ardea Klug, 1855, var. chloritus d’Orbigny, 1902.
names that deserve the status of valid species; type series
However, these varietal names are now recognised as
containing more than a single species; undescribed or de-
synonyms of the original species description, e.g. the
scribed species misidentified in the published literature.
green var. chloritus d’Orbigny, 1902, is a synonym of
the cupreous Phalops ardea (Klug, 1855) (see School-
A start has been made to revising the alpha taxonomy by
meesters 2017: Catalogue of Life).
describing new taxa and correcting errors. So far, effort has
concentrated mainly on the tribe, Scarabaeini although
new species have also been described in other tribes since Photographs
2017 (see Preamble). However, many more years of work
are required to revise the entire membership of the sub- A representative photograph is provided for each species.
family Scarabaeinae in the subcontinent. Thus, the species Because of space limitations, it has been possible to develop
included in this book probably comprise only an estimat- only a limited photographic library. This is not a problem
ed 80%, or so, of the scarabaeine fauna of South Africa, with species that show minor external sexual dimorphism.
Botswana and Namibia. However, many taxa show minor to extreme dimorphism
coupled with great morphological variation according to
body size (Figure 1). This means that the featured photo-
Valid names and synonyms graphs of so-called major males are not necessarily a good
guide for identifying all individuals of a species in six of
Most species accounts are headed by the valid species
the nine scarabaeine tribes represented in Africa. Although
name together with the author and year of description.
photographs of major females are also provided for those
They are followed by synonyms listed chronologically
species showing the most extreme sexual dimorphism,
under the generic name used in the original description.
there is insufficient space to depict the full range of mor-
Many names have since changed, particularly those de-
phological variation shown by such species.
scribed before 1900. In a few cases, names of synonyms
head the species accounts where it is considered that re- The photographs may be a reliable identification guide for
vision would result in formal revalidation. The absence of most species in the Canthonini, Scarabaeini and Gymno-
synonymous names is marked as ‘No synonyms’. pleurini, as they usually show minor external differences
between the sexes. The photo images are also helpful for
There are five different classes of synonyms.
identifying species in the tribes that mostly show limited
1. Subsequent redescriptions of the same species un- sexual dimorphism (Ateuchini, Sisyphini). Female imag-
der different species names. es are provided especially for those species in tribes that
2. Invalid duplication of names already used for dif- include genera or species groups showing extreme sexual
ferent species in the same genus at an earlier date dimorphism, i.e. a few Onitini and Oniticellini plus many
requiring creation of a new replacement name (nom Coprini and Onthophagini, particularly in the genera:
Copris, Catharsius, Heliocopris and some Proagoderus plus
nov.).
Onthophagus.
3. Citations of species described by earlier authors that
were misidentified by later authors. Such species mis- The wide range of morphological variation in sexually di-
identifications are marked as sensu the author respon- morphic species is correlated to body size. Depending on
sible for the error following the name and author of the taxon, horns and sculpturing on the head, protuber-
the cited species, e.g. Sebasteos westwoodi (Harold, ances on the prothoracic disc and/or shape and dentition
1869a) sensu Péringuey, 1908, which is a synonym of of the legs, show great prominence in larger-bodied, ma-
Scarabaeus geminogalenus Davis & Deschodt, 2016. jor males and females. These features change in shape and
SURICATA 6 (2020) 5

a e

b f

c g

d h

5 mm

♂ ♀
Figure 1. Extreme morphological variation related to body size in Copris elphenor Klug, 1855. So-called major to minor males (A–D)
and major to minor females (E–H).
6 SURICATA 6 (2020)

decline in prominence with smaller body size to a point Sometimes, subsequent publications create lectotypes or
where minor males and females may be close to one anoth- neotypes if original type material was inexactly designated
er in appearance (Figure 1). Plots of body size against size or has been lost. In such cases these lectotype or neotype
of secondary sexual armature generally describe a sigmoid localities are also quoted. It should be noted that subse-
curve (Moczek 2002). The switch from major to minor quent publications sometimes cite original type localities
features, thus, occurs over a relatively small range in body according to all of the listed names, the first listed name,
size. or that on the locality labels of type specimens. Thus, in
some cases, original type localities cited here may differ
The taxonomy section in each species account identifies from those cited in later published descriptions.
sexually dimorphic species with an indication of the body
parts involved (head, prothoracic disc and/or legs). Spe- Type localities may be useful for assisting validation of
cies showing colour or colour pattern variation are also species names. However, for many older types, localities
identified together with an indication of the nature of dif- were not stated with the original descriptions or they are
ferences, particularly in iridescent species (cupreous, green particularly inaccurate, ambiguous, or even erroneous.
and/or blue). Some of the recurring inaccurate type localities cited in
the late 18th and early to mid-19th centuries comprise:
Cape of Good Hope (in various formats and languages),
Type localities Caffraria (sometimes followed by tota, interiore or even
Ngami), in terra Natalensi, juxta fluvium Limpopo or juxta
Type localities are listed in chronological order for species fluvium Gariep [KwaZulu-Natal, near Limpopo River, or
and synonyms, if any. Where a holotype was designated, near Orange River]. Caffraria is a particularly abused cita-
a single locality is cited. Where status of types was not tion. In the narrowest sense, it comprises a section of the
specified, they are regarded as syntypes and all localities SE coastal region from the Eastern Cape of South Africa
are cited. Species names and localities are as quoted in the to southern Mozambique (see species account for Kurtops
original published descriptions. Where clarification is re- caffrarius), but dung beetle distribution data suggest it has
quired, modern name changes to localities follow in square often been used much less exactly for southern Africa in
brackets along with any necessary qualifying comments. general or any area therein.

D. DISTRIBUTION DATA AND METHODS

Dung beetle distribution data have been treated in various published literature. Measurements of EOO (Extent of
ways. Species distributions have been outlined on a map Occurrence) vary from fairly accurate to gross estimates
for the whole of Africa and plotted as data points on a for species with the largest ranges. Except for Kenya and
customised map panel for southern Africa. They have also northern Tanzania, validated data were severely limited
been described according to occurrence in African countries for species with distributions continuing north of 15°S.
(Schoolmeesters 2017) and coincidence with vegetation Except in cases where a species is known from only a sin-
regions of South Africa (Mucina & Rutherford 2006) or gle locality, a map of Africa showing the overall geograph-
ecoregions of Namibia, Botswana, Zimbabwe and the ex- ical range has been placed as an inset into the map panels
treme south of Mozambique (Olson et al. 2001). Locality for species distribution in southern Africa (Figure 2). In
data for southern Africa south of 15°S have also been used general, ranges varied from very large to very small with
to calculate climatic associations (altitude, rainfall, tempera- the latter bias variously representing (1) natural localisa-
ture). tion due to specialisation, (2) range fragmentation due to
loss of preferred vegetation or dung types, or (3) a collec-
tion artefact.
Geographical range
The overall geographical range of each species in Africa Distribution in southern Africa
has been determined from two sources: (1) countries of
occurrence derived from publications (Schoolmeesters Distribution data for each validated species recorded in
2017); and (2) available distribution data. These data are South Africa, Botswana and Namibia, were plotted onto
often less than ideal. The range represented by listed coun- a customised map for southern Africa that primarily en-
tries is divided into validated occurrences, non-validated compassed an area south of 17°S and west of 33°E (Figure
citations and likely errors resulting from mistakes in the 2). The map panel also included Lesotho and Swaziland
SURICATA 6 (2020) 7

(now Eswatini), most of Zimbabwe, and small parts of 15 to 17°S and 33 to 35°E. For species known from more
Mozambique, Angola and Zambia. For each species, than a single locality, these data are expressed as a mean
distribution was plotted onto the map panel as presence ± SD and a range. The data comprise only one value per
within ~15 × 15 km2 squares that represent the subdi- 5 × 5 km polygon in order to reduce the effects of asym-
vision of degree squares of latitude and longitude into metrical collecting intensity across southern Africa.
16 ~equal parts. Most distributions are plotted using
red squares although other colours are used (black, blue, Reliable distribution data are derived from three sources:
green, grey) where (1) more than a single taxon or single 1. Localities derived from validated, databased muse-
putative taxon are plotted onto a single map, where (2) um records.
there is overlap between multiple taxa, or where (3) some
2. Localities in South Africa and Botswana derived
records are questionable.
from validated raw data in unpublished and pub-
lished studies that are cited in the Field surveys
Based on mapped distributions, 281 species have been
section.
classified as endemic since available data were restricted to
one (228), two (31) or all three of the focal countries of 3. Validated locality data derived from recent, pub-
South Africa, Botswana and Namibia (22) (Table 1). Of lished, taxonomic revisions, particularly in the
these species, a total of 200 are recorded as solely South tribes, Canthonini and Ateuchini.
African endemics. A further 261 species are not marked as
endemic, since their ranges extend beyond the borders of In general, the data for South Africa was the most compre-
any one, two or all three of the target countries. hensive (see Synthesis section) although there were also
good records derived from serial collections and quantita-
tive work in Botswana and Namibia. Good data were also
Locality data for southern Africa available for the extreme south of Mozambique, but those
for Zimbabwe were much more limited.
The ranges shown by each species in southern Africa have
been summarised in terms of altitude (m), annual rainfall
(mm) and annual temperature (°C: max. + min. / 2). These Climatic associations
data have been extracted from GIS base maps comprising
The climate of southern Africa is influenced by tempera-
interpolated average climatic data for 5 × 5 km polygons
ture variation related to a latitudinal gradient from 34°S to
using spot grid references for all specimens recorded from
17°S, variable topography, and differences in the amounts
S of 15°S. This includes data for specimens recorded in the
(<100 to >1 500 mm) and seasonality of annual rainfall.
areas laying outside of the map panel boundaries between
In general topographical terms, the subcontinent is bor-
dered by a relatively narrow strip of coastal lowlands de-
17
Zambia
Angola lineated by mountain ranges. The Cape Fold Mountain
19
Zimbabwe
Belt borders the south coast whereas the east and west
Namibia coasts are bordered by mountain escarpments that define
21
a central plateau, which is tilted from high altitude in
Mozambique

Botswana
23 the east to lower altitude in the west (Partridge & Maud
25
1987; Maud 2012). Across this area, two cells of wind
currents show seasonal expansion bringing, respectively,
27 winter rainfall from the Atlantic Ocean to the southwest
29
and summer rainfall from the Indian Ocean to the north-
Eswatini

South Africa
east (Tyson 1986). Furthermore, upwelling of cold glacial
31
waters result in a cell of dry air and a desert climate on the
Africa
33 inset
Lesotho West Coast. The adjoining central parts of the subconti-
nent also have an arid climate as the sequential expansion
35
12 14 16 18 20 22 24 26 28 30 32 of air currents bringing rainfall only reach the area late in
each rainy season. Thus, the subcontinent may be divided
Figure 2. Map panel of southern Africa onto which spot dis- into four major climatic regions (Walter & Lieth 1964).
tributions (1/16th degree squares) were plotted for each
species within an area delimited by 17°S and 33°E; inset 1. Winter rainfall region in the cooler southwest.
onto which range within Africa was superimposed for 2. Bimodal spring/autumn rainfall region along the
species with more than a single geographical record.
cooler south coast.
8 SURICATA 6 (2020)

3. Arid late summer rainfall region on the cooler Bioregions and ecoregions
northwest coast and hotter west to south centre.
4. Mid-summer rainfall region in the cooler highlands Coincidence of dung beetles with defined ecological re-
and warmer lowlands to the more mesic northeast. gions assists in assessment of conservation status. It was
decided to use two different classification systems: one for
Numbers of dung beetle species in South Africa show a de- vegetation types in South Africa (Mucina & Rutherford
cline from the northeast to the southwest (Davis 2002). 2006) and one for the remainder of southern Africa that
Previously, a combined total of 386 species has been re- comprised part of a global classification of ecoregions (Ol-
corded to the northeast in the mid-summer rainfall region. son et al. 2001).
Although northeast sub-regions show smaller totals, there
are still around 150 species in the highlands or greater than The vegetation of South Africa has been classified at three
200 in the lowland sub-regions. This number is reduced to different spatial scales using regression trees (Mucina &
around 125 species in the central arid late summer rainfall Rutherford 2006). From larger to smaller scales, these
region (Davis et al. 2016). In the bimodal rainfall region comprise eight biomes (Figure 3), 35 bioregions and 439
of southern South Africa and the winter rainfall region of vegetation units. Principal associations of dung beetles
the southwest, respectively, only 86 and 62 species were with vegetation areas have been described according to the
previously recorded (Davis 2002). Numbers cited by Davis size of their ranges. Those showing larger ranges are most-
(2002) are updated in the Synthesis section. ly described at biome and bioregion scales. Those showing

Table 1. Numbers of species assessed for the IUCN Red List; numbers of endemic species and numbers also recorded outside of the
borders of South Africa, Botswana and Namibia; numbers of species assessed for each IUCN threat category (see IUCN 2001)

CATEGORIES Number of species accounts (% of total species accounts) per tribe

Ateuchini Canthon. Coprini* Gymnopl. Oniticell. Onitini Onthoph. Scarab. Sisyphini

IUCN Red List 4 15 9 7 6 13 25 23 8

Endemic status
South Africa (RSA)** 16 (55) 64 (78) 34 (40) 2 (10) 2 (8) 10 (22) 33 (24) 28 (30) 10 (38)
Namibia 0 12 (15) 3 (4) 1 (5) 0 2 (4) 1 (1) 8 (9) 0
Botswana 0 0 0 0 0 0 0 1 (1) 0
RSA/Botswana 0 0 2 (2) 0 0 0 1 (1) 4 (4) 0
RSA/Namibia 0 2 (2) 0 0 0 7 (16) 3 (2) 10 (11) 1 (4)
Namibia/Botswana 0 0 0 0 0 0 1 (1) 1 (1) 0
RSA/Namibia/Botswana 1 (3) 1 (1) 6 (7) 1 (5) 0 2 (4) 2 (1) 9 (10) 0
Total endemics 17 (58) 79 (96) 45 (53) 4 (20) 2 (8) 21 (46) 41 (30) 61 (66) 11 (42)
Not endemic 12 (42) 3 (4) 40 (47) 16 (80) 24 (92) 24 (54) 95 (70) 31 (34) 15 (58)

Conservation status
Data Deficient (DD) 16 (55) 41 (50) 24 (28) 4 (20) 5 (19) 15 (33) 26 (19) 26 (28) 5 (19)
Least Concern (LC) 12 (41) 25 (31) 56 (66) 16 (80) 21 (81) 28 (62) 110 (81) 62 (67) 20 (77)
Near Threatened (NT) 0 0 3 (4) 0 0 1 (2) 0 1 (1) 1 (4)
Vulnerable (VU) 0 15 (18) 0 0 0 1 (1) 0 2 (2) 0
Endangered (EN) 1 (3) 1 (1) 2 (2) 0 0 0 0 1 (1) 0
Critically Endangered (CR) 0 0 0 0 0 0 0 0 0

DD but ?threatened
DD/NT 0 1 0 0 0 0 1 1 0
DD/VU 5 9 8 0 0 1 0 4 1
DD/EN 2 4 3 0 0 1 0 2 0
DD/CR 2 1 0 0 0 1 0 0 0

Total N species accounts 29 82 85 20 26 45 136 92 26

*Catharsius bradshawi not included

**Includes Lesotho and Eswatini (formerly Swaziland): nine in RSA/Lesotho; five in RSA/Eswatini; one canthonine in Eswatini only.
SURICATA 6 (2020) 9

Figure 3. Map showing the vegetation classification for South Africa (Mucina & Rutherford 2006) at biome scale.

Figure 4. Map showing (1) the ecoregions of Namibia and Botswana plus parts of Angola, Zambia and Zimbabwe (Olson et al.
2001: Arid: AT1309–1322; Savanna: AT0702–0726; Lacustrine: AT0902–0908) and (2) how they abut onto the vegetation
classification for South Africa at biome scale (Mucina & Rutherford 2006).
10 SURICATA 6 (2020)

smaller ranges are also described at vegetation unit scale of Zimbabwe and the extreme south of Mozambique (Fig-
within the context of both bioregion and biome scales. ure 4). Distributions are described for 11 principal ecore-
Statements on the proportional modification of biomes, gions defined across that area. Except in the case of the
bioregions and/or vegetation units plus its potential im- north Botswana alluvial soil specialist, Escarabaeus remii,
pact on conservation status have been provided for many there are no records from three sizeable ecoregions of al-
species whose ranges partly or wholly coincide with highly luvial soils comprising Etosha Pan (Namibia), Okavango
modified and/or fragmented natural vegetation cover. Swamps and Makgadikgadi Pan (N Botswana). Further-
more, no data are provided for the 14 ecoregions recorded
Although the ecoregion classification of Olson et al. across South Africa as descriptions of distribution across
(2001) is global in extent, it has been used to describe the vegetation regions of Mucina & Rutherford (2006)
dung beetle distribution only in Botswana, Namibia, most are considered more comprehensive.

E. ECOLOGICAL ASSOCIATIONS
Dung beetles show a range of ecological associations from Associations with textural categories of soil type have been
relatively generalist to relatively specialist. Therefore, iden- provided for each species where available.
tification of preferred habitats (soil and vegetation types)
or preferred food (primarily carrion, fungi, millipede
body contents or dung type) is important for conservation Vegetation type
purposes. Identification of factors that contribute to the
Dung beetles are strongly influenced by structural proper-
observed associations are also important.
ties of vegetation at both macro- and microhabitat scales.
Macrohabitat associations with different vegetation types
Soil type have been defined primarily in terms of (1) increasing
height of the vertical profile from grassland to forest and
As fossorial insects, dung beetles are strongly influenced by (2) increasing cover density of the canopy in woody vege-
the textural (Davis 1996b) and hydrological properties of tation. In terms of the surface microhabitats occupied by
soil. Associations with soil type are mostly defined using dung beetles, this represents a gradient from exposure to
four categories from the soil texture, classification system radiant heat to potential protection by increasing shade.
(sand, sandy loam, sandy clay loam, clay) of the United Such differences in the surface microhabitat are known to
States Department of Agriculture (USDA). In rank order, strongly influence species representation and assemblage
this classification represents contrasting grain size profiles structure of dung beetles (Davis et al. 2013). Vegetation
from coarse to increasingly finer-grained, which vary ac- associations are, therefore, described in terms of (1) grass-
cording to proportions of sand, silt and clay in each soil land and scrub, which provide little shade, (2) open shrub-
type. Ease and depth of tunnelling varies with soil type land or woodland, which may provide partial shade, and
(Hanski & Cambefort 1991a). It increases across the gra- (3) dense woodland, thickets or forest, which provide the
dient from finer to coarser-grained soils and also with in- greatest shade.
creasing soil moisture content following rainfall. Species
richness and biomass of dung beetle assemblages tends to Vegetation type is also responsible for creating or enhanc-
be greater on sandy soils than on finer-grained soils (Davis ing the surface microhabitats favoured by species. For in-
1996b). stance, a number of small-bodied specialists is associated
with the carpet of leaf litter found in forests. Furthermore,
Influences of grain size profiles on soil hydrological prop- dung beetle activity is favoured by greater density of sur-
erties include faster rate of drainage on coarser sandy soils face cover in grassland (Jankielsohn et al. 2001), which
and higher water retention capacity in finer-grained soils. reduces surface evaporation. However, extreme surface
Such properties are modified by dung beetle tunnelling cover density in fertilised Kikuyu pasture may inhibit
activity (Brown et al. 2010). Variation in soil hydrolog- tunnelling activity (Davis et al. 1999). Also, the impor-
ical properties according to grain size may be associated tance of shade is illustrated by observations on the effect
with differences in survivorship of immatures of different of weather conditions. On cloudy days, the shade special-
species on different soil types (Fincher 1973). Differences ist (Sisyphus nanniscus cited as S. costatus) has been record-
in water content between drier and moister soils may also ed in abundance in tall grass 50 m from dense woodland
influence nest architecture (Rougon & Rougon 1982). thicket (Doube 1983). Associations with categories of
SURICATA 6 (2020) 11

vegetation type and particular microhabitats (if applica- This results in dung types that vary in volume, physical
ble) have been provided for each species where available. consistency and spectra of released volatiles (Dormont et
al. 2010) to which dung beetles are attracted by olfaction
(Shibuya & Inouchi 1982; Inouchi & Shibuya 1986).
Food type Therefore, proportional associations with dung or other
food types have been provided where available. Howev-
Although association with mammalian dung dominates in er, some citations merely list mammals from whose dung
the subfamily Scarabaeinae, various other specialist associ- collections were made. These citations may categorise
ations occur, including those with carrion, fungi and mil- mammals according to diet and digestive process: (1)
lipede body contents. In the dominant dung-associated carnivores or omnivores producing small strong smelling
component, many species show a bias to particular dung droppings; (2) ruminant herbivores producing pellets or
types (Davis 1994; Davis et al. 2014) that differ accord- large fine-fibred pads; (3) monogastric herbivores produc-
ing to mammalian body size, diet and digestive processes. ing large coarse-fibred droppings.

F. TEMPORAL ASSOCIATIONS
Temporal associations may have little relevance for cur- drier weather, the number of active species in grassland on
rent conservation purposes although drivers of such pat- sandy clay loam declined from over 40 to around 20 in
terns may be important, particularly in a future threat- the Gauteng bushveld (Davis 2002).
ened by climate change. Temporal activity is described at
three different scales, (1) diel (variation over 24 hours of
each day), (2) day-to-day variation according to weather Seasonal activity
conditions and (3) seasonal. The joint effects of activity at
these temporal scales may contribute to long-term, year- Although seasonal peaks in activity are described for only
to-year changes in diversity that should be monitored un- some species where quantitative data are available, a quali-
der a scenario of climate change. tative assessment of annual activity is provided for all spe-
cies by shading along a bar graduated in months. Such
data are derived from months of collection on locality
Diel periodicity labels of reference material and from raw data of quan-
titative studies. Most of these assessments are probably
If known, diel periodicity is classified as either diurnal representative of the annual pattern of activity. However,
or active under darkness. Additional notes are added for it should be noted that there may be some bias as most
a few species for which more detailed, quantitative data collecting trips have been mounted in the warmer parts
on daily activity peaks is available. The diel patterns are of rainy seasons, which are much more favourable for ac-
driven by light intensity and temperature (Caveney et al. tivity than the unfavourable cool and/or dry seasons. It
1989). should also be noted that chance records of single speci-
mens under unfavourable, out-of-season conditions have
the same weight as many specimens recorded during sea-
Short-term weather responses sonal peaks in activity. The recorded extent and patterns
of seasonal activity may even vary according to different
Dung beetle activity varies from day-to-day depending climatic conditions across a species range.
on incidence of favourable or unfavourable weather con-
ditions for activity (Davis 1995, 2002). During seasonal Overall, seasonality of dung beetle activity varies accord-
peaks in activity, variation is primarily mediated by inci- ing to the different rainfall patterns in the four main
dence of substantial rainfall and incidence of cloud cover, climatic regions of southern Africa (see under Climatic
which affect diel cycles of temperature and light intensi- associations pg. 7). In the summer rainfall region to the
ty. Rainfall also increases the ease with which soil may be northeast, rainfall usually coincides with warmer tempera-
tunnelled. As long as temperatures are sufficiently high, tures leading to a unimodal mid-summer peak in dung
numbers of active species increase after substantial rain- beetle activity (Davis 2002). In the winter rainfall region
fall and decline during periods of dryness (Davis 1995, to the southwest, rainfall usually coincides with cooler
2002). Over a 10-day period in early summer that com- temperatures, leading to bimodal peaks in dung beetle ac-
menced with heavy rainfall and progressed to warmer tivity during the warmer spring and autumn.
12 SURICATA 6 (2020)

Year-to-year activity species and their breeding opportunities. Between-species

differences in breeding success would drive different pat-
Dung beetle activity varies substantially from year-to- terns of diversity in the following year. Under a scenario
year in terms of relative abundance of species (Doube of long-term changes in climate, such responses would
1991). Although the reasons have not been established, have implications for conservation, particularly for
year-to-year differences in the frequency of rainfall species with small ranges showing ecological specialisa-
events would influence activity patterns of different tions.

G. CONSERVATION STATUS AND METHODS

For assessments of conservation status, we use the cate- Red List website (http://www.iucnredlist.org/) (Table 1).
gories recommended in the IUCN Red List, Version 3.1 For each species, the awarded IUCN threat category de-
(IUCN 2001). As trends in population dynamics are pends on how much is known concerning its EOO, AOO
mostly unknown, awarding of categories is largely based and ecological associations. A summary is provided below.
on spatial distribution patterns using the rules stipulat-
ed for range size and number of known locations (IUCN 1. Data Deficient (DD): Assessment for 160 species
2001: see below). However, assessments may also be mod- for which information is extremely limited. How-
ified after a consideration of ecological associations and ever, threat categories are also discussed for 40 of
representation in reserves, if known. these species (NT, VU, EN or CR, see below) if
they show small or highly fragmented ranges that
are clearly not collection artefacts. This would po-
Assessment rationale tentially augment the small numbers of species (30)
currently awarded threat categories (Table 1).
The starting point for each assessment is a measurement of
the Extent of Occurrence (EOO) derived from an outline 2. Least Concern (LC): Assessment for the majority
of available validated distribution data that are, in some of species (350) that show large ranges and high
cases, severely limited. This is usually only an estimate frequency or small ranges with high abundance
of actual occurrence as an Area of Occupancy (AOO) is within reserves. Because Scarabaeinae are primarily
generally unavailable. However, available ecological data coprophagous, the wide availability of dung results
may provide insight into likely influences on spatial fre- in assessments of Least Concern (LC) for many
quency, range restriction or range contraction within the southern African species. Similarly, assessments of
cited EOO. Generalisation versus specialisation to soil LC are also awarded to a number of species with
type may be influential if an association results in a small specialisations on other widely available food types,
range. Restricted occurrence of preferred vegetation types such as carrion, fungi or millipede body contents.
or coincidence with areas of highly modified vegetation
structure are particularly pertinent to the conservation 3. Near Threatened (NT): ‘A taxon is Near Threat-
status of dung beetles. Data on proportional transfor- ened when it has been evaluated against the crite-
mation of vegetation units within bioregions (Mucina & ria, but does not qualify for Critically Endangered,
Rutherford 2006) are sometimes included as a guide to Endangered or Vulnerable now, but is close to
the extent of possible threats to a species. Increasing range qualifying for or is likely to qualify for a threatened
restriction of mammals dropping preferred dung types category in the near future’ (verbatim from IUCN
may also be influential and is often identified by a current 2001); currently awarded to six species.
beetle distribution pattern centred on reserves. Although 4. Vulnerable (VU): Threat category awarded if
population data are usually limited, numbers in collec-
a species shows an EOO < 20 000 km2, AOO
tions or quantitative sampling may provide insights into
< 2 000 km2, fragmented range or records from only
relative temporal frequency or rarity. Based on these data,
6–10 locations; currently awarded to 18 species.
an IUCN threat category is provided.
5. Endangered (EN): Threat category awarded if a spe-
cies shows an EOO < 5 000 km2, AOO < 500 km2,
IUCN categories fragmented range or only five locations; currently
awarded to six species.
As indicated in relevant species accounts, conservation
status of 110 taxa found in South Africa, Botswana and/ 6. Critically Endangered (CR): Threat category
or Namibia has been previously assessed for the IUCN awarded if a species shows an EOO < 100 km2,
SURICATA 6 (2020) 13

AOO < 10 km2, fragmented range or was record- Conservation measures

ed from only one location; not currently awarded
to any species, but the flightless Copris sexdentatus Conservation measures are recommended for all but a
(EN) is currently known from only one location in few species. However, information on which to base as-
critically endangered Eastern Rûens Renosterveld sessments are often limited to some degree, whatever the
having been recently re-recorded at one small re- awarded threat category. In some cases, further work is re-
serve after a hiatus of many decades. quired on taxonomic status or geographical range. How-
7. Extinct (EX): No southern African scarabaeine spe- ever, most frequently, there are recommendations for data
cies is currently regarded as globally extinct. How- on ecological associations which may be absent, restricted
ever, at the time of writing, two flightless fynbos to qualitative observations, or do not represent sufficiently
taxa have not been re-recorded for many decades detailed quantitative support. Without such supporting
(Copris sphaeropterus) or over a century (Aphengoe- data it is difficult to determine if the AOO is likely to be
cus clypeatus). smaller than the EOO. Each section ends with citation of
a few reserves where the species is protected or a statement
VU, EN and CR categories are considered to represent a that current records of the species are not known to coin-
high risk of extinction in the wild. cide with any reserve.

H. POTENTIAL THREATS

Regional habitat transformation east. Also in Namibia, AT1310 and AT0702 are largely
protected by local and national conservancies although
In Namibia and Botswana, transformation of natu- poaching is a problem.
ral ecoregions (Olson et al. 2001; World Wildlife Fund
2017) is mostly fairly limited except for the moister, In South Africa, conversion of natural environment to ur-
more densely populated north and east. Assessments of ban centres or agro-ecosystems varies from limited to fair-
conservation status are not equivalent to those for South ly advanced (Mucina & Rutherford 2006). Assessments
Africa (see below) as they are made in terms of both veg- of conservation status are made in terms of vegetation
etation and natural mammal complement. Ranges of the transformation as the large-bodied members of the nat-
latter have been more highly modified than vegetation. ural mammal fauna are now largely restricted to reserves
Three ecoregions are assessed as relatively stable/intact and game farms. Vegetation of the arid regions from cen-
(World Wildlife Fund 2017), which may equate to least tral South Africa to the West Coast remain relatively lit-
concern (Namib Desert: AT1315; Kalahari Xeric Sa- tle transformed and are considered least concern (0–20%
vanna: AT1309); Zambezian and Mopane Woodlands: transformed) (Nama Karoo, Succulent Karoo, Desert bi-
AT0725). Five are assessed as vulnerable (Nama Karoo: omes; most of the Albany Thicket Biome and Kalahari Sa-
AT1314; Namibian Savanna Woodlands: AT1316; Ka- vanna bioregions; see Mucina & Rutherford’s Figures 3.8
lahari Acacia-Baikiaea Woodlands: AT0709; Southern and 16.5). However, the moister Grassland and Savanna
African Bushveld: AT0717; Zambezian Baikiaea Wood- biomes to the northeast are largely classified as vulnerable
lands: AT0726) (World Wildlife Fund 2017). Woodland (20–40% transformed) to endangered (60–85% trans-
clearance is cited as a problem in AT0726, overgrazing formed). This excludes the savanna bordering the Lim-
by domestic livestock as a problem in AT0709, AT1314, popo Valley and Kruger National Park, and high altitude
AT1316, with overhunting a problem in AT1316 and all areas, particularly the Drakensberg Grassland bioregion,
indigenous fauna hunted out in the Namibian part of which remain least concern (see Figure 3.10. in Mucina &
AT1314. Three ecoregions are assessed as critical/endan- Rutherford 2006). Similarly, much of the moister east and
gered (World Wildlife Fund 2017), which may equate to south coastal regions (Indian Ocean Coastal Belt, Fyn-
endangered (Succulent Karoo: AT1322; Kaokoveld Des- bos biomes) are highly modified with assessments varying
ert: AT1310; Angolan Mopane Woodlands: AT0702). from vulnerable to endangered or even critically endan-
However, this does not apply to the Namibian parts of gered (85–100% transformed) in large, lower-altitude ar-
these ecoregions. In Namibia, AT1322 is protected in the eas of the Western Cape that coincide, particularly with
Sperrgebiet National Park where the vegetation is most- Renosterveld. However, again, many mountainous areas
ly intact although there is disturbance by mining on the and some sandy south coastal areas in the Fynbos Biome
coast with fencing preventing mammal migration in the remain assessed as least concern.
14 SURICATA 6 (2020)

The relative influence of habitat transformation on dung of Renosterveld above), development of grassland
beetle species is dependent on the size of their geograph- pastures and urban expansion is a growing threat
ical range and the breadth of their ecological associations to many species in the Eastern and especially the
(generalist or specialist). Species with ranges that coincide Western Cape.
with the most modified coastal or northeast regions of
2. Indigenous forest patches along the coastline and
South Africa are potentially those most at risk. The de-
mountain escarpment in the south and east are
gree of risk would increase for species with small overall
known to harbour various often rare or uncom-
range and specialist ecological associations, particularly
mon endemic genera and species, most of which
specialisation to threatened vegetation and food types.
are assessed as threatened. These forest patches are
Threatened vegetation may be associated with particular
naturally restricted in extent. Thus, threats from
soil types, such as sands concentrated around much of the
unsanctioned and uncontrolled clearance, area
coastline or in outliers within savanna, although this does
reduction, degradation and exploitation would be
not apply to the sands of the Kalahari Basin, Namib Des-
detrimental.
ert and outliers in the Karoo.
3. A number of savanna species are recorded as show-
ing a bias to thickets or open woodland vegetation
Local habitat modification (Davis 1996b). Therefore, proportional estimates
of transformation and clearance in woody vegeta-
Potential threats stem from modification to the natural tion units provide some insight into potential local
environment. These may or may not impact on particular threats.
ecological associations shown by each dung beetle species. 4. Modification of Highveld grass cover by pasture
improvement has been little studied although dif-
ferences have been recorded for some species be-
Soil modification tween dense Kikuyu pasture and less dense natural
Themeda-dominated grassland (see species accounts
Widespread only with regard to ploughing. with data cited from ‘At Foggy Valley’).
1. Ploughing: Conversion of natural habitat to arable 5. The impact of conversion of Highveld Grassland
lands is perhaps one of the most extreme forms of to tree plantation by commercial forestry has been
transformation. Although only 10.3% of the land little measured.
surface of South Africa was transformed as arable
lands by 2014, it is the geographical concentration 6. Observations suggest that reduction in vegetation
of such farming that is important as a conserva- cover by overgrazing also influences diversity of
tion factor, particularly in the Renosterveld of the dung beetle assemblages in grassland regions. Al-
southwest Cape, to which two, now extremely rare though it has been little measured, one study has
flightless species, are apparently endemic. shown a reduction in abundance of larger-bodied
species from undisturbed to overgrazed grassland
2. Mining: In general, mining is usually spatially re- (Jankielsohn et al. 2001).
stricted in its effects although open cast mining is,
potentially, more of a threat. Measurements near
Phalaborwa indicated limited influences from min-
Food availability and toxicity
ing (Davis et al. 2014).
Availability of dung types of indigenous mammals is wide-
ly modified in southern Africa. Over wide areas, these
Vegetation transformation dung types have been replaced by those of domestic live-
or habitat fragmentation stock, which may be potentially toxic due to widespread
use of pesticide treatments.
Widely modified for both urban development and farm-
land in the southwest and northeast of South Africa as 1. Range reduction of large indigenous mammals
well as along much of the coastline. through hunting has lead to loss in availability of
particular dung types from most of South Africa
1. It has been shown that many winter rainfall en- and extensive areas of Namibia and Botswana. This
demics are associated primarily with the natural has been most influential on those species associat-
shrublands of the fynbos region (Davis 1993). Thus, ed, especially, with antelope pellets on the Highveld
loss of shruband to arable farming (see ploughing and coarse-fibred dung of monogastric herbivores
SURICATA 6 (2020) 15

over most of the subcontinent. Distributions of a Range contraction

number of species are now concentrated in areas
containing game reserves with the implication of In summary, range contraction by dung beetles may re-
past contraction in their ranges. sult from specialisation to particular dung types dropped
2. Several types of synthetic pesticide treatments by mammals that have become range restricted through
are used to control ecto- or endoparasites on the hunting, the so-called game reserve effect. It may also oc-
domestic livestock that have widely replaced the cur through clearance of woody vegetation that removes
large indigenous mammal fauna. These are ad- suitable conditions for specialists to shady or partially
ministered as pour-ons (pyrethroids) or also as shady conditions. Such effects of habitat degradation may
ingestables or injectables (avermectins, milbe- be exacerbated by specialisation to particular climatic or
mycins). All leave varying amounts of residues soil type conditions. Ranges of the most threatened spe-
in dung that are variously toxic to dung beetles, cies mostly coincide with areas that have undergone exten-
both globally and in southern Africa (Davis et al. sive to extreme modification due to urban development or
2004; Wall & Beynon 2012; Jacobs & Scholtz farming practices, particularly the southwest Cape, south
2015). Further monitoring of the effects would be and east coastline, and northeast South Africa (Highveld
advisable. Grassland, woody savanna, woody coastal vegetation).

I. USE AS INDICATORS
Scarabaeine dung beetles are used, globally, as an indicator local habitat and microhabitat. These studies have in-
group owing (1) to their sensitivity to habitat heterogene- vestigated (1) classification of geographical areas (Davis
ity and transformation, and (2) ease with which they may et al. 2016), (2) habitat and food associations (Davis
be sampled quantitatively using dung-baited pitfall traps 1996b; Tshikae et al. 2013b), (3) effects of vegetation
(Halffter & Favila 1993; McGeoch et al. 2002; Spector type fragmentation (Davis 1993) or restoration includ-
2006). Therefore, the accuracy of nomenclature and cor- ing influences of microhabitat (Davis et al. 2013), and
rect identification are important. (4) effects of food availability or quality as regards range
contraction in indigenous monogastric mammals (Davis
In southern Africa, dung beetles have been used to assess 1997) or treatment of domestic livestock with pesticides
patterns of distribution and association at various spatial leading to harmful residues in their dung (Kryger et al.
scales (Davis et al. 2004) from biogeographical down to 2006, 2007).

J. FORMAT OF SPECIES ACCOUNTS

The book comprises sections on the subfamily Scarabaein- 2. Global conservation status: IUCN category fol-
ae, the nine tribes found in Africa, and the 86 genera found lowed by citation of South Africa (RSA), Botswana
in South Africa, Botswana and Namibia. It also comprises and/or Namibia, if endemic within the borders of
542 species accounts for dung beetles found in these three any one, two or all three countries.
countries. Each species account is arranged according to a
3. Seasonal activity bar: Qualitative assessment based
standard format of 14 to 15 sections. Depending on geo-
on months in which collections or samples were taken.
graphical distribution, one or the other of sections 12 and
13 may be deleted or both may be included. The content 4. Map: Distribution in southern Africa according
under the headings varies according to the history of the to presence in 1/16th degree squares across a map
species and the amount of available information. panel extending from 17°S to 33°E (South Africa,
Botswana, Namibia, most of Zimbabwe, extreme
south of Mozambique); usually with an inset show-
Species accounts ing known or estimated range in Africa (Figure 2).
1. Species name(s): Cited in large font italics fol- 5. Type localities: One or more type localities quot-
lowed by synonyms (if any) in smaller font italics. ed from original published descriptions of valid
16 SURICATA 6 (2020)

species and their synonyms; current changes in increasing range restriction of mammals dropping
locality names and/or qualifying notes are added preferred dung types or association with threatened
where necessary. vegetation types are particularly pertinent to the
6. Taxonomy: Species status is usually cited as ‘Ac- conservation status of dung beetles.
cepted species’ although additional notes are pro- 15. Conservation measures: Either ‘None recom-
vided if status is disputed or if there is appreciable mended’ or a discussion of recommended measures
intraspecific variation. derived from the Assessment rationale (pg. 12)
7. Distribution: A general description is provided for section. Coincidence of each species with protected
the species range as many taxa show distributions areas and a listing of a few of the more important
extending beyond the confines of the map (see reserves, if any.
Map).
8. Locality data: Ranges and mean values ± SD for Summary tables
altitude, annual rainfall and annual temperature
are provided for each species. These data are de- Three summary tables (Appendices 1, 2 and 3) have been
rived from average values within 5 × 5 km polygons developed from information in the species accounts. Sum-
extracted from GIS base maps for all of southern mary Table 1 (Appendix 1) is for species found in South
Africa south of 15°S. Africa. This includes data on proportional distribution
9. Habitat: Soil and vegetation associations in south- across the biomes of Mucina and Rutherford (2006) based
ern Africa are described according to available data, on occurrence in 1/16th of a degree squares. It also includes
qualitative or quantitative, published or unpub- IUCN categories and summarises putative ecological bias
lished. in each species if known. Summary Table 2 (Appendix 2) is
for species recorded in Namibia and Botswana. It includes
10. Food types: Food type associations in southern Af-
data on proportional distribution across the ecoregions of
rica are described according to available data, qual-
Olson et al. (2001) based on occurrence in 1/16th of a de-
itative or quantitative, published or unpublished.
gree squares. It also lists IUCN categories for each species
11. Temporal activity: Diel and seasonal periodicity in that is either restricted to those two countries or is also
southern Africa are described according to available found in South Africa. The number of species found in
data, qualitative or quantitative, published or un- each biome or ecoregion has been added at the base of each
published. If available, data describing short-term summary table. These numbers are dependent on climate
responses to weather conditions are also included. and area of each region. There are some differences to the
12. Ecoregions: For species with distributions partly numbers defined for climatic regions reported above (see
or entirely within Botswana and/or Namibia, cita- last paragraph under Climatic associations). Appendix 3
tions are made for coincidence with the ecoregions summarises climatic, altitudinal, distributional and range
of Olson et al. (2001). These citations include the size data for each species included in the species accounts.
ecoregions of Zimbabwe and the extreme south of
Mozambique for species also occurring in those The numbers of species recorded for biomes do not clear-
countries. ly show the decline in species numbers from NE to SW.
The low numbers shown for winter (62) and bimodal (86)
13. Bioregions: For species occurring partly or entirely rainfall regions (Davis 2002) are hidden as the Fynbos Bi-
within South Africa, citations are made for coinci- ome combines the area of both regions although overall
dence with the floral areas of Mucina & Ruther- numbers are similar (130). Numbers of species recorded
ford (2006). For species with larger ranges citations at localities may be a better criterion on which to base NE
are at biome and bioregion scale. For species with to SW declines. For instance, over one year, species as-
smaller ranges, citations are often also at vegetation semblages at winter rainfall localities in the Western Cape
unit scale. (Cape Peninsula shrubland: 13; SW coast shrubland:
14. Assessment rationale: An EOO (Extent of Occur- 23–33), were much lower than those in the mid-summer
rence) is provided for most species and a general rainfall region on the Gauteng bushveld (sandy clay loam,
discussion is made to support the allotted IUCN open woodland: 66; deep sand, open woodland: 76). In
threat category. This may include notes on the sizes the Western Cape, highest temperatures coincide with the
of the EOO and AOO (Area of Occupation) as well dry season so that activity peaks are in spring and autumn
as generalisation versus specialisation to soil type, whereas activity peaks and highest temperatures both co-
vegetation structure, and/or food type. Restrict- incide with the rainy season on the Gauteng bushveld
ed range and occurrence of preferred soil types, (Davis 2002, see pg. 11 under Seasonal activity).
18 SURICATA 6 (2020)

Subfamily
SCARABAEINAE
Latreille, 1802

The Scarabaeinae is one of 16 extant subfamilies consti- Deltochilini, Coprini) and the added suffix of sensu novo
tuting the diverse beetle family Scarabaeidae. In terms of [new sense]. For those polyphyletic tribes with radically
numbers of species, the Scarabaeinae are primarily diver- reduced membership, 100 genera were given the status of
sified to feed and breed using vertebrate dung although incertae sedis (uncertain placement) and the geographical
there are various African groups with different ecological range of two tribes was reduced to the Americas (Ateuchi-
specialisations, including carrion or fungus-feeding, or ni, Canthonini/Deltochilini).
poorly known habits, in forest leaf litter.
Revision of tribal classification is necessarily incomplete due
Until recently, the subfamily has been divided into 12 to different sets of genera included in each molecular phy-
tribes based on morphological characters (Smith 2006). logenetic study and frequent differences between evolution-
However, both morphological and molecular phylogenies ary relationships claimed for the same genera. Out of ~261
show that only nine are monophyletic (three not repre- valid genera and ~6 293 valid species recorded worldwide,
sented in Africa) whereas the remaining three are exten- the phylogenies analyse relationships between only 38.3–
sively polyphyletic (Philips et al. 2004; Monaghan et al. 52.5% of the genera and a very much smaller proportion
2007). Although topology differs radically between differ- of the species. Therefore, further analyses are required to
ent global molecular phylogenies (Monaghan et al. 2007; support a full revision of tribal classification. Under these
Gunter et al. 2016; Tarasov & Dimitrov 2016), the most circumstances, we have retained the old 12 tribe classifi-
recent study was used to propose a partial revision of trib- cation system (see below) to arrange the species accounts
al classification. This proposal retained eight of the old in this book. However, we have modified the membership
tribal units with unchanged membership (Gymnopleu- of the Coprini and Ateuchini (old sense) according to the
rini, Scarabaeini, Onitini, Onthophagini, Oniticellini, morphological phylogeny of Montreuil (1998). In addition,
Phanaeini, Eucraniini, Eurysternini). Names of the other we have treated the membership of the three polyphyletic
four were also retained but with expanded (Sisyphini) or tribes according to the molecular phylogeny of Tarasov &
radically reduced membership (Ateuchini, Canthonini/ Dimitrov (2016) to better reflect evolutionary relationships.
SURICATA 6 (2020) 19

Twelve tribes (Smith 2006)

Nine tribes recorded in southern Africa
Polyphyletic
Centred on southern continents (Neotropical, Afrotropical, Australasia) with limited northern presence
1. Ateuchini Perty, 1830 (in new sense only Americas; all other genera incertae sedis).
2. Canthonini van Lansberge, 1874a (= Deltochilini Lacordaire, 1856) (in new sense only Americas; all other genera
incertae sedis; but see notes for Sisyphini below).

Afro-Eurasia, Americas and Australia

3. Coprini Leach, 1815 (in new sense only two genera in Afro-Eurasia and Americas; all other genera incertae sedis).

Monophyletic for the most part

Restricted to Afro-Eurasia (Afrotropical, Palaearctic, Oriental)
4. Gymnopleurini Lacordaire, 1856.
5. Onitini Castelnau, 1840.
6. Scarabaeini Latreille, 1802.

Centred in Afro-Eurasia, also variously in Americas and Australia

7. Oniticellini Kolbe, 1905 (also a few species in the Americas).
8. Onthophagini Burmeister, 1846 (Onthophagus species also widespread in the Americas and Australia).
9. Sisyphini Mulsant, 1842 (two species in central America; in new sense with addition of genus Epirinus Dejean,
1833 (formerly Canthonini), which is now removed to the revalidated tribe, Epirinini van Lansberge, 1874a (Dan-
iel et al. 2020)).

Three tribes restricted to the Americas

Monophyletic for the most part
10. Phanaeini Hope, 1838 (Nearctic, Neotropical).
11. Eucraniini Burmeister, 1873 (only Neotropical).
12. Eurysternini Vulcano, Martinez & Pereira, 1961 (only Neotropical).
20 SURICATA 6
X (2020)

TRIBE ATEUCHINI
Perty, 1830

Subtribe Ateuchina Perty, 1830: As Ateuchidae; Type genus: Ateuchus Weber,

1801.
Synonyms:
= Choerididae Harold, 1867a; Type genus: Choeridium LePeletier & Serville, 1828 [= Ateuchus Weber, 1801, senior
name].
= Demarziellini Balthasar, 1961a; Type genus: Demarziella Balthasar, 1961a [incertae sedis].

Subtribe Scatimina Vaz de Mello, 2008: Type genus: Scatimus Erichson, 1847a.

Smith (2006) and Bouchard et al. (2011) provide evidence for Ateuchini as the correct tribal name on the
basis of precedence. Unlike Bouchard et al. (2011), we do not list the recent tribal names of Dichotomiini
Perreira, 1954, and older Pinotinae Kolbe, 1905, as synonyms of the Ateuchini since their type genus (Di-
chotomius Hope, 1838 = Pinotus Erichson, 1847b), and other Copris-like genera, have been transferred to
the tribe, Coprini, as supported by the morphological phylogeny of Montreuil (1998) (see tribal account
for Coprini). However, nomenclature may change further following future revision (see below).

As defined by Montreuil (1998), the non-Copris-like membership of the tribe, Ateuchini, is represented
globally within areas of tropical and warm temperate climate. In the old sense, 37 genera may be assigned to
the tribe on morphological criteria (13 with morphological phylogenetic support). These occur in three bio-
geographical centres: Afrotropical/Oriental (7); Americas (mainly Neotropical: 29); Australia/New Guinea
(1). However, molecular phylogenies show that this tribal membership is polyphyletic, comprising at least
seven distantly related lineages for just 18.9% of the generic membership (Monaghan et al. 2007) or six
lineages for 32.4% of the membership (Tarasov & Dimitrov 2016). As a result, Tarasov & Dimitrov (2016)
redefine the Ateuchini as comprising three lineages restricted to the Americas. All other ateuchine genera are
given the status of incertae sedis, including all those in Africa. Under these circumstances, we use the old clas-
sification system for the Ateuchini as defined by Montreuil (1998), pending a full revision of membership.

In the old sense, many ateuchine genera are forest endemics, including Paraphytus Harold, 1877a, in tropi-
cal Africa and the Oriental region. Other African genera include some forest or shade endemics plus grass-
land and savanna species. Most of the Afrotropical genera show some form of specialisation, one possibly
kleptocoprid on other dung beetle species (Pedaria Castelnau, 1832), one constructing broods from materi-
al in rotten tree trunks (Paraphytus), three related genera possibly all primarily mycetophagous (Coptorhina
Hope, 1833, Delopleurus Erichson, 1847b, Frankenbergerius Balthasar, 1938).

In the old sense, a total of six genera have been recorded for the Ateuchini in the Afrotropical region. Mo-
lecular (Monaghan et al. 2007) and morphological phylogenies (Montreuil 1998) suggest that these genera
comprise three distinct lineages with those of Pedaria and Paraphytus, distant from that for the other four
basally derived genera (Sarophorus Erichson, 1847b; Delopleurus; Coptorhina; Frankenbergerius). The recent
partial revision of tribal classification (Tarasov & Dimitrov 2016) only partly supports these divisions with
Pedaria in a separate lineage but Paraphytus classified together with the other four ateuchine genera in basal
Scarabaeinae (Figure 5, pg. 699).
SURICATA 6 (2020) 21

Two lineages are represented in Botswana, Namibia and/or South Africa where a total of five genera and
possibly up to 37 species are found.
(1A) Coptorhina Hope, 1833: relatively species-poor genus with a southerly distributional bias including
four species centred on savanna or upland grassland in southern Africa.
(1B) Delopleurus Erichson, 1847b: two or three southern African savanna species in a relatively species-
poor genus that is widespread in the Afrotropical and Oriental regions.
(1C) Frankenbergerius Balthasar, 1938: comprises only seven species found in shrublands and forests
along the entire seaboard of South Africa.
(1D) Sarophorus Erichson, 1847b: relatively species-poor genus with a southerly distributional bias in-
cluding ten species centred on dense shrubland, forest or margins of upland grassland in southern
Africa.
(2) Pedaria Castelnau, 1832: Possibly represented by up to 13 species in savannas and upland grasslands
of southern Africa but species accounts are only provided for five validated taxa of this widespread
and species-rich Afrotropical genus.

ATEUCHINI
22 SURICATA 6 (2020)

Genus Coptorhina Hope, 1833

Type species and designation: Coptorhina africana Hope, 1833, by subsequent designation (Janssens 1939a).
Synonyms: None.
Last review: Entire genus reviewed by Frolov et al. (2008).

The long-established genus, Coptorhina Hope, 1833, is currently reduced to seven valid species after the revision of Frolov
et al. (2008) described one new species, but identified nine previous names as synonyms resulting from high intraspecific,
morphological variability with body size. The genus is restricted to the Afrotropical region where it is represented from
central and E equatorial regions southwards to southern Africa.

All species are macropterous comprising specialised mycetophages producing nests with a single brood pear of basidiomy-
cete fragments lined with a clay shell (observed for C. klugi Hope, 1833; C. auspicata Péringuey, 1901).

Four species are found S of 15°S. Two show an E bias, one from E tropical savanna to Highveld Grassland and one restrict-
ed to drier Highveld Grassland. Two show an E savanna bias with additional scattered records in the drier centre and W
of southern Africa. One of these is also distributed widely in the S and central African tropics.

Because of their widespread distributions and widely available food type, most species are assessed as Least Concern (LC)
although one remains poorly known and Data Deficient (DD).

ATEUCHINI
SURICATA 6 (2020) 23

Coptorhina auspicata
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Transvaal (Rustenburg), Southern Rhodesia (Salisbury, En-

keldoorn), Ovampoland (Humbe)’ [South Africa, Zimbabwe, Namib-
ia], lectotype: ‘Rhodesia. Enkeldorn. Darling.’ [Chivhu in Zimbab­we].
Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc),
characters vary in prominence with body size.
Distribution: Frequently recorded in moist upland woody savanna with
a widespread scatter of records elsewhere in the summer rainfall region
of southern Africa; Zambia, Zimbabwe, Namibia, Botswana, South
Africa.
Locality data (mean ± SD, range): Altitude: 1 139 ± 276, 165–1657 m;
annual rainfall: 610 ± 138, 329–948 mm; annual temperature (max. +
min. /2): 19.6 ± 1.6, 15.2–22.8°C (N=56).
Habitat: No quantitative assessment; limited DBRU collection records on
sand (4), sandy loam (2) in grassland/pasture (3), shrub/woodland (3).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (2), mushrooms (3), alighted on the soil surface (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to May); probably active under cooler conditions after rainfall
owing to observed probable primary association with fungi; recorded
in late afternoon (18:00) near Maun, Botswana.
Ecoregions Namibia, Botswana, Zimbabwe: Southern African Bush-
veld (AT0717), Southern Miombo Woodlands (AT0719), Zambezian
Baikiaea Woodlands (AT0726), Xeric Kalahari Savanna (AT1309).
Bioregions South Africa: Primarily in Central Bushveld (SVcb) (Savanna
Biome) with a few records in four other bioregions in the Savanna,
Grassland and Indian Ocean Coastal Belt biomes.
Assessment rationale: EOO = 898 960 km2; widespread EOO, but fre-
quency of records apparently biased to moist savanna, which may be
associated with food specialisation to fungi; poorly known ecologically, 2 mm

but possibly associated with sandy soils; assessment as Least Concern

(LC) probably justified on the basis of widespread EOO and AOO.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on soil and vegetation associations as well
as confirmation of specialist association with basidiomycete fungi; not
widely protected, but recorded from several nature reserves including
D’Nyala and Doorndraai (South Africa).

ATEUCHINI
24 SURICATA 6 (2020)

Coptorhina excavata
Frolov, Akhmetova & Scholtz, 2008

No synonyms
Global: DD
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type locality: ‘Mamathes Basutoland’ [Lesotho].

Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc),
characters vary in prominence with body size.
Distribution: Dry to moist, NW Highveld Grassland, South Africa.
Locality data (mean ± SD, range): Altitude: 1 451 ± 146, 1 293–1 662 m;
annual rainfall: 586 ± 121, 425–733 mm; annual temperature (max. +
min. /2): 16.4 ± 1.2, 14.7–18.3°C (N=8).
Habitat: No quantitative assessment; one DBRU collection record on
sandy loam in grassland.
Food types: No quantitative assessment; one DBRU collection record
from a ‘toadstool’ (basidiomycete fungus).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Feb.); probably active under cooler conditions after rainfall
owing to probable primary association with fungi.
Bioregions South Africa: Dry Highveld Grassland (Gh), W Mesic High-
veld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 115 840 km2; ecologically poorly known,
but a single observation suggests a specialist mycetophage like other
Coptorhina species; relatively low number of records probably related
to its specialist behaviour; probably deserves a Least Concern (LC)
assessment on basis of relatively large range, but currently assessed as
Data Deficient (DD) since possible effects of widespread habitat trans-
formation are unknown (Gh: 4–63%; Gm: 6–69%).
Conservation measures: To assess conservation status, quantitative data
are required on ecological associations and effects, if any, of habitat
transformation; protected in Telperion Nature Reserve (South Africa).

2 mm

ATEUCHINI
SURICATA 6 (2020) 25

Coptorhina klugi
Hope, 1833

= Coptorhina africana Hope, 1833

= Coptorhina obtusicornis Boheman, 1857
= Coptorhina vicina Péringuey, 1901
= Coptorhina optata Péringuey, 1901
= Coptorhina punctata Ferreira, 1954a
Global: LC

J A S O N D J F M A M J

Type localities: C. klugii: ‘Caput Bonae Spei’ [Cape of Good Hope, South
Africa]; C. africana: ‘Sierra Leone’ [?locality error]; C. obtusicornis: ‘jux-
ta fluvium Limpopo’ [near Limpopo River, southern Africa]; C. vicina:
‘Mozambique (Rikatla)’ [Ricatla, S Mozambique]; C. optata: ‘Southern
Rhodesia (Manica)’ [E Zimbabwe]; C. punctata: ‘Transvaal : Pretoria’
[Tshwane/Pretoria, Gauteng, South Africa].
Taxonomy: Accepted species cited as Coptorhina klugii Hope, 1835, by
Frolov et al. (2008); original description of C. klugii Hope, 1833:97
is preceded by that of a synonym, C. africana Hope, 1833:96, whose
cited type locality is far outside the known range of C. klugi and whose
name is now modified to modern style; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size.
Distribution: Primarily moist areas along SE African coastline, edge of E
escarpment, N edge of Highveld, Waterberg and Blouberg with scattered
records elsewhere: South Africa, Lesotho, Zimbabwe, Mozambique.
Locality data (mean ± SD, range): Altitude: 646 ± 584, 0–1 735 m; an-
nual rainfall: 751 ± 142, 315–1 016 mm; annual temperature (max. +
min. /2): 19.5 ± 2.6, 14.4–25.2°C (N=73).
Habitat: No quantitative assessment; limited DBRU collection records
on deep sand (3), sandy clay loam (3) in upland grassland (2), open
woodland (1), lowland woodland (3).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (3) or basidiomycete mushrooms (2); also recorded to
meat and banana bait. 2 mm

Temporal activity: Diurnal flight activity, primarily in the summer rainy

season (Oct. to Apr.); probably active under cooler conditions after
rainfall owing to probable primary association with fungi.
Ecoregions Zimbabwe, Mozambique: Southern Zanzibar-Inhambane
Coastal Forest Mosaic (AT0128), Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Primarily Indian Ocean Coastal Belt Biome; E
edge Sub-Escarpment Grassland (Gs), N and NE edges Mesic High-
veld Grassland (Gm) (Grassland Biome); Waterberg area in Central
Bushveld (SVcb), moist SE Lowveld (SVl) (Savanna Biome); scattered
records in six other bioregions.
Assessment rationale: EOO = 681 450 km2; in the laboratory, brood pear
constructed from macerated mushroom fragments supports speciali-
sation to mycetophagy; but, little support offered by published field
data; habitat associations also unclear; assessed as Least Concern (LC)
on basis of wide distribution, but limited ecological data suggests that
Data Deficient (DD) would be more appropriate.
Conservation measures: Assessment of conservation status would be
much improved by quantitative ecological data from both upland and
coastal situations, including precise food association data and effects, if
any, of soil type bias and habitat transformation; lowland populations
protected in uMkhuze Game Reserve (South Africa).

ATEUCHINI
26 SURICATA 6 (2020)

Coptorhina nitidipennis
Boheman, 1857

= Coptorhina bicolor Ancey, 1883

= Coptorhina seminitida Fairmaire, 1893
= Coptorhina subaenea Janssens, 1939a
= Coptorhina saganicola Müller, 1947
= Coptorhina pygmaea Balthasar, 1963a
Global: LC

J A S O N D J F M A M J

Type localities: C. nitidipennis: ‘prope fluvium Gariep’ [close to Orange

River, South Africa]; C. bicolor: ‘l’Usagara’ [Usagara, NE Tanzania];
C. seminitida: ‘Choa’ [Shewa, Central Ethiopia]; C. subaenea: ‘Congo
belge : Élisabethville : Lubumbashi’ [Democratic Republic of the
Congo (DRC): Lubumbashi]; C. saganicola: Not stated; C. pygmaea:
‘Kamerun: Joko’ [Cameroon: Joko].
Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc),
characters vary in prominence with body size.
Distribution: Widespread, mostly in moist savanna from equatorial to
southern Africa; Nigeria, Cameroon, Central African Republic, south-
ern Sudan, Ethiopia, Democratic Republic of the Congo, Uganda,
Kenya, Tanzania, Malawi, Zambia, Zimbabwe, Mozambique, Angola,
Namibia, Botswana, South Africa.
Locality data (mean ± SD, range): Altitude: 837 ± 562, 0–1 627 m; an-
nual rainfall: 722 ± 318, 181–1 170 mm; annual temperature (max. +
min. /2): 20.3 ± 2.5, 14.7–24.9°C (N=39).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam.
Food types: No quantitative assessment; one DBRU collection record:
caught in flight.
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Apr.); probably active under cooler conditions after rainfall
owing to observed probable primary association with fungi.
2 mm
Ecoregions Namibia, Botswana, Zimbabwe: Centred on Southern Af-
rican Bushveld (AT0717), Southern Miombo Woodlands (AT0719);
marginal in four other ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome); marginal in one other bioregion.
Assessment rationale: EOO = 4 106 325 km2 (gross estimate); assumed
to be a specialist mycetophage like other Coptorhina species; assessed as
Least Concern (LC) on basis of extremely large range although deserves
Data Deficient (DD) status since it is very poorly known ecologically.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
possible effects of habitat transformation; protected in uMkhuze Game
Reserve (South Africa).

ATEUCHINI
SURICATA 6 (2020) 27

Genus Delopleurus Erichson, 1847b

Type species and designation: Delopleurus pullus Boheman, 1857, by subsequent monotypy (Boheman 1857).
Synonyms: None.
Last review: Entire genus reviewed by Frolov (2014) with the addition of one new species (Král
2014).

The long-established genus, Delopleurus Erichson, 1847b, shows an Afro-Eurasian distribution pattern. It currently com-
prises ten valid species centred in the Afrotropical (7), W Oriental (India: 2) and extreme S Palaearctic (Yemen, Socotra: 1)
regions. They are suspected to be mycetophagous, like Coptorhina species, but records from mushrooms, dung and carrion
are too few to determine any specialisation.

Two, perhaps three species have been recorded S of 15°S. Two have been recorded from savanna in two or all three coun-
tries comprising South Africa, Botswana and Namibia. A single record for a third species represents an outlier occurrence
far outside its main range in the tropics.

ATEUCHINI
28 SURICATA 6 (2020)

Delopleurus darrenmanni
Frolov, 2014

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Namibia W. Caprivi Park Nova, 5 km north of Okavango

River 18°09'56’ S, 21°44’31’ E’.
Taxonomy: Accepted species.
Distribution: Known only from a small range on floodplains in central
southern Africa: N Botswana, NE Namibia, SW Zambia.
Locality data (mean ± SD, range): Altitude: 907 ± 21, 889–930 m; an-
nual rainfall: 609 ± 89, 541–710 mm; annual temperature (max. +
min. /2): 22.3 ± 0.3, 22.1–22.6°C (N=3).
Habitat: No quantitative assessment; no data.
Food types: No quantitative assessment; holotype recorded on fungi.
Temporal activity: Probably diurnal flight activity in the summer rainy
season (Nov., Dec.).
Ecoregions Namibia, Botswana: Centred on Zambezian Flooded Grass-
lands (AT0907) (two out of three records) (third record from flood-
plain at Savuti in Zambezian and Mopane Woodlands: AT0725).
Assessment rationale: EOO = 22 265 km2 (presumably underestimated);
probably a specialist mycetophage centred on moist floodplains that
would be favourable for fungal growth; however, ecologically poorly
known; in the absence of quantitative data, assessed as Data Deficient
(DD).
Conservation measures: In order to determine conservation status, a
quantitative survey is required to determine the full EOO, AOO and
ecological associations; this needs to be centred on the Okavango,
Kafue and Cuando (SW Angola) floodplains plus the edges of adjacent
ecoregions; protected in Chobe National Park (Botswana) and West
Caprivi Park (Namibia).

2 mm

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SURICATA 6 (2020) 29

Delopleurus gilleti
Janssens, 1939a

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Togo; Congo belge : Kasai : forêt de Luebo....; Katanga

: Lulua : rivière Kapelekese’ [holotype: Togo; paratypes: Democratic
Republic of the Congo (DRC)].
Taxonomy: Accepted species, but record from N Namibia (Frolov 2014)
falls within the range of D. pullus Boheman, 1857, and represents an
extreme outlier compared to remaining known localities for D. gilleti
(see range map).
Distribution: Widespread intertropical distribution, primarily in W and
SE African savanna either side of the rain forest belt; range may be truly
disjunct or a collection artefact; Côte d’Ivoire, Mali, Togo, Nigeria, S
Democratic Republic of the Congo (DRC), Tanzania, Malawi, Zam-
bia, Mozambique, Namibia.
Locality data (mean ± SD, range): Altitude: 791 ± 401, 329–1 046 m;
annual rainfall: 678 ± 379, 243–933 mm; annual temperature (max. +
min. /2): 21.0 ± 3.4, 17.5–24.3°C (N=3).
Habitat: No quantitative assessment; one DBRU collection record from
gravelly sandy loam in a fallow field (Nigeria), but paratype material
recorded from tropical forest.
Food types: No quantitative assessment; limited DBRU collection records
from mushroom (14 individuals) and cattle dung (1 individual).
Temporal activity: Probably diurnal flight activity in the summer rainy
season (Jan., Feb.).
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 1 773 210 km2 (range without Namibian
record); marginal occurrence in N southern Africa; assessed as Least
Concern (LC) on basis of extremely widespread EOO, and, as a prob-
able specialist mycetophage, apparent occupation of a wide range of
2 mm
vegetation types, including disturbed agricultural lands.
Conservation measures: Assessment of conservation status would be im-
proved by further validation of the Namibian record and quantitative
data on EOO, AOO and ecological associations; not currently known
from protected areas in southern Africa.

ATEUCHINI
30 SURICATA 6 (2020)

Delopleurus pullus
Boheman, 1857

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘prope fluvium Limpopo’ [close to Limpopo River, south-

ern Africa], lectotype: ‘Caffraria’ [inexact, southern Africa].
Taxonomy: Accepted species.
Distribution: Widespread in southern African dry to moist savanna;
disjunct pattern may be a collecting artefact; South Africa, Botswana,
Namibia, Angola, Zimbabwe, Malawi, Mozambique.
Locality data (mean ± SD, range): Altitude: 1 063 ± 352, 1–1 550 m;
annual rainfall: 701 ± 263, 311–1 515 mm; annual temperature (max.
+ min. /2): 19.7 ± 1.7, 17.1–22.9°C (N=23).
Habitat: No quantitative assessment; no data.
Food types: No quantitative assessment; one series of five individuals
recorded from a basidiomycete mushroom at Legonyane, North West
Province, South Africa.
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Feb.).
Ecoregions Namibia, Botswana, Zimbabwe: Southern African Bushveld
(AT0717), Southern Miombo Woodlands (AT0819), Angolan Mo-
pane Woodlands (AT0702), Namibian Savanna Woodlands (AT1316).
Bioregions South Africa: Centred on Central Bushveld (SVcb) with mar-
ginal records in Lowveld (SVl) and Eastern Kalahari Bushveld (SVk)
(Savanna Biome).
Assessment rationale: EOO = 186 855 km2 (underestimated); probable
specialist mycetophagous habits result in limited records and apparent
disjunct distribution; although poorly known ecologically, assessed as
Least Concern (LC) on the basis of wide EOO.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
confirmation of mycetophagous habits and effects of habitat trans-
2 mm
formation, if any; protected in several South African nature reserves:
D’Nyala, Ben Alberts.

ATEUCHINI
SURICATA 6 (2020) 31

Genus Frankenbergerius Balthasar, 1938

Type species and designation: Frankenbergerius mirabilis Balthasar, 1938, by monotypy.
Synonyms: = Pseudocoptorhina Ferreira, 1954a: Type species: Coptorhina armata Boheman, 1857,
by original designation.
Last review: Entire genus reviewed by Frolov & Scholtz (2005).

Frankenbergerius Balthasar, 1938, is endemic to the W, S and E seaboards plus NE edge of the Highveld, South Africa. It
is currently represented by seven species. One widespread, forest taxon occurs as two subspecies centred on the SE coast or
NE escarpment and is assessed as Least Concern (LC). Records for the remaining six species are severely limited resulting
in assessments of Data Deficient (DD).

Of the six poorly known species, three are centred in the summer rainfall region, primarily in forest, occurring variously
on the SE coast and/or NE escarpment. The other three are centred in the Western Cape winter rainfall region on the
SW or W coasts, possibly in shrubland. The two species centred in the highly transformed SW Cape may deserve threat
categories.

Ecological habits are unclear as records of food type are limited in number and have been made, sporadically, on fungi,
fruit and dung.

ATEUCHINI
32 SURICATA 6 (2020)

Frankenbergerius armatus
(Boheman, 1857)

ssp. armatus (Boheman, 1857)

= Coptorhina granulifera Harold, 1871a
= Frankenbergerius mirabilis Balthasar, 1938
ssp. tuberculatus Frolov & Scholtz, 2005
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: ssp armatus as Epirhinus armatus: ‘Caffraria tota’ [inex-

act, southern Africa], lectotype: ‘Caffraria’; C. granulifera: ‘Port Natal’
[Durban, South Africa]; F. mirabilis: ‘Columbien, Chiriguana’ [Co-
lombia, S America; presumably an error, = mislabeled material]; ssp
tuberculatus first of 13 listed type localities: ‘Entabeni Forest Station
[23°00’S, 30°14’E]’ [Limpopo Province, South Africa].
Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size; divided into two subspecies although the
differences are subtle.
Distribution: E South African endemic; SE coast and escarpment (sub-
sp. armatus: red squares); NE escarpment (subsp. tuberculatus: black
squares); citation from Angola is an error.
Locality data (mean ± SD, range): subsp. armatus: Altitude: 376 ± 429,
12–1 406 m; annual rainfall: 778 ± 178, 618–2 522 mm; annual tem-
perature (max. + min. /2): 18.0 ± 1.9, 13.5–20.8°C (N=14); subsp.
tuberculatus: Altitude: 1 216 ± 493, 19–1 635 m; annual rainfall: 839
± 171, 381–1 157 mm; annual temperature (max. + min. /2): 17.3 ±
3.2, 8.7–22.0°C (N=17).
Habitat: No quantitative assessment; no soil type data, however, 37% of
available records (13 out of 35) cited from forest, three from high alti-
tude grassland (remainder not stated); a number of records from litter
of indigenous and plantation forest (Pinus, Quercus, Eucalyptus).
Food types: No quantitative assessment; recorded from mushrooms, fungous
tree bark and trunks, rotten Cussonia fruit; also sampled to banana bait.
Temporal activity: Not known, but presumably shows diurnal flight ac-
2 mm
tivity, primarily in the summer rainy season (Oct. to May).
Bioregions South Africa: Although some records map onto bushveld, forest
and thicket vegetation units, they do not clearly define habitat associations;
in general, N records follow edge of E escarpment and Soutpansberg, rep-
resenting edges of the Savanna (SV) and Grassland Biomes (G) (Low-
er = Lowveld (SVl), Mopane (SVmp), Sub-Escarpment Grassland (Gs);
Upper = Mesic Highveld Grassland (Gm)); S records follow coast and
edge of SE escarpment (Lower = Indian Ocean Coastal Belt Biome (CB),
Albany Thicket Biome (AT); Upper = Sub-Escarpment Grassland (Gs) of
the Grassland Biome; all separated by Sub-Escarpment Savanna (SVs)).
Assessment rationale: EOO = 65 575 km2; widely distributed along SE
coast and E escarpment, but no clear assessment of habitat and food in
the absence of quantitative data; however, shade vegetation dominates
stated habitats and population density may be negatively influenced by
its clearance; only fungi and fruit recorded as food types; currently as-
sessed as Least Concern (LC) owing to large size of range, however AOO
would be much smaller and liable to threat if primarily a shade specialist.
Conservation measures: Although currently cited as Least Concern (LC),
quantitative ecological data are essential to adequately assess conserva-
tion status; protected in various South African forest reserves including
Alexandra (part of Addo Elephant National Park), Dwesa-Cwebe Na-
ture Reserve, Entabeni State Forest.

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SURICATA 6 (2020) 33

Frankenbergerius barratti
(Waterhouse, 1876)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Coptorhina barratti: ‘S. Africa (Transvaal)’ [South Africa].

Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: Primarily follows moist climate along the N and E edges of
the Highveld, South Africa and the W side of Lesotho.
Locality data (mean ± SD, range): Altitude: 1 320 ± 329, 699–1 800 m;
annual rainfall: 768 ± 62, 698–855 mm; annual temperature (max. +
min. /2): 16.3 ± 2.5, 12.0–20.3°C (N=8).
Habitat: No quantitative assessment; single records from both indigenous
forest and natural Highveld Grassland on sandy clay loam.
Food types: No quantitative assessment; one individual sampled to pig
dung, but food association essentially unknown.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity in the summer rainy season (Dec. to Mar.).
Bioregions South Africa: Within or close to the N and E edges of Mesic
Highveld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 16 485 km2; EOO based on spot records
that all lie along the edge of Mesic Highveld Grassland suggesting a
range that is < 20 000 km2; F. barratti is represented by few museum
specimens and is ecologically poorly known; apparent rarity may be
real or a collection artefact; it may deserve an IUCN threat category
of Vulnerable (VU), but is currently assessed as Data Deficient (DD).
Conservation measures: Quantitative ecological data is required to ad-
equately assess conservation status in both natural highland grassland
and forest patches in kloofs along the edge of the N Highveld; F. barratti
could equally be under threat or rarely collected owing to specialised
habits; not currently known from any officially protected area.
2 mm

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34 SURICATA 6 (2020)

Frankenbergerius forcipatus
(Harold, 1880)

= Coptorhina imitativa Péringuey, 1901

= Pseudocoptorhina forcipata Ferreira, 1954a
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Coptorhina forcipata: ‘Cap bon. spei.’ [Cape of Good

Hope, South Africa; presumably inexact]; C. imitativa: ‘Transvaal
(Lydenburg)’ [Mashishing, Mpumalanga, South Africa]; P. forcipata:
‘Colónia do Cabo’ [‘Cape Colony’, possibly Eastern Cape, South Af-
rica].
Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: South African endemic; scattered records along the Eastern
Cape and KwaZulu-Natal escarpment; also Magaliesberg on the N
edge of the Highveld.
Locality data (mean ± SD, range): Altitude: 1 220 ± 721, 443–2 403 m;
annual rainfall: 662 ± 98, 531–819 mm; annual temperature (max. +
min. /2): 15.2 ± 3.2, 9.7–19.2°C (N=6).
Habitat: No quantitative assessment; two of six available records cited as
forest; no soil type data.
Food types: No quantitative assessment; recorded in small numbers in
pitfall traps.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity in the summer rainy season (Dec. to Mar.).
Bioregions South Africa: Possibly shaded patches in Central Bushveld
(Moot Plains Bushveld (SVcb 8), Sub-Escarpment Savanna (Bhisho
Thornveld (SVs 7)) (Savanna Biome); Forest Biome: Southern Mistbelt
Forest (FOz 3), Northern Afromontane Forest (FOz 2) locality cited
as forest, but does not map onto nearest patch within Drakensberg
Grassland Bioregion (Gd) (Grassland Biome).
Assessment rationale: EOO = 21 625 km2; ecologically extremely poorly
known; limited records suggest possible forest or shade associations,
but quantitative support required; large range, but AOO would be
much smaller if it is a habitat specialist; currently assessed as Data De-
2 mm ficient (DD).
Conservation measures: A survey is required to accurately determine the
EOO; quantitative ecological data is also required to assess the AOO
and conservation status; protected in the uKhahlamba Drakensberg
Park (World Heritage Site).

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SURICATA 6 (2020) 35

Frankenbergerius gomesi
(Ferreira, 1954a)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Pseudocoptorhina gomesi: ‘Uitzicht, Zoutpansberg’ [Farm

Uitzicht, W Soutpansberg, Limpopo Province, South Africa].
Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: Records scattered along the W–E trending Soutpansberg,
NE escarpment edge (Limpopo and Mpumalanga provinces), and the
NE coast of KwaZulu-Natal, South Africa.
Locality data (mean ± SD, range): Altitude: 943 ± 488, 19–1 635 m;
annual rainfall: 805 ± 98, 604–985 mm; annual temperature (max. +
min. /2): 18.5 ± 2.1, 14.7–22.6°C (N=13).
Habitat: No quantitative assessment; limited records from riverine bush,
eucalypt plantation, forest litter or pitfall traps in Roodewal Forest,
Soutpansberg.
Food types: No quantitative assessment; limited records from fungus and
fungous [tree] trunk; also sampled to pig dung baits.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity in the summer rainy season (Oct. to Feb.).
Bioregions South Africa: Records straddle the edges of the Grassland (G)
/ Savanna (SV) / Indian Ocean Coastal Belt Biomes (CB); only three
map records close to or coinciding with Northern Mistbelt Forest (FOz
4) or Scarp Forest (FOz 5) patches [does not include samples from
Roodewal Forest (Soutpansberg)]; in N: mapped non-forest records fall
within Legogote Sour Bushveld (SVl 9), Sekhukune Montane Grass-
land (Gm 19), Soutpansberg Mountain Bushveld (SVcb 21) vegetation
units.
Assessment rationale: EOO = 18 695 km2; ecologically poorly known;
limited records suggest a shade specialist that could be either a food 2 mm

type specialist or generalist; mapping unreliable for determining habitat

associations, but, as a shade specialist, AOO would be much smaller
than EOO and population density would be susceptible to clearance of
woody vegetation; possibly deserves an IUCN threat category of Vul-
nerable (VU) on the basis of EOO <20 000 km2 and probable disjunct
AOO in small patches of shade vegetation; however, currently assessed
as Data Deficient (DD).
Conservation measures: Quantitative ecological data is required to ade-
quately assess conservation status; protected in Nelspruit and Lekgala-
meetse nature reserves.

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Frankenbergerius nanus
(Péringuey, 1888)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Coptorhina nana: ‘near Constantia, Cape Colony’ [near

Constantia, Cape Town, Western Cape, South Africa].
Taxonomy: Accepted species, but very close to F. opacus Frolov & Scholtz,
2005; little sexual dimorphism noted.
Distribution: Small coastal range in the SW Western Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 339 ± 374, 20–777 m; an-
nual rainfall: 436 ± 87, 347–555 mm; annual temperature (max. +
min. /2): 15.1 ± 1.8, 12.6–16.6°C (N=4).
Habitat: No quantitative assessment; three known localities from areas
comprising deep sand or sandy loam where the natural vegetation is
shrubland; one specimen sampled on deep sand in strandveld.
Food types: No quantitative assessment; one specimen sampled to cattle
dung, but probably a chance record.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity; recorded during winter and spring in the winter rainy season
(June to Sept.).
Bioregions South Africa: Vegetation units: Atlantis Sand Fynbos (FFd 4),
Cape Flats Dune Strandveld (FS 6), Overberg Dune Strandveld (FS7)
and Peninsula Granite Fynbos (FFg 3) (Fynbos Biome).
Assessment rationale: EOO = 2 995 km2; little ecological data; assessed as
Data Deficient (DD), but may deserve IUCN threat status approach-
ing Endangered (EN) on basis of small range, low number of locations
(3) and extensive transformation by urbanisation and arable farming
in parts of its range (FFd 4: 40%; FS 6: 40%; FS 7: 5%; FFg 3: 56%).
Conservation measures: A survey is required to establish the full EOO
and AOO to adequately assess conservation status; this survey should
2 mm be combined with the collection of quantitative ecological data, partic-
ularly testing response to clearance of natural shrubland; not currently
known to occur in any protected area although recorded at Pella Mis-
sion close to Pella Nature Reserve.

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Frankenbergerius nitidus
Frolov & Scholtz, 2005

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘RSA, Northern Cape Province, Hoekbaai, 31o11’S,

17o47’E’ [South Africa].
Taxonomy: Accepted species.
Distribution: Known only by a single female from the Namaqualand type
locality on the arid W coast of South Africa.
Locality data (average): Altitude: 1 m; annual rainfall: 140 mm; annual
temperature (max. + min. /2): 18.2°C (N=1).
Habitat: No quantitative assessment; known only from a vegetation unit
comprising deep sand and open scrub / shrubland.
Food types: No quantitative assessment; a single, probable chance record
from a white flower (Mesembryanthemaceae); food associations essen-
tially unknown.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity; recorded during spring in the winter rainy season (Aug.).
Bioregions South Africa: Possibly Namaqualand Seashore Vegetation
(Azonal: AZd 2) in which Mesembryanthemum guerichianum is listed as
an important element, unlike adjoining inland Namaqualand Strand-
veld (SKs 7) (Succulent Karoo Biome).
Assessment rationale: EOO/AOO unknown; the range could be small
and restricted to the coastline or wider; also ecologically extremely
poorly known; possible threats impossible to assess although, at pres-
ent, the area around the type locality appears little transformed, except
at Hoekbaai itself; under these circumstances, assessed as Data Defi-
cient (DD).
Conservation measures: A survey along the coastline and inland is
required to determine the EOO and AOO to adequately assess con-
servation status; this survey should be combined with the collection
of quantitative ecological data; not currently known to occur in any
protected area.
2 mm

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Frankenbergerius opacus
Frolov & Scholtz, 2005

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Stellenbosch’ [Western Cape, South Africa].

Taxonomy: Accepted species, but very close to F. nanus (Péringuey, 1888);
little sexual dimorphism noted.
Distribution: Known only by seven individuals, all but one from the type
locality in the SW Western Cape.
Locality data (average): Altitude: 228 m; annual rainfall: 435 mm; annu-
al temperature (max. + min. /2): 16.9°C (N=1).
Habitat: No quantitative assessment; known only from an area compris-
ing finer-grained soils in which the natural vegetation is Renosterveld
shrubland.
Food types: No quantitative assessment; food associations unknown.
Temporal activity: Not known, but presumably shows diurnal flight ac-
tivity; recorded in autumn during the winter rainy season (May).
Bioregions South Africa: Depending on accuracy of map coordinates, oc-
cupied vegetation units could be Swartland Shale Renosterveld (FRs 9)
with clay loam soils (at Stellenbosch) or nearby Swartland Granite
Renosterveld (FRg 2) with sandy loam soils (Fynbos Biome).
Assessment rationale: EOO/AOO unknown; no ecological data avail-
able; high levels of habitat transformation for arable farming or urbani-
sation around the single known locality (FRs 9: 90%; FRg 2: 80%); as-
sessed as Data Deficient (DD), but probably deserves an IUCN threat
category approaching Critically Endangered (CR) on the basis of few
known museum specimens, no records since 1949, and their known
occurrence at a single location in a highly transformed area.
Conservation measures: A survey is required to determine the EOO and
AOO to adequately assess conservation status; this survey should be
2 mm
combined with the collection of quantitative ecological data, particu-
larly to determine if its population density is influenced by clearance of
natural shrubland; not currently known to occur in any protected area.

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SURICATA 6 (2020) 39

Genus Pedaria Castelnau, 1832

Type species and designation: Pedaria nigra Castelnau, 1832, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Boucomont (1922); new species added, subsequently, by
Balthasar (1937, 1939a, 1963b), Janssens (1939a), Ferreira (1962a), Cambefort
(1982), Paulian et al. (1982); partial revisions and 27 new species added by Josso &
Prévost (2003: 19; 2006a: 4; 2009a: 4).

The long-established genus, Pedaria Castelnau, 1832, currently comprises 60 species restricted to Afrotropical forests,
savannas and grasslands, where they show kleptocoprid habits in association with dung buried by large tunnelling
Scarabaeinae, especially Heliocopris (Cambefort 1991). The species are often difficult to separate without extraction of
genitalia as external morphological differences are subtle. Illustrations of aedeagi are only provided for the new species
recently described by Josso & Prévost (2003, 2006a, 2009a).

Except for one new species, the known southern African taxa were described long ago. Therefore, illustrations of aedeagi
are lacking as in the revision of Boucomont (1922), which also seems to have missed the South African species, Pedaria
conformis Péringuey, 1901. Of the eight described species recorded in South Africa, Namibia and/or Botswana, species ac-
counts are provided for five more-or-less validated taxa. The remaining three taxa described from South Africa (P. conformis
Péringuey, 1901; P. aspera Péringuey, 1901; P. alternans Waterhouse, 1890) may or may not be included in a further eight
species that have now been recognised from collections made in this region.

The distribution patterns shown by Pedaria species in southern Africa are not fully represented by the five species for
which species accounts are provided. These species show widespread or restricted savanna distributions with one centred
along the E seaboard of Africa. However, two of the unnamed or unvalidated species show Kalahari or Highveld centred
distributions, respectively.

Despite the poor state of knowledge of conservation status for Pedaria, it is probable that none of the 13 southern African
species currently faces threats. Poor state of knowledge probably results partly from infrequent collection due to cryptic
appearance in dung as well as the subtle taxonomic differences that complicate identification of species.

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40 SURICATA 6 (2020)

Pedaria barrei
Josso & Prévost, 2006a

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘East Kenya, Tsavo East, Voi Lodge’ [Tsavo East National
Park, Kenya].
Taxonomy: Accepted species; very close in appearance to P. cylindrica
Fahraeus, 1857, but aedeagus quite different; four South African local-
ities supported by extraction of aedeagi, three cited by Josso & Prévost
(2006a).
Distribution: Mostly dry lowland savanna from S to E Africa: South Af-
rica, Tanzania, Kenya.
Locality data (mean ± SD, range): Altitude: 496 ± 285, 153–878 m;
annual rainfall: 560 ± 106, 403–710 mm; annual temperature (max. +
min. /2): 21.8 ± 1.0, 20.8–23.1°C (N=7).
Habitat: No quantitative assessment; limited DBRU collection records:
sand (1), sandy loam (1) in open shrub / woodland (2).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (2), buffalo (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Mar.); attracted to light.
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 460 055 km2 (gross estimate); wide-
spread, but difficult to assess conservation status as quantitative habitat
and food association data are unavailable; records from elephant (2)
and zebra dung (1) in Kenya are insufficient to support any bias to mo-
nogastric herbivore dung given southern Africa records from ruminant
herbivore dung; assessed as Data Deficient (DD).
Conservation measures: Quantitative ecological data are required to ad-
equately assess conservation status; protected in Kruger National Park
(South Africa), Tsavo East National Park (Kenya).

2 mm

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SURICATA 6 (2020) 41

Pedaria brancoi
Josso & Prévost, 2003

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Mozambique, Parc de Gorongoza’ [Gorongosa National

Park, central Mozambique].
Taxonomy: Accepted species.
Distribution: Lowland to upland, moist to dry savanna in S Central and
SE Africa: Namibia, Zimbabwe, Mozambique, Tanzania.
Locality data (mean ± SD, range): Altitude: 1 087 ± 528, 59–1 818 m;
annual rainfall: 729 ± 235, 354–1 136 mm; annual temperature (max.
+ min. /2): 20.9 ± 2.5, 18.3–25.2°C (N=8).
Habitat: No quantitative assessment; no data.
Food types: No quantitative assessment; no data.
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Nov. to Jan.; also June); attracted to light.
Ecoregions Namibia, Zimbabwe: N Zambezian and Mopane Woodlands
(AT0725), Southern Miombo Woodlands (AT0719), Zambezian
Baikiaea Woodlands (AT0726), Kalahari Acacia-Baikiaea Woodlands
(AT0709), NW Kalahari Xeric Woodlands (AT1309).
Assessment rationale: EOO = 809 330 km2 (gross estimate); widespread,
but difficult to assess conservation status in absence of ecological data;
assessed as Data Deficient (DD).
Conservation measures: Quantitative data on ecological associations
is required before conservation status may be assessed; protected in
Gorongosa National Park (Mozambique), Serengeti National Park
(Tanzania).

2 mm

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42 SURICATA 6 (2020)

Pedaria cylindrica
Fahraeus, 1857

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘prope fluvium Gariep’ [near Orange River, South Africa].
Taxonomy: Accepted species; very close in appearance to P. barrei Josso &
Prévost, 2006, but aedeagus quite different; out of 18 map localities,
ten are supported by extraction of aedeagi (red squares).
Distribution: Moist southern African savanna: Namibia, Botswana,
South Africa; also cited from Angola, Zimbabwe, Mozambique.
Locality data (mean ± SD, range): Altitude: 810 ± 457, 45–1 378 m;
annual rainfall: 612 ± 130, 306–900 mm; annual temperature (max. +
min. /2): 20.3 ± 1.7, 18.1–22.6°C (N=22).
Habitat: No quantitative assessment; DBRU collection records: sand (2),
sandy loam (5), sandy clay loam (1) in grassland/pasture (2), shrubland
(3), open woodland (3).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (7), wildebeest (1).
Temporal activity: Flight activity unknown, but presumably in darkness
like other Pedaria species; seasonal activity primarily in the summer
rainy season (Oct. to May).
Ecoregions Namibia, Botswana: Southern African Bushveld (AT0717),
Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome).
Assessment rationale: EOO = 127 365 km2 (gross estimate); apparently
widespread in various open vegetation types, on various soil soils types
and on ruminant herbivore dung, but quantitative support lacking;
assessed as Least Concern (LC), but essentially Data Deficient (DD).
Conservation measures: Asessment of conservation status would be
improved by quantitative data on ecological associations; protected
in uMkhuze, Hluhluwe–iMfolozi and Langjan game reserves (South
Africa).

2 mm

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SURICATA 6 (2020) 43

Pedaria picea
Fahraeus, 1857

= Pedaria cuprascens Harold, 1859

= Pedaria sobrina Péringuey, 1901
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: P. picea: ‘prope fluvium Gariep’ [near Orange River, South
Africa]; P. cuprasacens: ‘Cap der guten Hoffnung’’ [Cape of Good
Hope, South Africa]; P. sobrina: ‘Cape Colony (Graham’s Town, Port
Elizabeth)’ [Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Mostly moist, upland and coastal, southern African sandy
savanna: South Africa, Botswana, Namibia; uncommon in central Ka-
lahari, Botswana.
Locality data (mean ± SD, range): Altitude: 860 ± 525, 0–1 759 m; an-
nual rainfall: 664 ± 185, 245–1 003 mm; annual temperature (max. +
min. /2): 19.3 ± 2.0, 13.9–22.6°C (N=73).
Habitat: In Gauteng bushveld (as Pedaria sp. b): extreme bias to deep sand
(753) compared to sandy clay loam (1) with a bias to more open vege-
tation, grassland (347), open woodland (344), shaded thickets (63); in
uMkhuze Game Reserve: extreme bias to deep sand (17.4) compared
to duplex soil (sand over clay: 0.4), sandy clay loam (0), clay (0); large-
ly supported by DBRU collection records: sand (6), sandy loam (5),
sandy clay loam (3) in grassland/pasture (9), open shrub / woodland
(6) and thicket (1).
Food types: In Gauteng bushveld (as Pedaria sp.): bias to omnivore dung
(pig: 67) compared to herbivore dung (horse: 22, cattle: 44); somewhat
similar in Botswana (pig: 17, elephant: 4, cattle: 0); DBRU collection
records on dung of cattle (10), human (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to May); on Gauteng bushveld: seasonal activity peak,
Oct. to Jan., declining from Feb. to Apr. with mainly evening flight in
mid-summer (Nov. and Jan.: 18:00–20:00, fewer 20:00–22:00), but
also minor pre-dawn flight activity (04:00–06:00).
Ecoregions Namibia, Botswana: E Kalahari Acacia-Baikiaea Woodlands
(AT0709), NE Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), cooler,
2 mm
moister parts of Central Bushveld (SVcb) and Lowveld (SVl) (Savanna
Biome); warmer N or E sandy parts of Dry Highveld Grassland (Gh),
Mesic Highveld Grassland (Gm), Sub-Escarpment (Grassland (Gs)
(Grassland Biome); Indian Ocean Coastal Belt Biome (CB), Albany
Thicket Biome (AT).
Assessment rationale: EOO = 388 675 km2; AOO would be smaller and
restricted according to occurrence of sandy soils within EOO; however,
equally abundant in grassland and open woody habitats; conflicting
dung association data, but probably comes readily to dung of domestic
livestock; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by resolution of conflict in dung associations; protected in
uMkhuze Game Reserve, Leeuwfontein and Tswaing nature reserves
(South Africa).

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Pedaria segregis
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Mozambique (Rikatla)’ [Ricatla, S Mozambique].

Taxonomy: Accepted species with a possibly undescribed close relative
centred on Kalahari deep sands and outliers.
Distribution: Sandy soils in moist, lowland, grassy savanna along the E
seaboard of Africa: South Africa, Mozambique, Zimbabwe, Kenya; also
cited from Tanzania; citation from Angola probably an error.
Locality data (mean ± SD, range): Altitude: 213 ± 363, 0–1 354 m;
annual rainfall: 704 ± 112, 482–1 003 mm; annual temperature (max.
+ min. /2): 22.0 ± 1.2, 18.4–23.4°C (N=33).
Habitat: In uMkhuze Game Reserve: extreme bias to deep sand (23.3)
compared to duplex soil (sand over clay: 0.6), sandy clay loam (0.1),
clay (0); on sand in Maputo Special Reserve: very strong bias to fossil
lagoon grassland (10.7 per sample) compared to sand forest patches
(large: 0.1, medium: <0.1, small: 0.4) and coastal dune forest (0); large-
ly supported by DBRU collection records: sand (4), sandy loam (2) in
grassland/pasture (2), open shrub / woodland (4) and thicket (1).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (3), wildebeest (1), rhinoceros (1), horse (1).
Temporal activity: Flight activity unknown, but presumably in darkness
like other Pedaria species; seasonal activity in the summer rainy season
(Oct. to Apr.).
Ecoregions Zimbabwe, Mozambique: Sandy soils in Maputaland Coast-
al Forest Mosaic (AT0119), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Sandy soils in Central Bushveld (SVcb),
Lowveld (SVl), Mopane (SVmp) (Savanna Biome); N Indian Ocean
Coastal Belt Biome (CB).
Assessment rationale: EOO = 730 030 km2 (gross estimate); AOO would
be smaller and restricted according to occurrence of sandy soils and
more open vegetation within the EOO; well represented in reserves in
coastal regions; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on food associations; protected in uMkhuze
Game Reserve (South Africa), Maputo Special Reserve (Mozambique).

2 mm

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SURICATA 6 (2020) 45

Genus Sarophorus Erichson, 1847b

Type species and designation: Pedaria tuberculata Castelnau, 1840, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Frolov and Scholtz (2003a); new species added by Frolov
(2004), Roets et al. (2017).

The long-established genus, Sarophorus Erichson, 1847b, is usually treated as a full genus although Péringuey (1901)
treated it as a subgenus of Pedaria Castelnau, 1840, with which it has often been confused. Recent authors have identified
characters that may justify the separation of two species into another genus although all are currently treated as belonging
to a single genus that is endemic to the S part of the Afrotropical region.

Sarophorus currently comprises 12 valid species divided into two species groups. One (tuberculatus group) comprises six
species centred to the SW in the winter and bimodal rainfall regions of South Africa and at points along the E escarpment
in the summer rainfall region. The other (costatus group) also comprises six species that are mostly, though not entirely,
centred to the NE with most ranges occurring in moist, summer rainfall savanna. Two of these species show ranges entirely
outside of South Africa, Botswana or N Namibia in Zimbabwe, Zambia, SE Democratic Republic of the Congo (DRC)
or Tanzania.

Both species groups overlap either side of the transition between bimodal and summer rainfall areas in the Eastern Cape,
South Africa. All species seem to show a bias to denser vegetation providing shade, including forest in the Eastern Cape
and KwaZulu-Natal. Such shady or dense vegetation associations suggest clearance of shrubs and trees would be detrimen-
tal to population density.

Sarophorus is ecologically poorly known, even though S. costatus (Fahraeus, 1857) is frequently attracted to dung-baited
pitfalls, particularly to omnivore dung. Of the ten species occurring in southern Africa, eight are restricted to South Africa,
one is found in South Africa, Botswana and Zimbabwe with one in N Namibia and Angola.

tuberculatus group: The six group members are centred, respectively, in Renosterveld in Namaqaland (1) or SW Western
Cape (1) (may belong to a second genus), shrubland from SW coastal sands of the Western Cape to the Eastern Cape (1),
forest patches on the S Cape coast (bimodal rain) (1) or S KwaZulu-Natal (1), or upland woodland in E Mpumalanga (1)
(last two both summer rainfall).

costatus group: The four species found in the study region are centred in E savanna (2: but only one widespread), W
savanna (1), or on the S coast of the Eastern Cape (summer to bimodal rain) (1).

The four more widely distributed species have been assessed as Least Concern (LC), whereas six known by few records are
assessed as Data Deficient (DD) with, in particular, three putative forest or upland woodland species potentially Endan-
gered (EN).

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Sarophorus angolensis
Frolov, 2004

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Tschivinguire Huila’ [SW Angola].

Taxonomy: Accepted costatus group species; the Namibian material is not
yet validated as S. angolensis although it is close in appearance.
Distribution: Known from two savanna localities close to the W es-
carpment: type locality is on the highlands of the Bié Plateau, Huila
Province, SW Angola; non-validated locality is at Epembe in the arid
Kaokoveld, N Namibia.
Locality data (mean ± SD, range): Altitude: 1 372 ± 581, 961–1 783 m;
annual rainfall: 583 ± 289, 379–787 mm; annual temperature (max. +
min. /2): 18.2 ± 0.7, 17.7–18.7°C (N=2).
Habitat: No quantitative assessment; Epembe material from white sand
near river.
Food types: No quantitative assessment; no data.
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
at Epembe, recorded during the late summer rainy season (Feb.); at
type locality, recorded in early summer (Nov.).
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702); N Na-
mibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 3 760 km2; small range likely a collecting
artefact; ecological data virtually absent; alpha taxonomy uncertain as
to whether Namibian and Angolan material represents one or two spe-
cies, currently assessed as Data Deficient (DD).
Conservation measures: A quantitative survey of Kaokoveld and SW An-
gola is required to determine the EOO, AOO, ecological associations
and whether the material comprises one or two species; not currently
known from any protected region.

2 mm

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Sarophorus bidentatus
Frolov & Scholtz, 2003a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Namaqualand, Kamieskroon’ [South Africa].

Taxonomy: Accepted tuberculatus group species.
Distribution: Currently known from the type locality on the stepped W
escarpment of South Africa (holotype) and by a further 14 individuals
from ‘Cap bon Esp’ [Cape of Good Hope] deposited in the Muséum
National d’Histoire Naturelle, Paris.
Locality data (average): Altitude: 1 070 m; annual rainfall: 162 mm;
annual temperature (max. + min. /2): 15.5°C (N=1).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; no data available.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
in spring (Sept.) during the winter rainy season.
Bioregions South Africa: May be associated with Namaqualand Granite
Renosterveld (FRg 1) (Fynbos Biome), which occupies a higher rainfall
island within the arid Succulent Karoo Biome; however, survey support
required.
Assessment rationale: EOO not known; no ecological data available;
single known locality may be inexact; Kamieskroon occurs within the
Succulent Karoo at the edge of the upland island of higher rainfall (up
to 300–400 mm p/a) within the area of greater aridity; no known re-
cords since type specimen was recorded in 1930; close relatives show
an association with tall shrubland or shaded thicket and this may be
an important consideration for its conservation; currently assessed as
Data Deficient (DD) although an IUCN threat category approaching
Vulnerable (VU) might be justified on the basis of likely small range
and apparent rarity, even though the local Renosterveld is cited as least
2 mm
threatened.
Conservation measures: A survey to determine the full EOO, AOO and
ecological associations is required to assess conservation status; not cur-
rently known from any reserve.

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Sarophorus carinatus
Frolov & Scholtz, 2003a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Lydenburg Distr., Ohrigstad’ [Mpumalanga, South Africa].

Taxonomy: Accepted tuberculatus group species.
Distribution: Known from only two localities along the NE escarpment,
South Africa.
Locality data (mean ± SD, range): Altitude: 1 281 ± 142, 1 181–1 382 m;
annual rainfall: 875 ± 6, 870–879 mm; annual temperature (max. +
min. /2): 16.8 ± 0.9, 16.2–17.4°C (N=2).
Habitat: No quantitative assessment; holotype recorded from humus;
another eight individuals retrieved from top soil under leaf litter in a
shrub/woodland patch on an upland koppie within a grassland matrix.
Food types: No quantitative assessment; food types unknown, but not
attracted to pitfalls baited with cattle dung at one locality.
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
recorded during the summer rainy season (Nov., Mar.).
Bioregions South Africa: Holotype: Ohrigstad Mountain Bushveld
(SVcb 26) (Central Bushveld, Savanna Biome); eight individuals:
shrub/woodland patch in KaNgwane Montane Grassland (Gm 16)
(Mesic Highveld Grassland, Grassland Biome).
Assessment rationale: EOO = 755 km2 (presumably underestimated);
observations from humus or litter in shrub/woodland suggest the AOO
is much more restricted than the EOO, particularly in the Grassland
Biome; known from only nine individuals collected in 1962 and 2002
although this may be a collection artefact related to specialised habi-
tat and habits; assessed as Data Deficient (DD), but the small known
range at few wooded localities suggests that an IUCN threat category
of Endangered (EN) might be justified; however, quantitative data is
required in support.
Conservation measures: To assess conservation status, a survey is required
to determine the full EOO and AOO, also, to quantify ecological asso-
2 mm ciations; not currently recorded from any protected area.

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Sarophorus costatus
(Fahraeus, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Paedaria costata: ‘in terra Natalensi’ [KwaZulu-Natal,

South Africa].
Taxonomy: Accepted costatus group species.
Distribution: SE African savanna; distribution possibly influenced by
woody vegetation cover: South Africa, E Botswana, S Zimbabwe.
Locality data (mean ± SD, range): Altitude: 926 ± 461, 17–1 824 m;
annual rainfall: 674 ± 134, 339–1 038 mm; annual temperature (max.
+ min. /2): 19.3 ± 2.2, 14.5–24.4°C (N=139).
Habitat: In Gauteng bushveld: slight bias to deep sand (251) compared
to sandy clay loam (155) and strong bias towards woody vegetation of-
fering at least some shade: grassland (7), open woodland (128), shaded
thicket (271); in Suikerbosrand Nature Reserve: recorded only in open
woodland (20), not Highveld Grassland (0) or montane grassland (0); on
sandy clay loam at Roodeplaat Nature Reserve: bias to open woodland
(138), compared to grassland (39) and shaded thicket (18); however, in
uMkhuze Game Reserve, extreme bias to finer-grained soils: sand (0.0),
sand over clay (1.6), sandy clay loam (46.5), clay (4.7) with a 70% bias to
less-shaded vegetation within woodland; DBRU collection records simi-
lar: sand (4), sandy loam (15), sandy clay loam (6), clay (1) in grassland
/ pasture (10), open shrub/woodland (11), shaded woodland/forest (6).
Food types: In Gauteng bushveld: bias to omnivore dung: pig (248), horse
(77), cattle (43), carrion (8), rotting grass clippings (8); similar bias at
Phalaborwa: pig (62), elephant (11), cattle (16); DBRU collection re-
cords reflect commonly sampled dung types: cattle (23), elephant (4),
rhinoceros (5), horse (3), human/cattle mix (3).
Temporal activity: Diurnal flight activity; sampled in Gauteng bushveld
and recorded elsewhere throughout the summer rainy season (Oct. to
Apr.); at Phalaborwa: very strong bias to activity in warm weather soon
after heavy rainfall (75), little activity in hot, dry weather (12) or warm
cloudy weather following light rainfall (2).
2 mm
Ecoregions Botswana, Zimbabwe: Southern African Bushveld (AT0717),
Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl), marginal in Eastern Kalahari Bushveld (SVk) (Savanna
Biome); Indian Ocean Coastal Belt Biome (CB); N margins Mesic High-
veld Grassland (Grassland Biome); also margins of Sub-Escarpment
Savanna (SVs) and Sub-Escarpment grassland (Gs).
Assessment rationale: EOO = 283 960 km2; evidence for soil and vege-
tation associations shows some conflict, particularly soil associations;
however, it is probable that there is a bias to shade or partial shade;
if so, the AOO would be influenced by woody vegetation cover and
impacted upon by its clearance, which is extensive in some parts of
its range; however, S. costatus is widespread and attracted to the dung
of farm livestock despite a bias to omnivore dung; therefore, currently
assessed as Least Concern (LC).
Conservation measures: To improve the assessment of conservation status,
there is a need for further quantitative data on soil and vegetation associa-
tions, particularly the relative importance of shade offered by woody vege-
tation as this could seriously influence AOO; protected in Kruger Nation-
al Park, uMkhuze and Hluhluwe–iMfolozi game reserves (South Africa).

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Sarophorus diabolus
Roets, 2017

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘SOUTH AFRICA, Western Cape Province, Riebeeck-

Kasteel, Porseleinberg, –33.45995 18.88627’.
Taxonomy: Accepted tuberculatus group species.
Distribution: Known only from two adjoining Renosterveld localities in
the winter rainfall region: SW Western Cape in South Africa.
Locality data (mean ± SD, range): Altitude: 334 ± 112, 255–413 m;
annual rainfall: 350 ± 1, 349–351 mm; annual temperature (max. +
min. /2): 16.9 ± 0.5, 16.6–17.3°C (N=2).
Habitat: No published quantitative assessment; type series recorded in
dense, 2–3 m tall, Renosterveld.
Food types: No quantitative assessment; type series sampled to a com-
bined bait of pig dung and chicken liver.
Temporal activity: Diel flight periodicity unknown, but probably diur-
nal; type series recorded mainly during the winter rainy season in late
winter (July); two individuals also recorded in late summer (March).
Bioregions South Africa: Vegetation unit: Swartland Shale Renosterveld
(FRs 9) (West Coast Renosterveld (FO 7), Fynbos Biome).
Assessment rationale: EOO not known; current AOO may be dependent
on persistence of dense, tall, Renosterveld shrubland in a widespread
vegetation unit that is now 90% totally transformed, primarily as arable
farmland; S. diabolus would currently qualify for an IUCN threat cat-
egory approaching Critically Endangered (CR), although it is assessed
as Data Deficient (DD) in the absence of a survey to discover other
localities.
Conservation measures: A quantitative survey of surviving Renosterveld
patches in (FRs 9) and similar vegetation on finer-grained soils in
adjoining areas of the Western Cape is essential to determine the full
2 mm EOO; assessment of conservation status also requires quantitative data
on ecological associations, particularly the effects of removal of dense
shrubland on the AOO; protected in Kasteelberg Nature Reserve,
Western Cape.

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Sarophorus frolovi
Roets, 2017

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘SOUTH AFRICA, KwaZulu-Natal Province, Midlands,

Weza forestry plantations, –30.61680 29.683626’.
Taxonomy: Accepted tuberculatus group species.
Distribution: Known from a single forest locality in the summer rainfall
region: S KwaZulu-Natal in South Africa.
Locality data (average): Altitude: 1 187 m; annual rainfall: 815 mm;
annual temperature (max. + min. /2): 15.3°C (N=1).
Habitat: No quantitative assessment; type series recorded at edges between
Southern Mistbelt Forest and grassland.
Food types: No quantitative assessment; type series sampled to a com-
bined bait of pig dung and chicken liver.
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
recorded during the summer rainy season (Feb.).
Bioregions South Africa: Grid reference for type locality coincides with the
Midlands Mistbelt Grassland vegetation unit (Gs 9) (Sub-Escarpment
Grassland, Grassland Biome) near a patch of Southern Mistbelt Forest
(FOz 3) (Forest Biome).
Assessment rationale: EOO and AOO not known; ecologically extremely
poorly known; would currently qualify for an IUCN threat category of
at least Vulnerable (VU), but in the absence of surveys around other
FOz 3 forest patches, currently assessed as Data Deficient (DD).
Conservation measures: A survey in Sub-Escarpment Grassland around
other patches of FOz 3 is required to determine the full EOO of
S. frolovi; assessment of conservation status also requires quantitative
data on ecological associations, particularly relative abundance in forest
patches versus that in grassland; unclear if S. frolovi is protected under
Weza Plantation management.
2 mm

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Sarophorus latus
Frolov & Scholtz, 2003a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Farm Rhenosterpoort’ [Gauteng, South Africa].

Taxonomy: Accepted costatus group species; although close in appearance
to Sarophorus costatus (Fahraeus, 1857), the aedeagus is quite different.
Distribution: Bushveld savanna in the vicinity of E–W trending moun-
tain ranges (Magaliesberg; Soutpansberg), South Africa.
Locality data (mean ± SD, range): Altitude: 1 091 ± 165, 956–1 331 m;
annual rainfall: 633 ± 40, 594–689 mm; annual temperature (max. +
min. /2): 19.0 ± 0.9, 17.8–19.5°C (N=4).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; recorded on carrion (dead bird)
plus dung of baboon and horse.
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
recorded during the summer rainy season (Dec. to Apr.).
Bioregions South Africa: Vegetation units: Moot Plains Bushveld (SVcb
8), Loskop Mountain Bushveld (SVcb 13), Soutpansberg Mountain
Bushveld (SVcb 21) (Central Bushveld Bioregion, Savanna Biome).
Assessment rationale: EOO = 12 270 km2 (includes areas between moun-
tain ranges); AOO may be restricted to mountain bushveld, but quanti-
tative support required; could be susceptible to habitat transformation
(28%, SVcb 8; 3%, SVcb 13; 21%, SVcb 21), but no accurate data on
vegetation associations; assessed as Data Deficient (DD) although it
may currently warrant an IUCN threat category of Vulnerable (VU) on
the basis of restricted range at few known locations.
Conservation measures: To assess conservation status, a quantitative sur-
vey of mountain bushveld and adjoining vegetation units is required
to determine the EOO, AOO and ecological associations; protected in
Happy Rest and Rhenosterpoort nature reserves.

2 mm

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Sarophorus punctatus
Frolov & Scholtz, 2003a

No synonyms
Global: EN (see IUCN Red List – EN)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Keurboomstrand’ [Eastern Cape, South Africa].

Taxonomy: Accepted tuberculatus group species.
Distribution: Known only from the type locality on the coastline of the
Eastern Cape, South Africa.
Locality data (average): Altitude: 0 m; annual rainfall: 458 mm; annual
temperature (max. + min. /2): 18.0°C (N=1).
Habitat: No adequate quantitative assessment; sampled using ground
traps set from the edge into disturbed podocarp forest.
Food types: No quantitative assessment; unknown.
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
recorded during mid-summer (Dec.) in the bimodal rainfall region.
Bioregions South Africa: Sampled from Southern Afrotemperate Forest
(FOz 1) (Forest Biome) although grid reference coincides with adjoin-
ing South Outeniqua Sandstone Fynbos (FFs 19) (Fynbos Biome).
Assessment rationale: EOO and AOO not known, but AOO possibly
dependent on patches of forest along the Eastern Cape coastline, estab-
lished on sandstone and granite-derived soils; essentially Data Deficient
(DD) although tentatively assessed as Endangered (EN) in line with
current IUCN Red List, which focuses on the low number of known
individuals from a single locality recorded ~40 years ago (1976); further
data are required for a more accurate assessment.
Conservation measures: Assessment of conservation status as EN needs
to be supported by a survey of habitats at the type locality and of similar
habitats at distance from Keurboomstrand; this study should determine
the EOO, AOO, ecological associations and current levels of threat;
elements of FOz 1 are now protected in Garden Route National Park
close to the type locality.

2 mm

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Sarophorus striatus
Frolov & Scholtz, 2003a

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘23 km S of Avontuur’ [Eastern Cape, South Africa].

Taxonomy: Accepted costatus group species.
Distribution: Centred on Albany Thicket along the S coastal region of the
Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 371 ± 248, 27–871 m; an-
nual rainfall: 501 ± 100, 390–741 mm; annual temperature (max. +
min. /2): 17.2 ± 1.1, 15.0–19.2°C (N=13).
Habitat: No quantitative assessment; limited DBRU collection records
inconclusive: sandy loam (2), sandy clay loam (1) in grassland/pasture
(2), open woodland (1).
Food types: No quantitative assessment; DBRU collection records all
from cattle dung (6).
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
recorded in late summer and during the autumn peak (Jan. to May) of
the bimodal rainy season.
Bioregions South Africa: Vegetation units: Gamtoos Thicket (AT 4),
Sundays Thicket (AT 6), Kowie Thicket (AT 8), Albany Coastal Belt
(AT 9) (Albany Thicket Biome); also Bhisho Thornveld (SVs 7) (Sub-
Escarpment Savanna); Kouga Sandstone Fynbos (FFs 27), Langkloof
Shale Renosterveld (FRs 17) (Fynbos Biome).
Assessment rationale: EOO = 6 740 km2; relatively limited known range;
vegetation associations unclear, but some relatives are shade specialists;
therefore, as this species is centred on a thicket biome, clearance of
thicket may be detrimental to population density and/or lead to reduced
AOO; however, quantitative support required; current transformation
in thicket stands at 6–17% (AT 4, AT 6, AT 8, AT 9), mainly due to
cultivation and urbanisation; currently assessed as Least Concern (LC)
although a quantitative assessment is recommended.
Conservation measures: Assessment of conservation status would be
2 mm
much improved by quantitative data on ecological associations, partic-
ularly if there is a bias to shade and a threat due to clearance of woody
vegetation; not currently known from any protected area.

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SURICATA 6 (2020) 55

Sarophorus tuberculatus
(Castelnau, 1840)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Pedaria tuberculata: ‘Afrique’ [Africa].

Taxonomy: Accepted tuberculatus group species.
Distribution: SW Western Cape and Eastern Cape in the winter and
bimodal (spring/autumn) rainfall regions, South Africa; disjunction
between W/E localities possibly a collection artefact.
Locality data (mean ± SD, range): Altitude: 445 ± 464, 0–1 383 m;
annual rainfall: 386 ± 119, 242–667 mm; annual temperature (max. +
min. /2): 15.9 ± 1.2, 13.2–18.0°C (N=17).
Habitat: No adequate quantitative assessment; in Western Cape: sampled
on both sand (45) and sandy loam (73) in tall shrubland at Farm Pam-
poenvlei, but absent from low profile scrub on deep sands of nearby
coastal plain (Farm Modderrivier); also sampled on deep sand in tall
shrubland (69) at Farm Geelbek, but rare in adjoining sparse pasture
cleared of shrubs (2); supported by limited DBRU collection records
from sand (3), sandy loam (1), sandy clay loam (1) in grassland (1) and
shrubland (5).
Food types: No quantitative assessment, but sampled to cattle dung in
Western Cape; DBRU collection records entirely from dung of cattle
(8), but quantitative assessment required.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
and primarily in spring (July to Oct.) and autumn (Mar. to May) in the
winter and bimodal rainy seasons; at farms Geelbek and Pampoenvlei
in Western Cape: limited late July activity, but primarily active in Aug.,
declining in Sept. and rare in Oct., also rare autumn activity in May.
Bioregions South Africa: In the W: Strandveld (FS 3, FS 5), Granite
Renosterveld (FRg 2); in the E: Sand Fynbos (FFs 19, FFs 23, FFs 27)
(all Fynbos Biome); also Albany Thicket Biome (AT).
Assessment rationale: EOO = 46 695 km2; available ecological data sug-
gest the AOO would be much smaller according to occurrence of tall
shrubland, which is under particular threat in the W (transformation 2 mm
in the E: 2–28% in FFs 19, FFs 23, FFs 27; W: 35-80% in FS 3, FS
5, FRg 2); food associations not tested, but clearly attracted to dung
of farm livestock; currently assessed as Least Concern (LC) on basis of
large EOO and protection in at least one reserve in the W.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative ecological data, particularly the effects of
shrubland clearance; protected on Farm Geelbek, which is now part of
West Coast National Park where the sparse grassland pasture has been
permitted to regenerate as shrubland.

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TRIBE CANTHONINI
van Lansberge, 1874a
[TRIBE DELTOCHILINI Lacordaire, 1856]

As Deltochilides; Type genus: Deltochilum Eschscholtz, 1822.

As Canthonides; Type genus: Canthon Hoffmansegg, 1817.

Synonyms:
= Minthophilides Lacordaire, 1856; Type genus: Mentophilus Castelnau, 1840.
= Scatonomides Lacordaire, 1856; Type genus: Scatonomus Erichson, 1835.
= Epilissinides van Lansberge, 1874; Type genus: Epilissus Dejean, 1836.
= Coprobiadae Burmeister, 1873; Type genus: Coprobius Latreille, 1829 [= syn. of Canthon Hoffmansegg, 1817].
= Epirinides van Lansberge, 1874a; Type genus: Epirinus Dejean, 1833 (revalidated as tribe Epirinini van Lansberge,
1874a (Daniel et al. 2020)).
= Panelini Arrow, 1931; Type genus: Panelus Lewis, 1895.

Although the tribal name, Deltochilini, takes precedence (Bouchard et al. 2011), according to Smith
(2006), Canthonini should be maintained as the tribal name through long-term common usage. As a result,
either Deltochilini, Canthonini or both have been cited as the tribal name in recent publications. Even the
recent partial tribal revision of Tarasov and Dimitrov (2016) will not easily resolve this controversy as the
closely related type genera, Deltochilum and Canthon (Monaghan et al. 2007), occur in the same Neotrop-
ical and Nearctic clade now designated as the tribe, Deltochilini (Canthonini) sensu novo.

In the old sense, Canthonini are globally represented within areas of tropical, warm temperate and winter
rainfall climate. However, global molecular phylogeny clearly shows that the current tribal membership is
polyphyletic at generic level with seven distantly related, major lineages for just 36% of the tribal generic-
level membership (Monaghan et al. 2007) and at least 14 lineages in the recent partial tribal revision of Tara-
sov and Dimitrov (2016). As a result, their redefinition of the tribe as Deltochilini (Canthonini) sensu novo
comprises much reduced membership restricted to the Americas. Some American and all non-American gen-
era (Africa, Madagascar, Oriental, Australasia) are given the status of incertae sedis. Under these circumstanc-
es, we retain the old classfication system for arranging the genera in this book, pending a full tribal revision.

At the present time, 101 genera are assigned to the old tribe on morphological criteria. These primarily
occur in four continental biogeographical centres (Africa: 27; Oriental: 4; Americas/Caribbean: 31; Aus-
tralia/New Guinea: 19) with limited sharing: Africa/Oriental (1), Afro-Eurasia (1). A number of genera are
endemic to islands: Madagascar (7), Mauritius (1), New Caledonia (8), New Zealand (2).

Of the 27 genera and 90 species recorded for the Canthonini (old sense) in the Afrotropical region, most
are restricted to arid and winter rainfall regions of the SW or in forests and upland grasslands along the
S and E seaboards as far as SE Kenya. Essentially, only one canthonine genus is widely represented in the
African tropics (Chalconotus Dejean, 1833: 9 spp.) although two others have limited spot representation in
W Africa and/or NE Africa (Odontoloma Boheman, 1857: 20 spp.; Pycnopanelus Arrow, 1931: 2+1 Oriental
spp.). Past doubts have been expressed concerning the inclusion of some of these genera in the Canthonini
(Scholtz & Howden, 1987: Pycnopanelus; Howden & Scholtz, 1987: Odontoloma).
SURICATA 6 (2020) 57

A total of 23 canthonine genera and 76 species are represented in South Africa, Botswana and Namibia,
although none of these belong to the tribe in the new sense. Chalconotus and Pycnopanelus are represented
in all three countries, Dicranocara Frolov & Scholtz, 2003b, and Odontoloma are shared between Namibia
and South Africa, Gyronotus van Lansberge, 1874a (8 spp.) occurs along the E seaboard from South Africa
to Tanzania. The remaining genera are found only in Namibia (5) or South Africa (13).

Molecular phylogeny suggests that 18 of these 23 genera represent seven distinct lineages (Mlambo et al.
2015, Tarasov & Dimitrov 2016). In rank order from most basally to most terminally derived, the generic
lineages are typified by (numbers of southern African species in brackets):
(1) A branched clade, now comprising three new tribes (Davis et al. 2019):
(A) Extremely small-bodied Odontoloma (15) mostly centred in southern Africa (now in the new
tribe Odontolomini Davis, Deschodt & Scholtz, 2019).
(B) Arid SW seaboard genera: Byrrhidium Harold, 1869a (2); Dicranocara Frolov & Scholtz,
2003 (4); Namakwanus Scholtz & Howden, 1987 (2); that may be associated with rock
hyrax middens or concentrations of dung (now in the new tribe Byrrhidiini Davis, Deschodt
& Scholtz, 2019).
(C) SE seaboard forest litter genera: Outenikwanus Scholtz & Howden, 1987 (1); Silvaphilus
Roets & Oberlander, 2010 (1); Endroedyolus Scholtz & Howden, 1987 (1); Aliuscanthoniola
Deschodt & Scholtz, 2008 (1); Peckolus Scholtz & Howden, 1987 (3) (now in the new tribe
Endroedyolini Davis, Deschodt & Scholtz, 2019).
(2) Circellium Latreille, 1829 (1) in sandy winter and bimodal rainfall shrubland, S coast, South Africa
with close relationships to Scarabaeini.
(3) E seaboard genus (South Africa to SE Kenya), Gyronotus van Langsberge, 1874 in South African and
Eswatini forest patches or moist, cool grasslands (5).
(4) Bohepilissus Paulian, 1975, in S coast forest litter, South Africa (2) with close relationships to some
Madagascar, Neotropical and Australian genera.

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(5) Chalconotus Dejean, 1833 (cited as Anachalcos Hope, 1837) (1); single savanna species distributed
from S to E to W Africa.
(6) Epirinus Dejean, 1833 (29) in cooler South African climates: upland NE grassland and SW shru-
bland, SE forests, SW winter and bimodal rainfall shrubland; transferred to the tribe Sisyphini sensu
novo by Tarasov and Dimitrov (2016) but removed to the revalidated tribe, Epirinini van Lansberge,
1874a, by Daniel et al. (2020).
(7) Less typical canthonines in the arid SW: Pycnopanelus Arrow, 1931 (1); desert sands on the SW sea-
board: Hammondantus Cambefort, 1978 (1); or shrublands of the SW winter rainfall region: Aphen-
goecus Péringuey, 1901 (2); plus, surprisingly, one SE forest litter genus: Dwesasilvasedis Deschodt
& Scholtz, 2008 (1) grouped together with Lineage 4 Coprini genera, Macroderes Westwood, 1842
(SW Cape), Xinidium Harold, 1869a (SE highlands).

Several genera are missed from these phylogenies; two SE forest litter genera: Nebulasilvius Deschodt &
Scholtz, 2008 (2), Parvuhowdenius Deschodt & Scholtz, 2008 (1) (= new tribe: Endroedyolini); three arid
SW seaboard genera: Versicorpus Deschodt, Davis & Scholtz, 2011 (2), Drogo Deschodt, Davis & Scholtz,
2016 (1), Namaphilus Deschodt & Davis, 2017 (3) (= new tribe: Byrrhidiini).

The recent global phylogeny and partial tribal revision of Tarasov and Dimitrov (2016) largely supports
the divisions of Mlambo et al. (2015), but the lineages are more widely separated in the global analysis by
taxa from other biogeographical regions. Tarasov and Dimitrov (2016) classify Lineage 1 (above) as ‘Basal
Scarabaeinae’ along with most African genera of the Ateuchini (old sense), except Pedaria. Lineages 2 to 6
are also in separarate lineages with Epirinus removed to the tribe Sisyphini (but see notes under (6) above).
The genera of Lineage 7 are separated into four neighbouring lineages along with Macroderes and Xinidium,
here included in the Coprini (old sense).

Membership of this polyphyletic tribe (old sense) is frequently characterised by ball-rolling behaviour,
although in southern Africa this has been observed only in larger-bodied genera such as Circellium and
Chalconotus or in genera of smaller body size such as Epirinus. Different habits are probably shown by most
genera (see Dicranocara; also tunnelling observed in Aphengoecus). Habits of those genera showing smallest
body size are unknown, including Odontoloma and those many genera found in forest litter along the S
and SE seaboard.

Most flightless species with small ranges in South African fynbos and forest are assessed as facing threats
(Critically Endangered (CR): 1; Endangered (EN): 6; Vulnerable (VU): 17) including a few flying species
(VU: 4).

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Genus Aliuscanthoniola Deschodt & Scholtz, 2008

Type species and designation: Aliuscanthoniola similaris Deschodt & Scholtz, 2008, by original designation.
Synonyms: None.
Last review: Genus created by Deschodt and Scholtz (2008).

Aliuscanthoniola Deschodt & Scholtz, 2008 is monotypic and is currently known from only a single small patch of forest
on the SE coast of South Africa. In view of the very small range at a single known threatened locality, it is currently assessed
as Endangered (EN).

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Aliuscanthoniola similaris
Deschodt & Scholtz, 2008

No synonyms
Global: EN
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. Afr.; Transkei, Ntsubane For. St.’ [Ntsubane Forest Sta-
tion, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Known only from Ntsubane Forest, an element of the Crit-
ically Endangered Scarp Forest on the Pondoland coast.
Locality data (average): Altitude: 55 m; annual rainfall: 961 mm; annual
temperature (max. + min. /2): 19.7°C (N=1).
Habitat: No quantitative assessment; recorded on finer-grained soils in
forest leaf litter.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the summer rainy season (Nov., Dec.)
Bioregions South Africa: A single patch of the Scarp Forest vegetation
unit (FOz 5) (Forest Biome).
Assessment rationale: EOO not known but type was recorded from an
irregular forest patch comprising only 61.1 km2; Ntsubane is signifi-
cantly threatened by utilisation of resources and peripheral fires; in view
of the small known range at a single threatened location, assessed as
Endangered (EN).
Conservation measures: Continuing preservation of Ntsubane State For-
est (94.46 km2) is essential for the protection of the monotypic genus,
Aliuscanthoniola; as the EOO and AOO might extend southwestwards
to other nearby forest patches around Port St Johns, investigation of
these forests would be advantageous, particularly as all are similarly
threatened.

1 mm

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Genus Aphengoecus Péringuey, 1901

Type species and designation: Aphengoecus clypeatus Péringuey, 1901, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Scholtz and Howden (1987).

Aphengoecus Péringuey, 1901, comprises two flightless species of uncertain habits that are restricted to the SW of the
Western Cape. The possibility of association with termite nests needs to be investigated. In view of their small ranges and/
or low frequency of records in an area of high habitat modification, they presumably face threats. Available data result in
assessments of either Vulnerable (VU) or Data Deficient (DD).

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Aphengoecus clypeatus
Péringuey, 1901

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Colony (Stellenbosch)’ [Western Cape, South Afri-

ca].
Taxonomy: Accepted species.
Distribution: Known only from the type locality of Stellenbosch and in-
exact localities cited from the Cape and southern Africa.
Locality data (average): Altitude: 228 m; annual rainfall: 435 mm; annu-
al temperature (max. + min. /2): 16.9°C (N=1).
Habitat: Not known, but dominant natural habitat in two vegetation
units around Stellenbosch is shrubland on sandy loam.
Food types: Possibly associated with the refuse heaps of termites whose
nests were close to the traps that captured the type series of 27 individ-
uals; not recorded elsewhere in the study area.
Temporal activity: Flightless, nocturnal activity; seasonal activity un-
known.
Bioregions South Africa: Stellenbosch lies in the West Renosterveld
Bioregion (FO 7) (Fynbos Biome) at the edge of two vegetation units,
Swartland Granite Renosterveld (FRg2) and Swartland Shale Renos-
terveld (FRs 9).
Assessment rationale: EOO unknown; only known by the type series,
collected in 1892, and nine other individuals; unclear if subsequent
absence of collection is due to specialised habits or radical habitat trans-
formation in the Stellenbosch area (FRg 2: 80%; FRs 9: 90%); assessed
as Data Deficient (DD), but these circumstances probably warrant at
least an Endangered (EN) threat category, probably Critically Endan-
gered (CR).
2 mm
Conservation measures: A survey needs to be mounted in the Stellen-
bosch region to determine if this species is still extant; if rediscovered,
its EOO, AOO and ecological associations need to be determined to
assess conservation status; sampling in the vicinity of termite mounds
would be recommended; not known from any protected area.

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Aphengoecus multiserratus
Scholtz & Howden, 1987

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape, Cederberg, Jeep track, 1 280 m’ [Western Cape,

South Africa].
Taxonomy: Accepted species.
Distribution: Restricted to a small area along the dry SW coastline of the
Western Cape, South Africa and adjoining dry mountains (Cederberg).
Locality data (mean ± SD, range): Altitude: 194 ± 298, 3–1 280 m;
annual rainfall: 245 ± 64, 146–350 mm; annual temperature (max. +
min. /2): 17.1 ± 0.9, 15.2–18.5°C (N=16).
Habitat: Sampled only on deep sand in natural shrubland: on dunes at
Farm Pampoenvlei (31), in strandveld at Farm Modderrivier (9).
Food types: Attracted in small numbers to faeces, cattle dung, meat and
banana bait.
Temporal activity: Flightless, diel periodicity unknown; at S edge of
range: quantitative data for seasonal activity strongly biased to autumn
tailing off into winter (Apr. to July); strong bias to spring (Aug., Sept.)
in reference material, but from traps left 59–63 days.
Bioregions South Africa: Centred on Western Strandveld (FS) and Sand
Fynbos (FFd) vegetation units along the coast and in Sandstone Fynbos
(FFs) on the Cederberg (Fynbos Biome); also in Namaqualand Strand-
veld (SKs 7) at the S end of the Succulent Karoo Biome.
Assessment rationale: EOO = 3 700 km2; occurs on the coastal strip of
the farm, Modderrivier, run as the private Grotto Bay Nature Reserve,
but little of the known range coincides with officially protected areas;
furthermore, there is high habitat transformation across its small range
(FS, FFd: 25–55%; FFs: 15%); an EOO < 20 000 km2 and AOO prob-
ably < 2 000 km2 at a limited number of locations in an area subject 2 mm

to extensive transformation would qualify this species for Vulnerable

(VU) status.
Conservation measures: In view of possible ecological specialisation,
probable low vagility due to flightless habits, and limited protection
in reserves, a study of its AOO and ecological associations is required
to better assess conservation status; owing to observations made on the
habits of its congener, A. clypeatus Péringuey, 1901, possible association
with the refuse heaps of termites or ants should be investigated; not
recorded in West Coast National Park, but protected in the Cederberg
Wilderness Area.

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Genus Bohepilissus Paulian, 1975

Type species and designation: Epilissus subtilis Boheman, 1857, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Scholtz and Howden (1987).

Bohepilissus Paulian, 1975, currently comprises two very small-bodied, flightless, forest endemic species. One shows a
restricted range in the highly modified SW Cape and is assessed as Vulnerable (VU). The other is widespread in forest
patches along the S and SE coasts, but may represent a species complex. Both have been recorded in forest litter.

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Bohepilissus nitidus
(Balthasar, 1965a)

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Panelus nitidus ‘Süd-Afrika aus der Umgebung von Kap-
stadt’ [environs of Cape Town, Western Cape, South Africa].
Taxonomy: Accepted species.
Distribution: South Africa, Western Cape, Cape Town: primarily E slopes
of Table Mountain with one record from a mountainside at Muizen-
berg.
Locality data (mean ± SD, range): Altitude: 485 ± 376, 93–994 m; an-
nual rainfall: 487 ± 63, 415–555 mm; annual temperature (max. +
min. /2): 14.1 ± 1.8, 11.7–16.1°C (N=6).
Habitat: Probably primarily a forest litter specialist; sampling on Table
Mountain supports forest bias (J. Pryke, unpublished data): young for-
est (5.5/trap), natural forest (2.2/trap), riverine forest (1.7/trap), also in
nearby pine plantation (1.8/trap) plus fynbos, regenerating fynbos and
cultivated gardens (all 1/trap).
Food types: No quantitative assessment; type series recorded in humus;
sampled using pig dung.
Temporal activity: Flightless; diel periodicity not recorded; sampled in
spring, early summer and autumn (July to Jan., Apr.).
Bioregions South Africa: Centred on the most westerly extent of South-
ern Afrotemperate Forest (FOz 1) patches from which most samples
were recorded.
Assessment rationale: EOO ~25 km2, known AOO ~3 km2 (area of FOz
1 on E slopes and ravines of Table Mountain); assumed to be originally
associated with FOz 1 forest patches like its congener, B. subtilis (Bo- 1 mm
heman, 1857); owing to small EOO and even smaller AOO within the
municipal boundaries of a major city, assessed as Vulnerable (VU), even
though most known localities occur within a national park and some
recent records originate from transformed habitats.
Conservation measures: Continuing protection of forest patches in Cape
Town is probably essential for the conservation of this species despite
some records from transformed habitats where it occurs at lower popu-
lation density; protected within Table Mountain National Park.

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Bohepilissus subtilis
(Boheman, 1857)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Epilissus subtilis: ‘Caffraria meridionali’ [inexact, proba-

bly S South Africa, possibly near Port Elizabeth].
Taxonomy: Accepted species, but may comprise a species complex owing
to variation in setae, punctation, eye shape and aedeagi; this is repre-
sented as a geographical mosaic rather than a cline.
Distribution: S coastal region of Western and Eastern Cape with outliers
on the NE coast of the Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 422 ± 479, 0–1 993 m;
annual rainfall: 492 ± 130, 356–961 mm; annual temperature (max. +
min. /2): 16.2 ± 2.3, 9.6–19.7°C (N=29).
Habitat: No quantitative assessment; many records from relict patches of
forest.
Food types: No quantitative assessment; mostly sifted from forest litter;
also sampled to dung and carrion.
Temporal activity: Flightless; diel periodicity unknown; recorded in
spring to early summer and autumn (Aug. to Mar.) reflecting seasonal
rainfall of the bimodal rainfall region in the centre of its range.
Bioregions South Africa: Primarily in Southern Afrotemperate Forest
(FOz 1) patches on the S coast and coastal mountains; outliers in Scarp
Forest to the NE (FOz 5) (Forest Biome).
Assessment rationale: EOO = 38 100 km2; assuming that B. subtilis
represents a single variable species that differs between most neigh-
bouring localities, it would not currently be seriously threatened as it
is widespread and occurs in rather more than 10 separate locations,
many of which are now protected within a national park; an unknown
proportion of FOz 1 has been cleared in the past; assessed as Least
1 mm Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; however,
continuing protection of FOz 1 forest patches would presumably be
essential for the conservation of this species and its varieties; protected
in the Garden Route National Park.

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Genus Byrrhidium Harold, 1869

Type species and designation: Byrrhidium ovale Harold, 1869a, by subsequent designation (Janssens, 1938).
Synonyms: = Elassocanthon Kolbe, 1908: Type species: Elassocanthon brevipes Kolbe, 1908, by
monotypy.
Last review: Entire genus reviewed by Scholtz and Howden (1987); synonymy of two species De-
schodt et al. (2016).

Following the revision of Scholtz and Howden (1987) and the recent synonymy of two species (Deschodt et al. 2016),
Byrrhidium Harold, 1869, is now reduced from three to just two species restricted to arid areas of Namaqualand in South
Africa. Conservation of this genus may be dependent on persistence of concentrations of dung at hyrax middens or near
refuges for rodents. However, this possibility needs to be tested.

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Byrrhidium convexum
Scholtz & Howden, 1987

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Nunies/Sendelingsdrift at 28°18’S, 16°58’E’ [South Africa

on Namibian border].
Taxonomy: Accepted species.
Distribution: Extremely arid mountains of the Richtersveld, Northern
Cape, in the arid north of the winter rainfall region, NW South Africa.
Locality data (mean ± SD, range): Altitude: 452 ± 187, 167–732 m;
annual rainfall: 57 ± 11, 42–74 mm; annual temperature (max. + min.
/2): 16.6 ± 0.7, 15.6–17.9°C (N=7).
Habitat: Sampled on sandy loam amongst shrubs and boulders of a rocky
outcrop.
Food types: Sampled close to hyrax middens using pig dung bait.
Temporal activity: Flightless; diel periodicity unknown; recorded during
spring (Oct., Nov.) and autumn (Apr.).
Bioregions South Africa: Vegetation unit: N tip of Central Richtersveld
Mountain Shrubland (SKr 1) (Richtersveld, Succulent Karoo Biome);
peripheral records in mountain or hilly vegetation units of the Desert
Biome (Dg 2, Dg 3: inland Gariep Desert Bioregion; Dn 5: coastal
Namib Desert Bioregion).
Assessment rationale: EOO = 163 km2; poorly known with just a few
qualitative observations of ecological associations; would qualify for
an IUCN threat category on the basis of small known range at few
locations; however, all but one of these known localities lie within a
protected area; currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative sur-
vey of the Richtersveld is required to determine the full EOO, AOO
and ecological associations; protected in Richtersveld National Park.

2 mm

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Byrrhidium ovale
Harold, 1869a

= Elassocanthon brevipes Kolbe, 1908

= Byrrhidium namaquensis Scholtz & Howden, 1987
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: B. ovale: ‘Port Natal’ [Durban, KwaZulu-Natal, South

Africa; undoubtedly a locality error for this species, currently known
only from Namaqualand]; E. brevipes: ‘Klein-Namaland: Kamaggas’
[Kommaggas, Namaqualand, South Africa]; B. namaquensis: ‘Garies,
Namaqualand’ [South Africa].
Taxonomy: Accepted species.
Distribution: Arid regions above and below the W escarpment: Central to
S Namaqualand, in the winter rainfall region of South Africa; citation
for B. ovale from Namibia represents confusion over the location of
‘Kamaggas’ (E. brevipes type locality), which is situated in Little Nama­
qualand, South Africa, NOT Great Namaqualand in Namibia.
Locality data (mean ± SD, range): Altitude: 689 ± 373, 298–1 179 m;
annual rainfall: 143 ± 25, 98–163 mm; annual temperature (max. +
min. /2): 16.8 ± 1.2, 15.2–18.7°C (N=6).
Habitat: Sampled on sandy loam in relatively undisturbed shrubland with
greater observed frequency close to heuweltjies (fossil termite mounds)
inhabited by small mammals.
Food types: Sampled to pig dung, but possibly supported primarily by
dung of small mammals inhabiting heuweltjies.
Temporal activity: Flightless, diel periodicity uncertain; either active in
late afternoon, during darkness, or in early morning; recorded during
the winter rainy season (July to Sept.); sampled under cool tempera-
tures after rainfall in spring.
Bioregions South Africa: Vegetation unit: NW Namaqualand Klipkoppe
Shrubland (SKn 1) (Namaqualand Hardeveld Bioregion, Succulent
Karoo Biome).
Assessment rationale: EOO = 2 370 km2; poorly known with ecological 2 mm
data supported only by qualitative observations; frequency apparent-
ly lower in disturbed habitat (e.g. overgrazing); probably deserves an
IUCN threat category of at least Vulnerable (VU) on the basis of small
range, < 10 known localities and probable sensitivity to grazing pres-
sure; however, currently assessed as Data Deficient (DD).
Conservation measures: To adequately assess conservation status, a quan-
titative survey is required to document the full EOO, AOO and eco-
logical associations, particularly bias to heuweltjies and possible effects
of overgrazing and habitat disturbance; a small portion of the SKn 1
vegetation unit (4%) is protected in Namaqua National Park.

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Genus Chalconotus Dejean, 1833

Type species and designation: Scarabaeus cupreus Fabricius, 1775, by monotypy.
Synonyms: = Anachalcos Hope, 1837: Type species: Scarabaeus cupreus Fabricius, 1775, by mono-
typy.
Last review: Entire genus reviewed as Anachalcos Hope, 1837, by Janssens (1938a); new species of
Anachalcos added by Müller (1942), Josso and Prévost (2001).

Chalconotus Dejean, 1833, was recently revalidated as the senior generic name for species long classified in the genus,
Anachalcos Hope, 1837 (Branco 2011). The large-bodied genus is endemic and widely represented in savannas and forest
of the Afrotropical region where it is represented by nine species, all but one of which show intertropical distributions.
The one exception that is also represented in southern Africa shows a pan-dry savanna distribution S of the Sahara and is,
therefore, assessed as Least Concern (LC).

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Chalconotus convexus
Boheman, 1857
= Anachalcos obscurus van Lansberge, 1882
= Anachalcos sericeus Felsche, 1907
Global: LC

J A S O N D J F M A M J

Type localities: C. convexus: ‘Caffraria interior’ [inexact, interior of South

Africa]. A. obscurus: ‘Somalis (Afrique Équatoriale)’ [inexact, Somalia];
A. sericeus: ‘Kionga’ [Quionga, N Mozambique].
Taxonomy: Accepted species; recently transferred to Chalconotus Dejean,
1833, from Anachalcos Hope, 1837.
Distribution: Widespread in drier savanna, S to E to W Africa: South
Africa, Mozambique, Zimbabwe, Botswana, Namibia, Angola, Kenya,
Tanzania, Somalia, Nigeria; also reported from Malawi, Democratic Re-
public of the Congo, Uganda, Ethiopia, Eritrea, Benin, Côte d’Ivoire,
Burkina Faso, Senegal.
Locality data (mean ± SD, range): Altitude: 790 ± 509, 0–1 850 m; an-
nual rainfall: 646 ± 191, 22–1 254 mm; annual temperature (max. +
min. /2): 20.3 ± 2.3, 13.4–25.9°C (N=335).
Habitat: In Gauteng bushveld: bias to finer-grained soils, sandy clay loam
(346), deep sand (110); in uMkhuze Game Reserve: more equitable
distribution, deep sand (13.8), sand over clay (22.2), sandy clay loam
(15.8), clay (7.7); in Gauteng bushveld: strong bias to shaded thickets
(270), open woodland (167), grassland (19); largely supported by DBRU
collection records: sand (10), sandy loam (5), sandy clay loam (7), clay
(4) in forest (16), open shrub/woodland (19) and grassland/pasture (5).
Food types: In Gauteng bushveld: readily attracted to cattle dung in hab-
itat sampling, but quantitative trophic sampling shows extreme bias
to carrion (60) and omnivore dung (pig: 112) compared to herbivore
dung (horse: 1, cattle: 4); not so extreme at Phalaborwa: dung of pig
(542), elephant (108), cattle (36); DBRU collection records on dung:
mixed human/cattle or buffalo (13), elephant (13), rhinoceros (2),
5 mm
horse (1), cattle (9), wildebeest (1).
Temporal activity: Flight activity in darkness, but also observed construct-
ing cattle dung balls in early morning light following rainfall; in Gauteng
bushveld: active during the summer rainy season with peak activity in
early to mid-summer (Oct. to Feb.) tailing off into late summer (Mar.
to Apr.); also spring activity (Aug.) at the warmer KwaZulu-Natal coastline; at Phalaborwa: abundant activity in warm weather
after light rain (185), heavy rain (302) or in dry conditions (202); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Centred on Southern African Bushveld (AT0717), Zambezian and Mopane
Woodlands (AT0725), Southern Miombo Woodlands (AT0719), Namibian Savanna Woodlands (AT131), Angolan Mopane
Woodlands (AT0702); E and W separated by deep Kalahari sand ecoregions.
Bioregions South Africa: Primarily Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl); also Sub-Escarpment Savanna
(SVs) (Savanna Biome), Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 6 750 790 km2 (gross estimate); although bias to shaded and woody habitats suggest that local
clearance of woodland would be a threat, it would be little threatened over most of its extremely widespread range throughout
drier savanna of Africa where it readily colonises cattle dung despite bias to omnivore dung and carrion; assessed as Least
Concern (LC).
Conservation measures: None required; protected in various national parks and game reserves, including Kruger (South Africa),
Etosha (Namibia), Lake Mutirikwe (Zimbabwe), Tsavo West (Kenya), Yankari (Nigeria).

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Genus Circellium Latreille, 1829

Type species and designation: Scarabaeus bacchus Fabricius, 1781, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Scholtz and Howden (1987).

Usually cited as Circellium Latreille, 1825 but first listing as ‘Circellie’ is in Latreille, 1829. Circellium Latreille, 1829 is
very large bodied, flightless and monotypic with a patchy distribution restricted to the S coast of South Africa. It is assessed
as Vulnerable (VU) owing to its flightless condition and high levels of disturbance over much of its range. Nevertheless,
despite patchy and mostly uncommon occurrence along the S Cape coastal regions, it occurs with high frequency in the
least disturbed, sandy, shrubland areas of Addo Elephant National Park.

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Circellium bacchus
(Fabricius, 1781)
= Scarabaeus hemisphaericus Pallas, 1781
= Circellium lyceus Westwood, 1837a
= Circellium bacchus var. waterhousei Shipp, 1895a
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus bacchus: ‘Cap bon. sp.’ [Cape of Good

Hope, South Africa]; S. hemisphaericus: Not stated; C. lyceus: ‘Sierra
Leone’ [undoubtedly an erroneous locality]; C. bacchus var. waterhou-
sei: ‘Matabili-land’ [Matabeleland, SW Zimbabwe; considered a most
doubtful locality].
Taxonomy: Accepted species.
Distribution: Centred on the S coastline, South Africa, in the winter and
bimodal rainfall regions; E and W populations are genetically distinct,
presumably, due to disjunction after Pliocene sea level rise; published
records from NE South Africa, Namibia, Zimbabwe, Mozambique,
Tanzania and Sierra Leone are considered to be errors; claims for range
contraction, linked to that of hooked-lipped (black) rhinoceros and
their dung, are not supported by food type associations.
Locality data (mean ± SD, range): Altitude: 151 ± 146, 0–535 m; annual
rainfall: 431 ± 42, 346–531 mm; annual temperature (max. + min. /2):
17.2 ± 0.9, 14.9–18.7°C (N=24).
Habitat: No quantitative assessment of soil type association; in Addo El-
ephant National Park: abundant in dense shrub/woodland on sandy
soils; most uncommon in adjacent disturbed open vegetation.
Food types: In Addo Elephant National Park: elephant dung preferred for
feeding, buffalo and cattle dung for breeding; also recorded on dung of
monkey, human, rhinoceros, hare, ostrich.
Temporal activity: Flightless, ectothermic and diurnal with maximal ac-
tivity between 18–26°C, particularly after rainfall; recorded in every 5 mm
month of the year, but low abundance in autumn and winter (Apr.
to July); female makes and rolls balls of dung for brood construction,
which is done by males in many other species.
Bioregions South Africa: E populations: Albany Thicket Biome (AT); W
populations: vegetation units in Shale Renosterveld (FRs), Limestone
Fynbos (FFl), Sandstone Fynbos (FFs), Sand Fynbos (FFd), Strandveld
(FS) (Fynbos Biome).
Assessment rationale: EOO = 11 950 km2; AOO would be smaller as
much of the mapped range falls within or borders vegetation units in
which the natural vegetation is threatened (transformation: AT: 1.5–
35%, FS: 5–50%, FR: 58–61%, FF: 17–70%); this is a cause for concern
as clearance of natural shrubland results in a radical decline in population
density whereas range fragmentation is exacerbated by low vagility due
to flightlessness; only 5% of the range is protected within natural shrub­
land reserves, mostly comprising small fragments; it has the distinction
of being the only South African dung beetle specifically protected by law
(Proclamation 24/1992 – Western and Eastern Cape Provinces).
Conservation measures: E populations are abundant in the relatively large
Addo Elephant National Park (1 640 km2) where elephant and buffalo
dung is common; however, rangers report that traffic road kill is a problem
in Addo where dung from elephants is common on roads that are used for
ease of passage through dense shrubland; even so, genetic subunits of W
populations might be more vulnerable as they are only protected in vari-
ous small nature reserves, including De Hoop, Goukamma, Salmonsdam.

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Genus Dicranocara Frolov & Scholtz, 2003b

Type species and designation: Dicranocara deschodti Frolov & Scholtz, 2003b, by original designation.
Synonyms: None.
Last review: All species reviewed by Deschodt and Scholtz (2007); one species added by Moretto
(2016).

The flightless genus, Dicranocara Frolov & Scholtz, 2003b, is entirely restricted to rocky river valleys and mountains just
to the N and S of the lower Orange River where it has been recorded close to rock hyrax middens. Although a new species
has now been described from the rocky edges of the upper Fish River, no Dicranocara species were recorded during a survey
along the lower Fish River that is bordered by sand and lacked hyrax colonies.

All species appear to occupy very small ranges where they may be supported mainly by concentrations of dung in hyrax
middens, which currently represent an assured dung supply. Although the ranges of three species lie entirely within pro-
tected areas, one of these areas has now been granted as a mining concession so that the assessment for that species has
been changed from Least Concern (LC) to Vulnerable (VU).

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Dicranocara deschodti
Frolov & Scholtz, 2003b

No synonyms
Global: VU (see IUCN Red List – LC)
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Namibia, Boom River, 27°59’S 17°03’E’.

Taxonomy: Accepted species; sexually dimorphic (head); prominence of
characters varies with body size.
Distribution: Only known from the Boom River Valley; probably endem-
ic to this usually dry, S Namibian river system that drains southwards
into the lower Orange River; nearby rocky-edged tributaries are occu- ♂
pied by other species.
Locality data (mean ± SD, range): Altitude: 769 ± 243, 339–930 m;
annual rainfall: 81 ± 19, 50–99 mm; annual temperature (max. + min.
/2): 16.5 ± 0.9, 15.9–17.9°C (N=5).
Habitat: Accumulations of sand at the bases of mountains and rocks bor-
dering the river bank near rock hyrax colonies.
Food types: Associated with piles of dung pellets contained in or near
communal hyrax middens; mouthparts modified, probably for the use
of dry food.
Temporal activity: Flightless, active in darkness; uses the clypeal projec-
tions at the front of the head to transport hyrax pellets; these are carried
across rocks and buried in the soil at the base where a single breeding
mass of dung containing an egg is constructed at the end of a short
tunnel.
Ecoregions Namibia: Near the NW tip of the Nama Karoo (AT1314).
Assessment rationale: EOO = 190 km2; despite flightless condition and
small known EOO, formerly assessed as Least Concern (LC) since its
range occurs entirely within a protected area where it would probably
not face current threats as long as a supply of dung is maintained, which 2 mm
seemed relatively assured given the widespread occurrence of the rock
hyrax in arid regions; however, the entire known range in the Boom
River Valley has now been granted as a diamond mining concession
and access has been barred in terms of the Diamond Act of 1999; as
disturbance by mining could impact on its habitat and the density of
hyrax colonies, the assessment has been revised to Vulnerable (VU).
Conservation measures: Although protected within the Ai-Aïs/Rich-
tersveld Transfrontier Park, granting of the entire known EOO as a
mining concession means that impact of mining activities will need to
be assessed at a later date.

♀ 2 mm

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Dicranocara inexpectata
Deschodt & Scholtz, 2007

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Richtersveld, Witputs RD’ [S Namibia].

Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: Possibly highly localised within an area to the N of the
source of the Boom River, the Boom River Valley lies 45 km to the S
♂ and forms the type locality of D. deschodti Frolov & Scholtz.
Locality data (average): Altitude: 867 m; annual rainfall: 99 mm; annual
temperature (max. + min. /2): 16.9°C (N=1).
Habitat: No quantitative assessment; type series recorded on a small
boulder-strewn outcrop.
Food types: No quantitative assessment; type series collected around
hyrax middens.
Temporal activity: Flightless; diel periodicity unknown, but presumably
active in darkness like D. deschodti; recorded during the late summer
rainy season (Mar., Apr.).
Ecoregions Namibia: Near the NW tip of the Nama Karoo (AT1314).
Assessment rationale: EOO unknown; probably not threatened as the
area is used primarily for grazing domestic livestock, but it would qual-
ify for an IUCN threat category if the EOO proves to be very small;
currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative sur-
vey is required to determine the EOO and AOO, also to confirm the
observed ecological associations; known only from the type locality,
which lies outside of protected areas.

2 mm

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Dicranocara tatasensis
Deschodt & Scholtz, 2007

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Tatasberg, Richtersveld Nat. Park’ [NW South Africa].

Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: Known only by the type series collected in higher parts of
the Tatasberg at the S edge of the lower Orange River Valley, South
Africa.
♂
Locality data (average): Altitude: 372 m; annual rainfall: 53 mm; annual
temperature (max. + min. /2): 17.7°C (N=1).
Habitat: No quantitative assessment; type series collected dead or alive
amongst boulders close to hyrax middens in a vegetation unit charac-
terised by sandy loam and scrub.
Food types: No quantitative assessment; probably primarily hyrax dung
pellets.
Temporal activity: Flightless; diel periodicity unknown, but presumably
active in darkness like D. deschodti Frolov & Scholtz, 2003; recorded
during the late summer rainy season (Mar., Apr.).
Bioregions South Africa: Vegetation unit: Tatasberg Mountain Succulent
Shrubland (SKr 9) (Richtersveld Bioregion, Succulent Karoo Biome).
Assessment rationale: EOO = 3.3 km2 (based on area of SKr 9); known
only from a very small upland island of Succulent Karoo vegetation;
SKr 9 is entirely isolated in the Gariep Desert Bioregion of the Desert
Biome; despite a possibly minute range it was found close to a relatively
assured food supply; also, the entire range occurs within a protected
area; therefore, currently assessed as Least Concern (LC) although it
has been described as rare and would be vulnerable to decline in hyrax
2 mm
population density.
Conservation measures: A survey is required to confirm that this species
is restricted to the Tatasberg and does not occur in the slightly moister
desert mountains to the S; if so, conservation status might require re-
vision on the basis of small range at a single location; protected in the
Richtersveld National Park (South Africa).

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Dicranocara vandersmisseni
Moretto, 2016a

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Namibie, Karas, Ai-Aïs’ [Ai-Aïs, S Namibia].

Taxonomy: Accepted species; sexually dimorphic (head), characters vary
in prominence with body size.
Distribution: Known only by the type series of three individuals collected
along those parts of the Fish River that are bordered by rocky edges,
♀ which support hyrax colonies.
Locality data (mean ± SD, range): Altitude: 449 ± 138, 351–547 m;
annual rainfall: 91 ± 44, 60–122 mm; annual temperature (max. + min.
/2): 19.6 ± 0.1, 19.5–19.7°C (N=2).
Habitat: No quantitative assessment; soil and vegetation associations un-
known.
Food types: No quantitative assessment; probably, primarily hyrax dung
pellets; one individual found dead in a hyrax midden (Aug.).
Temporal activity: Flightless; diel periodicity unknown, but possibly ac-
tive in darkness like D. deschodti Frolov & Scholtz, 2003; recorded alive
in the soil in mid-summer (Jan.); dead specimens recorded in spring
and mid-summer (Aug., Jan.).
Ecoregions Namibia: NW edge, Nama Karoo (AT1314).
Assessment rationale: EOO = 300 km2; ecologically very poorly known;
therefore, currently assessed as Data Deficient (DD); however, assumed
to be associated primarily with hyrax middens, and would therefore be
vulnerable to decline in hyrax population density.
Conservation measures: A survey is required to confirm that this species
is restricted to rocky areas of the arid Fish River Valley; protected with-
in the borders of the Ai-Aïs/Richtersveld Transfrontier Park.

2 mm

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Genus Drogo Deschodt, Davis & Scholtz, 2016

Type species and designation: Drogo stalsi Deschodt, Davis & Scholtz, 2016, by original designation.
Synonyms: None.
Last review: Genus created by Deschodt et al. (2016).

Drogo Deschodt, Davis & Scholtz, 2016, is a flightless, monotypic genus that is known only from the type locality in
mountains just N of the lower Orange River in Namibia. The genus has been recorded only in farm rangeland although
habitat disturbance is, apparently, limited.

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Drogo stalsi
Deschodt, Davis & Scholtz, 2016

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Zebrafontein, ± 30 km, NNE of Rosh Pinah’ [Farm in S

Namibia].
Taxonomy: Accepted species; known only by the male holotype.
Distribution: Known only from the type locality in arid S Namibia just
N of the Orange River.
♂ Locality data (average): Altitude: 1 608 m; annual rainfall: 123 mm;
annual temperature (max. + min. /2): 12.9°C (N=1).
Habitat: Unknown.
Food types: Unknown.
Temporal activity: Flightless, diel periodicity unknown; recorded during
the late summer rainy season (Apr.).
Ecoregions Namibia: NW edge, Nama Karoo (AT1314).
Assessment rationale: EOO unknown; known only by the male holo-
type; arid type locality unprotected, not apparently heavily disturbed
although used for the grazing of domestic livestock; unclear if D. stalsi
faces any threats; assessed as Data Deficient (DD) since so little is
known about the species.
Conservation measures: To adequately assess conservation status, a
quantitative survey is required to document the full EOO, AOO and
ecological associations; the type locality falls outside of and between the
boundaries of Sperrgebiet and Richtersveld National Parks.

2 mm

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Genus Dwesasilvasedis Deschodt & Scholtz, 2008

Type species and designation: Dwesasilvasedis medinae Deschodt & Scholtz, 2008, by original designation.
Synonyms: None.
Last review: Genus created by Deschodt and Scholtz (2008).

Dwesasilvasedis Deschodt & Scholtz, 2008, is flightless, small-bodied and monotypic; known only from leaf litter in a sin-
gle small patch of forest on the SE coast of South Africa. By virtue of its habits and small range in a single known location,
it is assessed as Vulnerable (VU).

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Dwesasilvasedis medinae
Deschodt & Scholtz, 2008

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Dwesa Forest, Eastern Cape’ [South Africa].

Taxonomy: Accepted species; sexually dimorphic (head).
Distribution: Known only from the type locality of Dwesa Forest.
Locality data (average): Altitude: 112 m; annual rainfall: 899 mm; annu-
al temperature (max. + min. /2): 18.9°C (N=1).
♂ Habitat: No quantitative assessment; all specimens in type series (19) sift-
ed from leaf litter in indigenous forest in an area dominated by sandy
loam soils; assumed to be a forest specialist.
Food types: Not known.
Temporal activity: Diel periodicity not known; recorded in the summer
rainy season (Nov.).
Bioregions South Africa: Scarp Forest (FOz 5) (Forest Biome).
Assessment rationale: EOO = 28.6 km2 (area of Dwesa Forest); almost
contiguous with adjoining Cwebe Forest (18.4 km2); Dwesa-Cwebe
Forest is an isolated patch of Transkei Coastal Scarp Forest; it is
unknown whether or not this species occurs in other much smaller
patches of FOz 5 to the S and N; although officially protected as the
Dwesa-Cwebe Wildlife Reserve, it is utilised for cutting of poles and
hunting; owing to small known range in few locations that are under
exploitation pressure D. medinae is assessed as Vulnerable (VU).
Conservation measures: Continuing protection of indigenous forest in
Dwesa-Cwebe Wildlife Reserve would be essential for the conservation
of this monotypic genus; a survey of other nearby FOz 5 Scarp For-
est patches to N and S would provide a better supported assessment
1 mm
of EOO and conservation status; vegetation specialisation and AOO
should be confirmed by a gradsect across the forest edge.

♀
1 mm

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Genus Endroedyolus Scholtz & Howden, 1987

Type species and designation: Endroedyolus paradoxus Scholtz & Howden, 1987, by original designation.
Synonyms: None.
Last review: Genus created by Scholtz and Howden (1987).

Endroedyolus Scholtz & Howden, 1987, is small-bodied, flightless and monotypic. It has been recorded from leaf litter in
a few small upland forest patches in the mistbelt of SE South Africa. Because of its flightless condition, very small known
range and threats to its only two known locations, the genus has been assessed as Endangered (EN).

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Endroedyolus paradoxus
Scholtz & Howden, 1987

No synonyms
Global: EN (See IUCN Red List – EN)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transkei, Umtata, Nquadu Mt.’ [Eastern Cape, South

Africa]
Taxonomy: Accepted species.
Distribution: Forest patches on coastal hills in the vicinity of Mthatha,
Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 1 170 ± 61, 1 126–1 213 m;
annual rainfall: 1 111 ± 136, 1 015–1 207 mm; annual temperature
(max. + min. /2): 15.6 ± 0.4, 15.3–15.8°C (N=2).
Habitat: No quantitative assessment; recorded from only two indigenous
forest localities.
Food types: No quantitative assessment; type series sifted from forest floor
leaf litter; subsequently sampled using dung baits.
Temporal activity: Flightless; diel periodicity not known; recorded during
the summer rainy season (Dec.).
Bioregions South Africa: Southern Mistbelt Forest (FOz 3) (Forest
Biome).
Assessment rationale: EOO = 80 km2; AOO = 8.9 km2; EOO and AOO
possibly underestimated as there is a group of > 20 isolated FOz 3
forest patches between Mthatha and Cala; however, this monotypic
genus is currently known only from the type locality and one other
nearby forest patch (Mhlanlane Forest Station) at the E end of these
forest patches; assessed as Endangered (EN) in the IUCN Red List;
both known localities are currently state-protected forests although
Nqadu Forest is subject to a land claim; also past fires have destroyed
1 mm
much of the habitat and commercial plantations have replaced much
of the native forest; even if future survey demonstrates a larger EOO,
probably deserves at least Vulnerable (VU) status although somewhat
Data Deficient (DD) at present.
Conservation measures: Protection of FOz 3 forest patches is essential
for the conservation of this species, particularly Nqadu and Mhlanlane;
to provide a more balanced assessment of EOO, a survey is required
in forest patches between Umtata and Cala, and those to the S and
N; a gradsect from grassland into forest patches would confirm the
vegetation specialisation and restricted AOO; it would also generate
data on population density and assist assessment of conservation status.

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Genus Epirinus Dejean, 1833

Type species and designation: Scarabaeus flagellatus Fabricius, 1775, by subsequent designation (Ferreira, 1972 – ear-
lier designations by Reiche, 1841 and Janssens 1938a ruled invalid by Daniel, 2019).
Synonyms: = Endroedyantus Cambefort, 1978; Type species: Endroedyantus silvestris Cambefort,
1978, by original designation.
Last review: All species reviewed by Scholtz and Howden (1987); new species, revised key and
morphological phylogeny (Medina & Scholtz 2005); new species and synonymies
(Deschodt et al. 2019) (see Preamble).

The long-established genus, Epirinus Dejean, 1833, comprised 29 species when the manuscript was completed at the end
of 2017. Subsequently, Deschodt et al. (2019) have described five new species that are not included in the species accounts
and have synonymised two others (see species accounts for E. hluhluwensis, E. ngomae, and Preamble). At the end of 2017,
available data indicated that the genus was endemic to South Africa, Eswatini and Lesotho, but one recent record for Epir-
inus flagellatus (Fabricius, 1775) extends the range to N Namibia (Deschodt et al. 2019). Six species have been observed
to roll balls of dung despite small body size.

As there is some conflict between relationships defined by morphological (Medina & Scholtz 2005) and partial molecu-
lar phylogenies (Mlambo et al. 2015), it is uncertain how to define centres of distribution. However, the morphological
phylogeny suggests a complex history of radiation and speciation along four main lineages with the most basally derived
lineages containing species centred more to the SW and the most derived containing some species centred more to the NE.
Species centred on cooler SW regions are primarily active during the winter rainy season from autumn to spring. Those
centred on cooler uplands or forest patches to the NE are primarily active during the summer rainy season.

Flightless species (10, reduced to 8) are primarily centred on coastal and mountain forest patches along the S and E
seaboard. Phylogenies variously divide them into two lineages centred to the SW or NE. All but one species show small
EOOs from 81 to 2 300 km2. Distributions are centred on the S Cape (3), E Cape (4) or NE KwaZulu-Natal (3 – now
synonymised as a single species, E. davisi Scholtz & Howden, 1987).

Species centred primarily on moister E Highveld grasslands and S Cape fynbos (8) occur in two different lineages. Most
show large EOOs although known ranges of a few are very small (80–170 km2). Distributions are centred on NE Highveld
(4, one in forest), E Highveld (1), NE Highveld to S Cape (2), S Cape (1).

The remaining 11 species show SW centres of distribution, but occur in three different lineages. Most show large EOOs
other than those (4) on the SW coast and mountains (15–3 400 km2). Distributions are centred on W (2) or SW (2)
coastal sands, SW mountains (1), SW fynbos (1), SE shrubland and Highveld Grassland (1), W Highveld Grassland (1),
Upper Karoo (1) or they are widespread primarily in SW fynbos and Karoo (2).

On the basis of widespread occurrence 11 species are assessed as Least Concern (LC) with the remainder assessed as Data
Deficient (DD: 14) or Vulnerable (VU: 4). Many of the known distributions are relatively small, particularly those found
in forest. Many are centred on or overlap into regions of high habitat transformation. It is probable that many of those
assessed as DD would qualify for an IUCN threat category. Severely limited available data suggest that clearance of indig-
enous SW shrubland is detrimental to population density in three species. Disturbance in Highveld Grassland may also
reduce population density. Only small parts of the ranges of most species are under protection.

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Epirinus aeneus
(Wiedemann, 1823)

= Epirhinus deplanatus Boheman, 1858

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Canthon aeneus: ‘Prom. bon. sp.’; E. deplanatus: ‘Prom-

ontorium Bonae Spei’ [both, Cape of Good Hope, South Africa].
Taxonomy: Accepted species.
Distribution: Widespread in the Nama Karoo and the winter and bimod-
al rainfall regions of South Africa comprising Fynbos and Succulent
Karoo.
Locality data (mean ± SD, range): Altitude: 652 ± 466, 0–1 614 m; an-
nual rainfall: 231 ± 94, 66–539 mm; annual temperature (max. + min.
/2): 17.6 ± 1.6, 12.4–21.7°C (N=125).
Habitat: In Western Cape: abundant in shrubland on deep sand (Modder-
rivier: 306; Pampoenvlei: 353) and pasture grassland cleared of shrubs
on sandy loam (Oranjefontein: 159; Waylands: 258); on deep sand in
West Coast National Park: less abundant in natural shrubland (320)
than in sparse-grass pasture (607) resulting from shrubland clearance;
DBRU collection records primarily from coarse-grained soils: sand
(28), sandy loam (3), sandy clay loam (4) in scrub/shrubland (18), also
pasture/grassland (7), fallow crop fields (6).
Food types: No quantitative data; DBRU collection records from cattle
(31), buffalo (1), horse (4), sheep (1) dung.
Temporal activity: Diurnal flight; probably in sunshine as long as
temperatures are suitable; in winter rainfall region of Western Cape:
autumn, winter, spring activity by adults, sclerotised (Apr. to Oct.),
teneral (Sept. to Nov.).
Bioregions South Africa: Known from all three bioregions of the Nama
Karoo Biome; also S Namaqualand Sandveld (SKs), Namaqualand
Hardeveld (SKn) and Rainshadow Valley Karoo (SKv) (Succulent
Karoo Biome); as well as units of Fynbos (FF), Strandveld (FS) and
Renosterveld (FR) (Fynbos Biome).
2 mm
Assessment rationale: EOO = 335 500 km2; widespread in both highly
transformed regions in the SW Western Cape (25–80%; FF, FS, FR)
and little-transformed regions used primarily for grazing of livestock
in Namaqualand and the Nama Karoo; abundance in pastures cleared
of natural shrubland and records from fallow crop fields suggest that
habitat transformation is less detrimental to E. aeneus than to many
other species with endemic SW distribution patterns; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tected in Bontebok and West Coast national parks.

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Epirinus aquilus
Medina & Scholtz, 2005

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Grahamstown’ [Eastern Cape, South Africa].

Taxonomy: Accepted species.
Distribution: Known only from the isolated coastal Alexandria State For-
est and one other, nearby, hillside forest patch (holotype locality), close
to Grahamstown, Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 402 ± 394, 123–681 m;
annual rainfall: 539 ± 80, 482–595 mm; annual temperature (max. +
min. /2): 16.8 ± 1.3, 15.9–17.7°C (N=2).
Habitat: No quantitative assessment; recorded only in forest established
on sandy soils; the coastal Alexandria Forest borders the Alexandria
dune field.
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Diurnal, flightless, primarily active in the summer
rainy season (Jan. to May).
Bioregions South Africa: Recorded from two vegetation units: Southern
Coastal Forest (FOz 6) (Forest Biome); also a forest patch in Suurberg
Quartzite Fynbos (FFq 6) (Fynbos Biome).
Assessment rationale: EOO = 130 km2; flightless condition and severely
limited range at only two locations would qualify this species for an
Endangered (EN) threat category; however, it is currently assessed as
Vulnerable (VU) since, (1) the EOO may be underestimated, and, (2)
it is protected in Alexandria Forest, which forms part of the extended
Addo Elephant National Park; nevertheless, re-assessment may be re-
quired as heavy exploitation has been reported at the forest periphery,
which extends onto private land.
Conservation measures: Assessment of conservation status would be
improved by a quantitative survey of forest patches and adjoining ter-
rain in the Alexandria and Grahamstown areas; this would confirm the
1 mm
EOO, AOO and ecological habits; continuing protection of Alexandria
Forest (Addo Elephant National Park) would be be essential for conser-
vation of this species.

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Epirinus asper
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Natal (Durban, Ladysmith), Transvaal (Lydenburg)’

[Durban, Ladysmith (KwaZulu-Natal), Mashishing (Mpumalanga),
South Africa], lectotype: ‘Durban, Natal’.
Taxonomy: Accepted species.
Distribution: Disjunct pattern on moist highlands along the edge of the E
escarpment: NE Lesotho border and Mpumalanga Province, South Af-
rica; absence of records from intervening highlands may be a collection
artefact; inexact Durban type locality (black square) may lie in coastal
hills, not recently recorded from Durban area.
Locality data (mean ± SD, range): Altitude: 1 763 ± 541, 1 073–3 252 m;
annual rainfall: 854 ± 131, 706–1 157 mm; annual temperature (max.
+ min. /2): 13.7 ± 3.3, 4.6–17.5°C (N=18) (inexact Durban lectotype
locality, not included).
Habitat: No quantitative assessment; DBRU collection records from
finer-grained soils: sandy clay loam (1), sandy loam (4) in grassland/
pasture (5).
Food types: No quantitative assessment; DBRU collection records from
cattle dung (4).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: NE edges of Drakensberg Grassland (Gd) and
Mesic Highveld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 40 500 km2 (E escarpment range only);
fairly widespread range in an area where natural habitat has been partly
transformed (Gd: 3–16%; Gm: about 23% in occupied vegetation
units); there are no quantitative data to assess possible detrimental
effects of pasture improvement and development of tree plantations;
currently assessed as Least Concern (LC) on basis of large range.
2 mm Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, including
a survey of natural grassland patches along the NE escarpment edge,
coastal hills around the Durban type locality, and intervening localities;
possible effects of pasture modification should also be tested; protected
in uKhahlamba Drakensberg Park (World Heritage Site).

CANTHONINI
SURICATA 6 (2020) 89

Epirinus bentoi
Ferreira, 1964a

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘África do Sul: Província do Cabo (Eland’s Bay,...)’ [Elands

Bay, Western Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Restricted to the West Coast, Western Cape, in the winter
rainfall region of South Africa.
Locality data (mean ± SD, range): Altitude: 91 ± 63, 15–199 m; annual
rainfall: 226 ± 68, 157–350 mm; annual temperature (max. + min. /2):
17.3 ± 1.2, 15.7–18.9°C (N=12).
Habitat: No quantitative assessment of soil type; on Farm Geelbek: strong
bias to natural shrubland (71) as opposed to sparse pasture (17) result-
ing from shrubland clearance; farm now restored to shrubland after
incorporation into West Coast National Park; all DBRU collection
records on deep coastal sands (4) in natural scrub/shrubland (4);
Food types: No quantitative assessment; DBRU collection records on
cattle dung (4).
Temporal activity: Diurnal flight activity in spring to early summer (Aug.
to Dec.); sclerotised adults (Aug. to Oct.), teneral adults (Oct. to Dec.).
Bioregions South Africa: Known primarily from strandveld vegetation
units: Namaqualand Strandveld (SKs 7) (Succulent Karoo Biome);
Lamberts Bay Strandveld (FS 1), Langebaan Dune Strandveld (FS 5)
(Fynbos Biome); also Atlantis Sand Fynbos (FFd 4).
Assessment rationale: EOO = 2 700 km2; localised within a relatively
small range in an area that is subject to high transformation pressures
through mining, cultivation and urbanisation (SKs 7: 10%, FS: 25-
35%, FFd 4: 40%); because of small range, sensitivity to habitat trans-
formation, and limited protection, this species is assessed as Vulnerable 2 mm
(VU).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tection of natural shrubland and continuing protection of reserves are
recommended for the conservation of this species; S parts of range pro-
tected in West Coast National Park and the private Grotto Bay Nature
Reserve.

CANTHONINI
90 SURICATA 6 (2020)

Epirinus comosus
Péringuey, 1901

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Cape Colony (Stellenbosch)’ [Western Cape, South Af-

rica], but later cited as the nearby ‘Strand’ on the basis of a specimen
bearing a type label in Péringuey’s handwriting.
Taxonomy: Accepted species.
Distribution: Mountain and coastal localities of the Western and East-
ern Cape in the winter and bimodal (spring/autumn) rainfall regions,
South Africa; disjunct pattern may be a collection artefact.
Locality data (mean ± SD, range): Altitude: 381 ± 536, 0–1 885 m;
annual rainfall: 375 ± 85, 186–475 mm; annual temperature (max. +
min. /2): 16.0 ± 2.2, 10.1–17.7°C (N=14).
Habitat: No adequate quantitative assessment; in Cape of Good Hope
Nature Reserve: sampled uncommonly (6) on deep sand at a locality
burnt 9–10 yr previously comprising open fynbos; absent (0) from
nearby localities, one burnt 2–3 yr previously comprising a dense cover
of Restio spp., another comprising Kikuyu pasture; one DBRU collec-
tion record from deep sand in sparse pasture.
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Diurnal flight activity in late winter to early summer
(July to Dec.) and autumn (Apr.) coincidental with the winter, spring
and autumn rainfall peaks in the winter and bimodal rainfall regions.
2 mm
Bioregions South Africa: Vegetation Units in Sand Fynbos (FFd 5, FFd
10), South Strandveld (FS 7), Sandstone Fynbos (FFs 4, FFs 9, FFs 19),
Granite Fynbos (FFg 3) (all Fynbos Biome); two marginal records in
Shale Renosterveld (FRs).
Assessment rationale: EOO = 27 500 km2; widespread, but AOO pri-
marily in vegetation units characterised by sandy soils and open natural
shrubland; transformation in occupied sandy vegetation units varies
from 5 to 80%; may be Near Threatened (NT) in parts of its range, but
currently assessed as Data Deficient (DD).
Conservation measures: A quantitative survey of the EOO, AOO and
ecological associations is required to adequately assess the conservation
status of E. comosus, particularly soil associations and effects of vegeta-
tion transformation; protected in Cape of Good Hope Nature Reserve.

CANTHONINI
SURICATA 6 (2020) 91

Epirinus convexus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transkei, The Haven’ [Eastern Cape, South Africa].

Taxonomy: Accepted species.
Distribution: E Eastern Cape, South Africa: coastal forest patches plus
one record from forest on the S escarpment.
Locality data (mean ± SD, range): Altitude: 225 ± 319, 0–863 m; annual
rainfall: 869 ± 165, 537–899 mm; annual temperature (max. + min.
/2): 18.8 ± 1.2, 16.4–19.7°C (N=6).
Habitat: No quantitative assessment; all locality records from forest (8).
Food types: Unknown.
Temporal activity: Diel periodicity unknown; presumably diurnal like
other Epirinus species; recorded sporadically in both summer rainy
season (Dec., Feb.) and cool dry season (July).
Bioregions South Africa: Primarily S coastal limits of Scarp Forest (FOz
5) and S inland limits of Southern Mistbelt Forest (FOz 3).
Assessment rationale: EOO = 2 300 km2; EOO < 5 000 km2 with validat-
ed AOO limited to two clusters of small coastal forest patches, which,
although protected, qualify this species for a threat status category,
perhaps Vulnerable (VU); however, must currently be assessed as Data
Deficient (DD) since precise data on range and population density are
lacking; furthermore, inland forest occurrence requires validation.
Conservation measures: To assess the conservation status of this species a
quantitative survey needs to be conducted in and around forest patches
from E coastal Scarp Forest to Southern Mistbelt Forest; this survey
should also assess ecological associations; currently protected in several 1 mm
coastal forest reserves, including Dwesa, Silaka, Ntsubane.

CANTHONINI
92 SURICATA 6 (2020)

Epirinus davisi
Scholtz & Howden, 1987

No synonyms (but see Taxonomy)

Global: DD
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type locality: ‘Dhlinza Forest, Eshowe Dist.’ [KwaZulu-Natal, South

Africa].
Taxonomy: Accepted species, recently re-evaluated (Deschodt et al. 2019)
as the senior name for the junior synonyms, E. hluhluwensis (also Scarp
Forest) and E. ngomae (Southern Mistbelt Forest) (both Medina &
Scholtz 2005) (see Preamble).
Distribution: Isolated patches of forest on the lower edges of the coastal
escarpment, KwaZulu-Natal, South Africa; described from Dhlinza
and oNgoye forests (see map above), but now includes records for ju-
nior synonyms, E. hluhluwensis and E. ngomae, plus new records from
Eswatini and Nkandla Forest, towards the upper edges of the coastal
escarpment (Deschodt et al. 2019).
Locality data (mean ± SD, range): Dhlinza and oNgoye forests; altitude:
258 ± 71, 208–308 m; annual rainfall: 988 ± 11, 980–995 mm; annual
temperature (max. + min. /2): 20.6 ± 0.3, 20.4–20.8°C (N=2).
Habitat: All records from forest characterised by sandy loam soils; record-
ed from humus [forest litter].
Food types: Not known (but see data for E. hluhluwensis and E. ngomae).
Temporal activity: Flightless; diel perodicity unknown; recorded in the
summer rainy season (Oct. to Dec.) in Dhlinza and oNgoye forests;
data for E. ngomae extends seasonal extent to March.
2 mm
Bioregions South Africa: Scarp Forest (FOz 5) (Forest Biome), but see
data for E. ngomae.
Assessment rationale: EOO = 81 km2; AOO = 30 km2 (estimated com-
bined area of Dhlinza and oNgoye) although addition of synonyms
much increases the EOO to 11 700 km2; known AOO would remain
sufficiently limited to qualify for a threat category of Vulnerable (VU),
but currently so poorly known, that it must be assessed as Data Defi-
cient (DD); not recently recorded in either Dhlinza or oNgoye (Dec.
2014).
Conservation measures: Continuing protection of Dhlinza and oNgoye
forest reserves (plus other forest patches occupied by new synonyms) is
essential for the conservation of this flightless taxon; improved assess-
ment of conservation status also requires a survey of similar regional
forest patches plus their environs as well as quantitative data on the
species’ ecological habits.

CANTHONINI
SURICATA 6 (2020) 93

Epirinus drakomontanus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: Lesotho

J A S O N D J F M A M J

Type locality: ‘Lesotho, Drakensberg (Tseke Tseke Gorge)’.

Taxonomy: Accepted species.
Distribution: Known only from extreme high altitude along the upper
edge of the E escarpment just within the NE border of Lesotho.
Locality data (mean ± SD, range): Altitude: 2731 ± 473, 2 274–3 218 m;
annual rainfall: 877 ± 125, 751–1 001 mm; annual temperature (max.
+ min. /2): 7.8 ± 2.9, 4.8–10.6°C (N=3).
Habitat: No quantitative assessment; sampled in low numbers (2) from
sandy clay loam in short grassland with scattered shrubs.
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Diurnal flight activity in the summer rainy season
(Jan. to Mar.).
Bioregions South Africa: Vegetation units: Northern Drakensberg High-
land Grassland (Gd 5), Lesotho Highland Basalt Grassland (Gd 8)
(Drakensberg Grassland, Grassland Biome).
Assessment rationale: EOO = 80 km2 (probably underestimated); small
known range limited to wet escarpment edge, but localities in Gd 8
suggest possible wider occurrence in the poorly protected Lesotho
Highlands, which are subject to habitat transformation by cultivation
and overgrazing (Gd 5, Gd 8: 7–10%); assessed as Data Deficient
(DD) since it is poorly represented in collections and poorly known.
Conservation measures: A quantitative survey is required to ascertain
the full EOO and ecological associations before an assessment may be
made of its conservation status; the survey should determine if it is
protected within the upper margins of uKhahlamba Drakensberg Park
(World Heritage Site) (South Africa) or whether its range lies entirely
2 mm
outside of the reserve within Lesotho.

CANTHONINI
94 SURICATA 6 (2020)

Epirinus flagellatus
(Fabricius, 1775)

= Scarabaeus granulatus Olivier, 1789

= Scarabaeus scabratus Fabricius, 1794
= Ateuchus callosus Thunberg, 1818
= Epirinus vicinus Ferreira, 1964a
Global: LC
Endemic: RSA, Lesotho, Namibia

J A S O N D J F M A M J

Type localities: As Scarabaeus flagellatus: ‘Cap B. S.’ [Cape of Good Hope,

South Africa]; S. granulatus: Not stated; S. scabratus: ‘Cap Bon. Spei’;
A. callosus: Not stated; E. vicinus: ‘África do Sul: Província do Cabo
(Willowmore,...’ [Willowmore, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Widespread in drier parts of the winter and bimodal rainfall
regions, South Africa; scattered, high altitude occurrence to the NE in
the summer rainfall region, particularly Lesotho highlands. After com-
pletion of this book at the end of 2017, this species has been recorded
from Okonjima Nature Reserve, representing a first record for Epirinus
from Namibia (Deschodt et al. 2019; see Preamble).
Locality data (mean ± SD, range): Altitude: 540 ± 557, 0–2 264 m;
annual rainfall: 302 ± 164, 79–955 mm; annual temperature (max. +
min. /2): 16.1 ± 1.9, 9.6–19.3°C (N=110).
Habitat: No adequate quantitative assessment; abundant in SW coastal
samples: Farm Pampoenvlei shrubland on sand (159) or sandy loam
(62); Modderrivier shrubland on sand (206), Groote Post pasture on
sandy loam (133); DBRU collection records: clay (1), sandy clay loam
(1), sandy loam (10), sand (5) in grassland/pasture (11) or Karoo scrub/
shrubs (4); also fallow crop fields (2).
Food types: No quantitative assessment; DBRU collection records on
cattle (18), horse (1), human (1) dung.
Temporal activity: Diurnal flight activity throughout year, but in win-
ter rainfall region, mainly autumn to early summer seasonal activity
by adults, sclerotised (mainly Mar. to Oct., peaking Mar. to May and
Aug.), tenerals (Oct. to Nov.).
Bioregions South Africa: Centred on various drier bioregions of the
2 mm
Succulent Karoo (SK), Fynbos (F) and Albany Thicket (AT) biomes;
scattered outlier highland occurrence in the Grassland Biome.
Assessment rationale: EOO = 258 000 km2; adaptable habits indicated
by widespread range across winter, bimodal and summer rainfall cli-
mates on sand and sandy loam in both shrubland and grassland cleared
of natural shrubland; little transformation over the arid NW part of its
range; S part of range with variable degrees of transformation; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tected in Namaqua and Bontebok national parks; also Cape of Good
Hope Nature Reserve.

CANTHONINI
SURICATA 6 (2020) 95

Epirinus granulatus
Scholtz & Howden, 1987

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Namaqualand, Koekenaap’ [South Africa].

Taxonomy: Accepted species.
Distribution: Restricted to the West Coast, Western Cape and Northern
Cape, in the winter rainfall region of South Africa.
Locality data (mean ± SD, range): Altitude: 130 ± 73, 15–279 m; annual
rainfall: 216 ± 90, 157–286 mm; annual temperature (max. + min. /2):
17.3 ± 1.1, 15.7–18.5°C (N=11).
Habitat: No quantitative assessment; in SW Cape: sampled from natural
shrubland on inland sand dunes (37) on Farm Pampoenvlei, not re-
corded nearby in cleared pastures (Farm Groote Post) or nearby towards
the cooler coastline (Farm Modderrivier).
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Diurnal flight activity during autumn and spring, scle-
rotised adults (mainly May to July), teneral (mainly Oct.).
Bioregions South Africa: Known from few vegetation units primarily
in strandveld; Namaqualand Strandveld (SKs 7) (Succulent Karoo
Biome); Lamberts Bay Strandveld (FS 1), Langebaan Dune Strandveld
(FS 5) (Fynbos Biome); also Atlantis Sand Fynbos (FFd 4).
Assessment rationale: EOO = 3 400 km2; localised within a relatively
small range in an area that is subject to transformation pressures through
mining, cultivation and urbanisation (SKs 7: 10%, FS: 25–35%, FFd
4: 40%); because of small range and AOO, possible sensitivity to trans-
formation with limited protection, assessed as Vulnerable (VU).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
effects of natural shrubland clearance; protection of existing reserves
probably essential for the conservation of this species; parts of S range
2 mm
protected in West Coast National Park.

CANTHONINI
96 SURICATA 6 (2020)

Epirinus gratus
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Colony (Griqualand West)’ [Northern Cape, South

Africa].
Taxonomy: Accepted species.
Distribution: Centred on the dry NW Highveld, South Africa.
Locality data (mean ± SD, range): Altitude: 1 379 ± 150, 1 176–1 824 m;
annual rainfall: 542 ± 121, 324–733 mm; annual temperature (max. +
min. /2): 17.2 ± 1.0, 14.8–19.0°C (N=23).
Habitat: No adequate quantitative assessment; in Suikerbosrand Nature
Reserve: sampled on sandy clay loam in grassland (5) or open wood-
land (1) at 1 631–1 639 m; DBRU collection records from sand (1),
sandy loam (1) in grassland (2).
Food types: No quantitative assessment; DBRU collection records from
cattle dung (2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Nov. to May); a preponderance of Feb. collection records may indicate
a late summer peak in activity.
Bioregions South Africa: Eastern Kalahari Bushveld (SVk) (Savanna
Biome); N Dry Highveld Grassland (Gh) (Grassland Biome).
Assessment rationale: EOO = 74 300 km2; widespread, but range largely
coincides with vulnerable and endangered vegetation units mainly due
to cultivation and invasive shrubs (mesquite in the W); extensive trans-
formation in occupied vegetation units in both the E (Gh: 25–63%)
and the W (SVk: 18–42%); however, assessed as Least Concern (LC)
due to large range and protection in a number of reserves.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations, particu-
2 mm larly possible effects of transformation of natural grasslands; protection
offered in nature reserves over only a small part of its range, including
Suikerbosrand, Abe Bailey, Sandveld and Nascott.

CANTHONINI
SURICATA 6 (2020) 97

Epirinus hilaris
Péringuey, 1901

= Endroedyantus youngae Cambefort, 1978

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Cape Colony (Cape Town, Stellenbosch)’ [South Africa],

lectotype: ‘Cape Town’; E. youngae: ‘Hawequas / 33.34S-19.08E’ [all
Western Cape, South Africa]
Taxonomy: Accepted species, but outlier records from higher altitude
should be validated after further comparison with E. montanus Scholtz
& Howden, 1987.
Distribution: Primarily in low-altitude localities along the S coast of
South Africa with a few records from higher altitude; distribution ex-
tends across two climatic regions, winter and bimodal rainfall to edge
of summer rainfall in the E; records from three disjunct centres may
variously be a collecting artefact, or reflect unfavourable South Coast
Renosterveld habitat (FO 8), or extensive habitat transformation.
Locality data (mean ± SD, range): Altitude: 609 ± 491, 16–1 993 m;
annual rainfall: 451 ± 63, 355–546 mm; annual temperature (max. +
min. /2): 15.3 ± 2.3, 9.6–18.3°C (N=17).
Habitat: No quantitative assessment; known primarily from fynbos shrub­
land localities with a few in forest.
Food types: No quantitative assessment; recorded from jackal dung.
Temporal activity: Diel periodicity unknown; presumably diurnal like
other Epirinus species; recorded during spring (Aug.) to early summer
(Nov. to Jan.).
Bioregions South Africa: Centred on Sandstone Fynbos vegetation units
(FFs 9, 10, 11, 13, 19) in Southwest Fynbos (FO 2) and Eastern Fynbos
(FO 6) bioregions (Fynbos Biome); also Albany Thicket Biome (AT).
Assessment rationale: EOO = 29 550 km2; ecologically poorly known;
1 mm
AOO may be much smaller if influenced by habitat transformation,
which is sometimes extensive, sometimes limited across range (2–75%);
transformation may account for disjunct records; however, assessed as
Least Concern (LC) on basis of widespread EOO.
Conservation measures: Assessment of conservation status would be
improved by quantitative ecological data, particularly ecological asso-
ciations and possible effects of clearance of natural shrubland; a survey
is also required to determine the full EOO and AOO; probably not
seriously threatened, but limited known protection in a few nature
reserves including Hottentots Holland.

CANTHONINI
98 SURICATA 6 (2020)

Epirinus hluhluwensis
Medina & Scholtz, 2005 (but see Taxonomy)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Zululand, Hluhluwe Game Res.’ [Hluhluwe–iMfolozi

Game Reserve, KwaZulu-Natal, South Africa].
Taxonomy: Accepted species when the book was completed at the end of
2017, but, together with E. ngomae Medina & Scholtz, 2005, now a ju-
nior synonym of E. davisi Scholtz & Howden, 1987 (also Scarp Forest).
Distribution: Known from Hluhluwe–iMfolozi Game Reserve.
Locality data (average): Hluhluwe Game Reserve, altitude: 273 m; av-
erage rainfall: 900 mm; annual temperature (max. + min. /2): 20.5°C
(N=1).
Habitat: Assumed to be forest on finer-grained soils.
Food types: Sampled to faeces bait.
Temporal activity: Not known; assumed to be flightless and diurnal like
close relatives; recorded in the summer rainy season (Nov.).
Bioregions South Africa: Scarp Forest (FOz 5) (Forest Biome).
Assessment rationale: EOO = 1 580 km2 (estimated from area occu-
pied by an isolated scatter of FOz 5 forest patches within and close to
Hluhluwe–iMfolozi Game Reserve); AOO = 50 km2 (estimated com-
bined area of actual forest patches); not known if it occupies all FOz 5
patches in area; although these number >10, small range might justify
a Vulnerable (VU) category; however, poorly known, so currently as-
sessed as Data Deficient (DD).
Conservation measures: Continuing protection of the scatter of Scarp
Forest patches isolated in Hluhluwe–iMfolozi Game Reserve may be
essential for the local conservation of this species, now synonymised
2 mm with E. davisi by Deschodt et al. (2019); a survey of each patch in the
area would be useful to accurately determine the AOO and ecological
associations.

CANTHONINI
SURICATA 6 (2020) 99

Epirinus minimus
Medina & Scholtz, 2005

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Prov. Alexandria For. St.’ [Eastern Cape, South Af-
rica].
Taxonomy: Accepted species.
Distribution: SE coastal area of Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 61 ± 87, 0–123 m; annual
rainfall: 462 ± 29, 441–482 mm; annual temperature (max. + min. /2):
17.4 ± 0.4, 17.1–17.7°C (N=2).
Habitat: No quantitative assessment; 47 individuals in type series sifted
from indigenous forest litter; three individuals from thicket at Cape St
Francis appear to be the same species.
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Flightless; diel periodicity assumed to be diurnal like
other Epirinus spp.; recorded in mid-summer (Dec.).
Bioregions South Africa: Southern Coastal Forest (FOz 6) (Forest
Biome), Algoa Dune Strandveld (AZs 1) (Azonal Coastal Vegetation).
Assessment rationale: EOO = 1 130 km2; poorly known, but AOO
may be even smaller if soil and vegetation specialisation is supported
by quantitative data; would qualify for IUCN threat category status
of Endangered (EN) on basis of range < 5 000 km2 at < 5 localities,
but, owing to a measure of statutory protection, currently assessed as
Vulnerable (VU).
Conservation measures: As soil and vegetation associations may influence
the AOO, quantitative data on ecological associations are required to
improve the assessment of conservation status; to determine the full
1 mm
EOO, it is also necessary to conduct a survey of forest patches and
dense fynbos thicket along the SE coastline, particularly on deep coast-
al sands; Alexandria Forest is protected in the Greater Addo Elephant
National Park.

CANTHONINI
100 SURICATA 6 (2020)

Epirinus montanus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Swartberge, Blesberg-W, 1 820 m’ [mountain in Eastern

Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Restricted known EOO at higher altitude on low rainfall
highland blocks of the Swartberg; lays to NE of coastal forest range of
close relative, E. silvestris; one cited grid reference for Swartberg Pass
(see black square) requires validation.
Locality data (mean ± SD, range): Altitude: 1 442 ± 778, 892–1 993 m;
annual rainfall: 331 ± 120, 246–415 mm; annual temperature (max.
+ min. /2): 12.5 ± 4.1, 9.6–15.4°C (N=2) (does not include southerly
record – black square).
Habitat: No quantitative assessment; two most accurate records coincide
with areas of highland fynbos shrubland on sandy soils.
Food types: No quantitative assessment; unknown.
Temporal activity: Diel periodicity unknown; presumably diurnal like
other Epirinus species; recorded during early summer (Nov., Dec.).
Bioregions South Africa: N records: North Swartberg Sandstone Fynbos
(FFs 23) vegetation unit (Fynbos Biome).
Assessment rationale: EOO = 1 750 km2 (includes non-validated south-
erly record); poorly known species with a very small known range
across a few isolated highland blocks in the Swartberg; would qualify
for IUCN threat category status of Endangered (EN) on basis of range
< 5 000 km2 at < 5 localities, but, on basis of limited information and
possible under-collection, assessed as Data Deficient (DD).
Conservation measures: To adequately assess the full EOO and conser-
vation status, a survey is required to the E and W of known localities
along the higher-lying areas of the Swartberg in FFs 23, which stretches
for 270 km along the mountain range; 70% of this vegetation unit is
conserved; therefore, survey may result in re-assessment as Least Con-
cern (LC).

2 mm

CANTHONINI
SURICATA 6 (2020) 101

Epirinus mucrodentatus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Mount Sheba, E. Tvl’ [Mpumalanga Province, South Af-

rica].
Taxonomy: Accepted species.
Distribution: Known only from a small, moist, high altitude area close to
the edge of the E escarpment in Mpumalanga, South Africa.
Locality data (mean ± SD, range): Altitude: 1 534 ± 239, 1 338–1 800 m;
annual rainfall: 885 ± 78, 803–958 mm; annual temperature (max. +
min. /2): 15.3 ± 1.5, 13.7–16.5°C (N=3).
Habitat: Uncertain; one Ditsong Museum record from high altitude
grassland; type series known to have been recorded in a survey of mist-
belt grassland and forest on Mt Sheba, but precise records not available.
Food types: A single individual sampled to carrion (Ditsong Museum);
but, probably, primarily a dung-frequenting species as the type series
was sampled to pig or cattle dung (precise records unavailable).
Temporal activity: Flight periodicity unknown, but assumed to be diurnal
like other Epirinus species; recorded in the summer rainy season (Jan.).
Bioregions South Africa: Known only from the NE element of a single
highland vegetation unit: Lydenburg Montane Grassland (Gm 18)
(Mesic Highveld Grassland, Grassland Biome) that contains patches of
Northern Mistbelt Forest (FOz 4) (Forest Biome).
Assessment rationale: EOO = 170 km2 (presumably underestimated); size
of AOO dependent on habitat association; EOO encompasses small
forest patches in a grassland matrix where transformation is 23% (Gm
18), primarily due to alien vegetation; only 2.4% of the area is formal-
ly protected; would qualify for an IUCN threat category on basis of
small range, but currently poorly known and assessed as Data Deficient
(DD).
Conservation measures: A survey is required to determine the EOO, 2 mm

AOO and ecological associations before an assessment may be made

of conservation status; if it is a grassland specialist, the survey should
determine if the range extends to the SW element of Gm 18 and if
population density is influenced by grassland modification; known to
have been relatively abundant at the type locality in the private Mt
Sheba Nature Reserve in 1974; current status uncertain.

CANTHONINI
102 SURICATA 6 (2020)

Epirinus ngomae
Medina & Scholtz, 2005 (but see Taxonomy)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘KZN Ngome State Forest’ [KwaZulu-Natal, South Africa].

Taxonomy: Accepted species when the book was completed at the end of
2017, but together with E. hluhluwensis Medina & Scholtz, 2005 now
a junior synonym of E. davisi Scholtz & Howden, 1987.
Distribution: Known from Ngome Forest on the upper edge of the coast-
al escarpment, N KwaZulu-Natal, South Africa.
Locality data (mean ± SD, range): Ngome Forest, altitude: 999 m; an-
nual rainfall: 858 mm; annual temperature (max. + min. /2): 17.6°C
(N=1).
Habitat: All records from forest characterised by finer-grained sandy clay
loam soils.
Food types: Poorly known; sampled using pitfalls either baited with pig
dung or unbaited.
Temporal activity: Flightless; diel periodicity unknown, but presumably
diurnal like other Epirinus species, recorded in the summer rainy season
(Nov. to Mar.).
Bioregions South Africa: Southern Mistbelt Forest (FOz 3) (Forest
Biome).
Assessment rationale: EOO = 765 km2 (estimated from area occupied by
an isolated scatter of forest patches at the northernmost extent of FOz
3); AOO = 40 km2 (estimated combined area of actual forest patches
2 mm with Ngome Forest by far the largest proportion, ca. 85%); not known
if it occupies all FOz 3 patches in the area; as these number < 10 and
represent a very small range, a Vulnerable (VU) category would be jus-
tified; however, currently assessed as Data Deficient (DD).
Conservation measures: Continuing protection of Ngome Forest Reserve
would be essential for the local conservation of this flightless taxon now
synonymised with E. davisi by Deschodt et al. (2019); to assess conser-
vation status, a study of ecological habits is required as well as a survey
of similar regional forest patches to determine population density in
its AOO.

CANTHONINI
SURICATA 6 (2020) 103

Epirinus obtusus
Boheman, 1857

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: Misspelt as Epirhinus obtusus: ‘Portum Elisabeth’ [Port Eliz-

abeth, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Somewhat disjunct pattern in dry to moist SE Highveld
and mostly drier parts of transitional SE coastal regions, South Africa;
disjunction may or may not be a collection artefact.
Locality data (mean ± SD, range): Altitude: 1 039 ± 513, 112–1 984 m;
annual rainfall: 494 ± 147, 277–808 mm; annual temperature (max. +
min. /2): 15.6 ± 1.9, 11.1–18.5°C (N=26).
Habitat: No adequate quantitative assessment; across a 500–2 800 m
gradsect at 30°S: recorded only at 1 500 m (49 on sandy clay loam in
natural grassland); DBRU collection records on sand (2), sandy loam
(3), sandy clay loam (4) in pasture/grassland (7), scrub/shrubland (2),
open woodland (1).
Food types: No quantitative assessment; DBRU collection records on
dung of buffalo (3), cattle (12), zebra (1), elephant (1).
Temporal activity: Diurnal flight activity, primarily in the summer rainy
season (Oct. to May).
Bioregions South Africa: In the NE: S Dry Highveld Grassland (Gh),
Sub-escarpment Grassland (Gs) (Grassland Biome); in the SW: centred
on Albany Thicket Biome (AT) extending into margins of Succulent
Karoo (SK), Nama Karoo (NK) and Savanna (SV) Biomes.
Assessment rationale: EOO = 106 900 km2; poorly known; may be a
widespread soil and open vegetation generalist; on this assumption,
assessed as Least Concern (LC), although approaching Data Deficient
(DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, including 2 mm
possible effects of habitat transformation; protected in Addo Elephant
National Park.

CANTHONINI
104 SURICATA 6 (2020)

Epirinus pseudorugosus
Medina & Scholtz, 2005

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘West Coast N. Park’ [West Coast National Park, Western
Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Currently known primarily from more open vegetation in
a small area dominated by west coastal strandveld shrubland just to the
north of Cape Town, South Africa.
Locality data (mean ± SD, range): Altitude: 63 ± 62, 15–199 m; annual
rainfall: 302 ± 29, 271–350 mm; annual temperature (max. + min. /2):
15.8 ± 0.2, 15.6–16.2°C (N=7).
Habitat: No quantitative assessment of soil type; recorded on deep sands
at Farm Geelbek (now West Coast National Park): few in natural shrub-
land (11), much more abundant where shrubland had been transformed
to sparse pasture grass (170); on deep sands in shrubland at nearby
Modderrivier: only 12 individuals; DBRU collection records from sand
(2) in sparse pasture (2) created by clearance of natural shrubland.
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Diurnal flight activity during the winter rainy season
(Apr. to Sept.) peaking in mid-winter (late June to early July) with lim-
ited activity in autumn (May) and spring (Aug.).
Bioregions South Africa: Known from only few vegetation units: Lange-
baan Dune Strandveld (FS 5), Atlantis Sand Fynbos (FFd 4), coastal
patches of Hopefield Sand Fynbos (FFd 3) and Saldanha Flats Strand-
veld (FS 3) (all Fynbos Biome).
Assessment rationale: EOO = 230 km2; high habitat transformation of
the species range by cultivation and urbanisation (FFd 3, FFd 4: 40%;
FS 3, FS 5: 35–50%), but higher abundance in pasture vegetation sug-
gests that this species is less influenced by transformation than most
endemic species; it may be less abundant than in the past at the type lo-
cality where shrubland has been permitted to regenerate across pastures
after Farm Geelbek was incorporated into West Coast National Park;
probably deserves at least Vulnerable (VU) status by virtue of highly
restricted known range, but currently assessed as Data Deficient (DD).
2 mm
Conservation measures: Further data on EOO, AOO and habitat as-
sociations are required to adequately assess the conservation status of
this species; protected at type locality in West Coast National Park, but
survey for relative abundance would be useful given the species bias to
occurrence in pastures and the current dominance of natural shrubland
within the park boundaries.

CANTHONINI
SURICATA 6 (2020) 105

Epirinus punctatus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Drakensberg, Cathedral Peak’ [1 800 m, KwaZulu-Natal,

South Africa].
Taxonomy: Accepted species, but specimens on Eswatini border should be
validated as E. punctatus.
Distribution: Recorded from both high rainfall forest and grassland locali-
ties on the cool highlands of the Drakenberg Escarpment, South Africa,
with the lowest locality at 1 167 m receiving high rainfall (1 157 mm
p/a); unclear if disjunction is a collection artefact.
Locality data (mean ± SD, range): Average altitude: 1 820 ± 549,
1 167–2 497 m; average annual rainfall: 954 ± 242, 751–1 271 mm;
average annual temperature (max. + min. /2): 13.2 ± 3.3, 9.2–17.4°C
(N=5).
Habitat: No quantitative assessment; probably primarily wet highland
forest patches although one available record is cited from grassland.
Food types: No quantitative assessment; not known.
Temporal activity: Presumably diurnal flight activity during the summer
rainy season (Oct. to Mar.).
Bioregions South Africa: Although five out of eight localities are cited
from forest (3) or forest stations (2), none of the cited grid referenc-
es map onto forest patches although patches occur nearby; in the S:
Northern Afrotemperate Forest (FOz 2); in the N: Northern Mistbelt
Forest (FOz 4). 1 mm
Assessment rationale: EOO = 17 500 km2; AOO probably much smaller,
centred on small patches of highland forest; known from < 10 locations
across a range of < 20 000 km2, therefore, qualifies for Vulnerable (VU)
status; part of range is protected; however, poorly known ecologically,
therefore, currently assessed as Data Deficient (DD).
Conservation measures: Before assessment may be made of conservation
status, a quantitative survey is required along the E escarpment between
the northeast border of Lesotho and northwest border of Eswatini;
this should examine the EOO, AOO and ecological associations with
regard to Northern Afrotemperate Forest patches, the S tip of North-
ern Mistbelt Forest patches, and adjacent grassland; protected in the
uKhahlamba Drakensberg Park (World Heritage Site).

CANTHONINI
106 SURICATA 6 (2020)

Epirinus pygidialus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Rep. South Africa: 75 km WSW Estcourt, Cathedral Peaks

For. Stn.’.
Taxonomy: Accepted species; individuals from Western and Eastern Cape
(black squares) require validation as E. pygidialus or a close undescribed
relative.
Distribution: Drakensberg escarpment from NE Lesotho border to
Mpumalanga (Mashishing); possibly also S coastline of Western and
Eastern Cape, South Africa (black squares – two individuals).
Locality data (mean ± SD, range): N range: average altitude: 2 100 ±
582, 1 575–2 700 m; average annual rainfall: 815 ± 43, 782–878 mm;
average annual temperature (max. + min. /2): 11.5 ± 3.5, 8.0–14.7°C
(N=4). S Cape range: average altitude: 81 ± 114, 0–161 m; average
annual rainfall: 435 ± 1, 434–435 mm; average annual temperature
(max. + min. /2): 16.5 ± 1.0, 15.9–17.2°C (N=2).
Habitat: No quantitative assessment; NE populations recorded at 1 500 m
in podocarp forest and on sandy clay loam at 2 400 m close to shrub­
land patches with Cape floral affinities; S Cape populations recorded
on coastline, possibly close to the beach.
Food types: Unknown.
Temporal activity: Diel flight periodicity unknown; assumed to be diur-
nal like other Epirinus spp.; recorded during spring in winter rainfall
region (Aug.), autumn in bimodal rainfall region (Apr.) and summer in
mid-summer rainfall region (Dec., Jan.).
Bioregions South Africa: In the S Cape: Fynbos biome; in KZN: North-
2 mm
ern Afrotemperate Forest (FOz 2); in E high altitude units: Drakensberg
Grassland (Gd), Mesic Highveld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 18 240 km2 or 7 810 km2; poorly known
with few, sometimes inexact, records in habitats ranging from shrub
patches in grassland to fynbos or forest; assessed as Data Deficient
(DD) owing to uncertain taxonomy and few widespread records.
Conservation measures: Further individuals and taxonomic study are re-
quired to determine if Eastern Cape and KwaZulu-Natal/Mpumalanga
populations belong to a single species; further survey and quantitative
data on ecological associations are required to determine conservation
status as the few available records apparently originate from various
habitat types; NE populations protected in the uKhahlamba Drakens-
berg Park (World Heritage Site).

CANTHONINI
SURICATA 6 (2020) 107

Epirinus relictus
Scholtz & Howden, 1987

No synonyms
Global: LC (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Drakensberg, Cathedral Peak’ [KwaZulu-Natal, South

Africa].
Taxonomy: Accepted species.
Distribution: Drakensberg escarpment along NE Lesotho border and
central S coast of Eastern Cape, South Africa; unclear if disjunct pat-
tern is real or a collection artefact; Mpumalanga record may represent
mislabeled material (black square).
Locality data (mean ± SD, range): Average altitude: 1 761 ± 968,
398–3 377 m; average annual rainfall: 701 ± 220, 354–1 037 mm;
average annual temperature (max. + min. /2): 11.8 ± 4.1, 3.9–16.4°C
(N=11).
Habitat: No adequate quantitative assessment; recorded in fynbos shrub­
land and grassland at higher altitude (> 1 850 m) and in forest at lower
altitude (400–1 300 m); along a gradsect (500 m to 2 800 m on lat-
itude 29–30°S): sampled in natural grassland on sandy clay loam at
1 900 m (2.0 per trap) and close to shrubland patches with Cape floral
affinities at 2 400 m (0.3 per trap).
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Diel flight periodicity unknown; assumed to be di-
urnal like other Epirinus spp.; recorded during summer in both the
bimodal (spring autumn) and mid-summer rainfall regions (Nov. to
Mar.).
Bioregions South Africa: Vegetation units: North Swartberg Sandstone
Fynbos (FFs 23) (Fynbos Biome); Southern Afrotemperate Forest (FOz
1), Southern Mistbelt Forest (FOz 3); also Drakensberg Grassland
Bioregion (Gd) (Grassland Biome). 2 mm
Assessment rationale: EOO = 50 250 km2 (excludes Mpumalanga re-
cord); EOO assumes continuous rather than disjunct distribution;
AOO would be very much smaller; relatively poorly known ecological-
ly, meriting a Data Deficient (DD) rating; however, assessed, here, as
Least Concern (LC) as most records are from reserves.
Conservation measures: Assessment of conservation status would be im-
proved by a quantitative survey across the known EOO; this should
determine if distribution is continuous or naturally disjunct; it should
also assess food and soil associations as well as apparent change in veg-
etation associations with altitude; protected in Tsitsikamma National
Park, uKhahlamba Drakensberg Park (World Heritage Site); conserva-
tion may be dependent on continuing protection in reserves.

CANTHONINI
108 SURICATA 6 (2020)

Epirinus rugosus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. Afr., Cape, Cederberg, Jeep track, 1 550 m, 32.23 S
19.08 E’ [mountain range, Western Cape, South Africa].
Taxonomy: Accepted species; but paratype from Cape Town (63 km N) is
E. pseudorugosus Medina & Scholtz, 2005; paratype from Blesberg-W
in the Swartberg, (cited as Blesberg-S) subsequently described as a new
species (see Preamble: E. inparrugosus Deschodt & Davis, 2019).
Distribution: After removal of misidentified paratypes; known by only
seven individuals recorded on the Cederberg, Western Cape, South
Africa.
Locality data (mean ± SD, range): Altitude: 1 293 ± 515, 944–1 885 m;
annual rainfall: 273 ± 39, 247–318 mm; annual temperature (max. +
min. /2): 13.2 ± 2.7, 10.1–15.1°C (N=3).
Habitat: Not known.
Food types: Not known.
Temporal activity: Flight activity assumed to be diurnal like other Epir-
inus species; recorded during the winter rainy season (Sept.) from
ground traps set for 63 days.
Bioregions South Africa: Vegetation unit: only Cederberg Sandstone
Fynbos (FFs 4) (Fynbos Biome).
Assessment rationale: EOO = ca. 15 km2; (undoubtedly underestimated
as this is the area covered by three study sites within a vegetation unit
that covers 2 124 km2); even if this potential range is confirmed, with
a third under protection, E. rugosus would qualify for an IUCN threat
category; however, as EOO is uncertain and there are no recorded eco-
logical data, it is assessed as Data Deficient (DD).
2 mm
Conservation measures: A quantitative survey of EOO, AOO and eco-
logical associations on the Cederberg is required to assess conservation
status; probably protected within the 710 km2 of the Cederberg Wil-
derness Area.

CANTHONINI
SURICATA 6 (2020) 109

Epirinus scrobiculatus
Harold, 1880

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cap bon. spei (Berg!)’ [Cape of Good Hope (mountain),
South Africa].
Taxonomy: Accepted species.
Distribution: West Coast, South Africa; primarily in the lowland sand-
veld below the escarpment; mountain type locality would be atypical;
citation from Namibia presumably an error.
Locality data (mean ± SD, range): Altitude: 177 ± 213, 0–899 m; annual
rainfall: 197 ± 80, 98–356 mm; annual temperature (max. + min. /2):
17.3 ± 0.9, 15.7–18.9oC (N=25).
Habitat: No adequate quantitative assessment; in Western Cape: sampled
primarily in shrubland on deep sand, Modderrivier (18), Pampoenvlei
(30); fewer on sandy loam in shrubland, Pampoenvlei (10), or pasture,
Groote Post (1); DBRU collection records on deep sand (3) in sparse
grass (2) or Succulent Karoo shrubs (1).
Food types: No quantitative assessment; DBRU collection records on
cattle (1) and horse (1) dung; sampled to cattle dung.
Temporal activity: Diurnal flight activity centred on the winter rainy sea-
son; in the SW Cape, active (Mar. to Nov.) peaking in autumn (May)
and spring (Aug.).
Bioregions South Africa: Vegetation units in Namaqualand Sandveld
(SKs), Western Strandveld (FS), Sand Fynbos (FFd) (Fynbos Biome);
marginal records in Namaqualand Hardeveld (SKn) (Succulent Karoo
Biome).
Assessment rationale: EOO = 15 275 km2; widespread within a restricted
range, apparently primarily on deep sand in shrubland; transformation 2 mm
limited in N of range (SKs: 4–8%) but extensive in the S (FFs, FS
vegetation units: 25–70%); assessed as Least Concern (LC) owing to
distribution centred primarily in a little-transformed region with ready
attraction to dung of domestic livestock in farm rangeland.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
West Coast and Namaqua national parks.

CANTHONINI
110 SURICATA 6 (2020)

Epirinus sebastiani
Medina & Scholtz, 2005

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘E. Transvaal [probably Uitsoek]’ [type locality (black

square) considered to be an error owing to mislabeling; several para-
types from the Amatole Mountains, Eastern Cape, South Africa (red
square), are considered to represent the true range of the species].
Taxonomy: Accepted species.
Distribution: Probably restricted to the Amatole Mountains, Eastern
Cape, South Africa.
Locality data (mean ± SD, range): Amatole Mountains data: Altitude:
1 242 ± 232, 1 078–1 406 m; annual rainfall: 706 ± 25, 688–723 mm;
annual temperature (max. + min. /2): 14.3 ± 1.2, 13.5–15.1°C (N=2).
Habitat: No quantitative assessment; recorded from litter on the floor of
indigenous forest.
Food types: No quantitative assessment; sampled to faeces bait in ground
traps.
Temporal activity: Flightless; diurnal activity; recorded during the sum-
mer rainy season (Nov., Dec.).
Bioregions South Africa: Southern Mistbelt Forest (FOz 3) (Forest
Biome).
Assessment rationale: EOO = 130 km2 (underestimated), AOO =
116 km2 (area of Pirie and Isidenge forests); currently known only
from two large patches of FOz 3 forest comprising the easternmost
in a group centred on the Amatole Mountains; these form the SW
limit of FOz 3 forest along the eastern seaboard of South Africa; species
assessed as Data Deficient (DD), but may deserve at least Vulnerable
2 mm (VU) status due to small EOO and restricted AOO in forest patches,
at least one of which is under pressure from resource utilisation (Pirie).
Conservation measures: Protection of FOz 3 forest in the Amatole
Mountains is essential for the conservation of E. sebastiani; a survey
of these patches, and of other groups of forest patches to N and S, is
required to determine its full EOO and ecological habits; a gradsect
from forest fragments into grassland would confirm its limited AOO
due to vegetation specialisation; protected in Isidenge State Forest.

CANTHONINI
SURICATA 6 (2020) 111

Epirinus silvestris
(Cambefort, 1978)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Endroedyantus silvestris: ‘Harkerville Forest’ [near Knys-

na, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on a small area of forest patches along the south
coast of Eastern Cape Province, South Africa; rare outlier occurrence in
Dwesa Forest (black square) requires validation after further compari-
son with E. convexus Scholtz & Howden, 1987.
Locality data (mean ± SD, range): Altitude: 237 ± 244, 26–841 m;
annual rainfall: 481 ± 38, 421–556 mm; annual temperature (max.
+ min. /2): 16.9 ± 1.1, 14.3–17.8°C (N=14) (excludes Dwesa Forest
outlier (black square)).
Habitat: No quantitative assessment; many locality labels specify forest
collection; soil types unknown.
Food types: No quantitative assessment; recorded on elephant dung.
Temporal activity: Flightless; diurnal activity; active in spring to early
summer in the bimodal autumn/spring rainfall region (Sept. to Dec.).
Bioregions South Africa: Largely localised in S Cape patches of Southern
Afrotemperate Forest (FOz 1).
Assessment rationale: EOO = 1 175 km2 (excludes Dwesa Forest outlier);
could qualify for a threat category on the basis of small EOO and even
smaller AOO; specialist association with forest of which an unknown
proportion has been transformed to tree plantation (at least 30 km2 in
2 mm
S Cape); currently assessed as Least Concern (LC) due to distribution
centred on a protected area.
Conservation measures: Conservation status is dependent on the con-
tinuing protection of low-altitude Afrotemperate forests along the
coastline of the Knysna region; as these occur on both sand or finer-
grained soils, an investigation is required to determine if there is any
localisation in population density according to soil type; protected in
the Garden Route National Park encompassing Tsitsikamma, Knysna
Lakes and Wilderness.

CANTHONINI
112 SURICATA 6 (2020)

Epirinus striatus
Scholtz & Howden, 1987

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Karroo, Zwartskraal Farm’ [N edge of Swartberg,

Western Cape, South Africa].
Taxonomy: Accepted species, but status of closely related, larger-bodied
populations at E edge of range needs assessment; that on the Eastern
Cape highlands N of Queenstown remains unassessed; that extending
across the Lesotho highlands to the Drakensberg on the NE Lesotho
border has subsequently been described as a new species (see Preamble,
E. muellerae Deschodt & Davis, 2019).
Distribution: Centred on the arid, cooler Upper Karoo with scattered
records elsewhere, particularly along the N edge of the Swartberg and
uplands on the SE Lesotho border.
Locality data (mean ± SD, range): Average altitude: 1 352 ± 176,
833–1 860 m; average annual rainfall: 306 ± 97, 195–696 mm; average
annual temperature (max. + min. /2): 15.2 ± 1.1, 12.2–17.9°C (N=53).
Habitat: No standardised quantitative assessment; in the Northern Cape:
extreme bias to cooler Upper Karoo (63.2%; 43 out of 68 study sites)
compared to warmer Karoo S of Orange River (11.6%; 14 out of 121
sites); absent from deep sands of SW Kalahari and arid Bushmanland
Karoo; bias to deep sand or loamy sand (5.9 per sample) compared to
sandy loam (4.0/sample) or sandy clay loam (1.0/sample).
Food types: No quantitative data; attracted to composite cattle/sheep
dung baits.
Temporal activity: Diurnal flight activity; seasonal occurrence reflects
distribution across cool, arid areas of the bimodal (spring/autumn) and
2 mm late summer rainfall regions (Sept. to June).
Bioregions South Africa: Primarily Upper Karoo (NKu) with scattered
records in four other bioregions.
Assessment rationale: EOO = 98 000 km2; depicted AOO may be a
collection artefact emanating from high collection intensity in a quan-
titative study on the Upper Karoo; widespread and not under threat
in natural vegetation on the farms of the Upper Karoo where habitat
transformation is minimal; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by standardised quantitative data on ecological associations;
not known from any protected area but well represented on farms in
natural Karoo shrubland.

CANTHONINI
SURICATA 6 (2020) 113

Epirinus sulcipennis
Boheman, 1857

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type localities: Misspelt as Epirhinus sulcipennis: ‘Caffraria meridionali et

ad Cap. Bonae Spei.’ [southern Caffraria and also Cape of Good Hope;
inexact, probably = S coast of South Africa], lectotype: ‘Cap B. Spei’
[Cape of Good Hope].
Taxonomy: Accepted species but individuals revised by Scholtz and How-
den (1987) require re-examination as they probably comprise three spe-
cies; (1) E. sulcipennis: W localities (Grootberg confirmed); (2) closely
related E. validus Péringuey, 1901: E localities (Eastern Cape and S
Free State, but not confirmed for Scholtz and Howden citations); (3)
Epirinus sp. nr striatus: N of Queenstown (confirmed).
Distribution: Probably restricted to S coastal uplands, South Africa;
possibly in both winter and bimodal rainfall regions; two confirmed
records at high altitude and low rainfall on Grootberg; cited occurrence
in wet coastal areas at Knysna requires validation.
Locality data (average): Altitude: 1 450 m; annual rainfall: 379 mm;
annual temperature (max. + min. /2): 10.9°C (N=1).
Habitat: No quantitative data; one confirmed record from an area com-
prising natural shrubland and sandy soils.
Food types: Unknown.
Temporal activity: Diel flight periodicity not known, but probably diur-
nal like other Epirinus species; recorded in Dec.
Bioregions South Africa: Confirmed occurrence only in one vegetation
unit: North Langeberg Sandstone Fynbos (FFs 15) (Fynbos Biome).
Assessment rationale: EOO and AOO unknown; however, FFs 15 veg-
etation unit extends from Western to Eastern Cape, is only 8% trans- 2 mm
formed, and may represent a refuge (13% conserved, 45% in mountain
catchment areas) where the species is not under threat; currently as-
sessed as Data Deficient (DD).
Conservation measures: A quantitative survey of the S upland and coastal
habitats in the Western and Eastern Cape is required to determine the
EOO, AOO, ecological associations and conservation status as well as
provide specimens for taxonomic revision; not currently known from
any protected area.

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Epirinus validus
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal (Lydenburg)’ [Mashishing, Mpumalanga, South

Africa].
Taxonomy: Accepted species.
Distribution: Edge of E escarpment, South Africa: at high altitude in N
(Mpumalanga, KwaZulu-Natal), descending to coastal hills in S (East-
ern Cape).
Locality data (mean ± SD, range): Average altitude: 1 772 ± 754,
303–3 218 m; average annual rainfall: 793 ± 119, 518–1 011 mm;
average annual temperature (max. + min. /2): 13.0 ± 3.8, 4.8–21.8°C
(N=26).
Habitat: No adequate quantitative assessment; at 30°S on Sani Pass: sam-
pled in natural montane grassland on sandy clay loam: abundant at
1 900 m (42.5/trap), rare at 2 400 m (0.5) and 2 800 m (0.04), absent
at lower altitude (1 500 m, 1 000 m, 500 m); DBRU collection records
on sandy loam (1) in grassland (1).
Food types: No quantitative assessment; DBRU collection records on
cattle dung (1); sampled in abundance to cattle dung.
Diel activity: Diurnal flight activity, primarily in the summer rainy season
(Sept. to May).
Bioregions South Africa: In the N: E edge Mesic Highveld Grassland
(Gm), NE edge Drakensberg Grassland (Gd); in the S: E edge Sub-
Escarpment Grassland (Gs), E edge Drakensberg Grassland (Gd) (all
Grassland Biome).
Assessment rationale: EOO = 52 550 km2; widespread, but poorly known
ecologically; transformation of escarpment edge vegetation units by
cultivation and tree plantations would presumably be detrimental (in
the N: 6–44%; in the S: 25–40%); however, assessed as Least Concern
2 mm
(LC) owing to abundance in protected areas along the NE border of
Lesotho where transformation is only 0–7%.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; as regards the
AOO, this should include possible detrimental effects on population
density of natural grassland modification to improved pastures; pro-
tected in uKhahlamba Drakensberg Park (World Heritage Site).

CANTHONINI
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Genus Gyronotus van Lansberge, 1874a

Type species and designation: Chalconotus pumilus Boheman, 1857, by subsequent designation (van Lansberge
1874a).
Synonyms: None.
Last review: Entire genus reviewed by Scholz and Howden (1987); new species added by Davis
et al. (1999b), Moretto and Perissinotto (2013); new species and a revalidation from
synonomy (Deschodt & Davis 2019)

When this book was completed at the end of 2017, the long-established genus, Gyronotus van Lansberge, 1874a, com-
prised eight flightless species centred on grasslands and forests of moist coastal and upland regions along the SE seaboard
of Africa from the Eastern Cape, South Africa, to NE Tanzania. However, a further three species have recently been de-
scribed and one other has been revalidated from synonomy (Deschodt & Davis (2019), see Preamble). The range has been
extended to SE Kenya.

Grassland spp.: Species associated with natural coastal or upland grasslands (3) have so far only been recorded in southern
Africa along the SE coast (KwaZulu-Natal, South Africa) or NE escarpment (South Africa, Eswatini). Conservation status
probably depends on the preservation of natural grassland.

Forest spp.: Five species associated with forest or dense shady vegetation have been recorded from the Eastern Cape in
South Africa to SE Kenya. Two South African species are found in forest patches along the SE coast with the more threat-
ened northerly species assessed as Vulnerable (VU).

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Gyronotus carinatus
Felsche, 1911

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘N. Calabar’ [Nigeria] or ‘Madagascar’ [Felsche doubted

that either were correct; known range of genus is SE to E Africa].
Taxonomy: Accepted species.
Distribution: Restricted to indigenous forest patches on the coastline and
coastal hills of NE KwaZulu-Natal, South Africa.
Locality data (mean ± SD, range): Altitude: 115 ± 120, 2–273 m; annual
rainfall: 964 ± 43, 900–998 mm; annual temperature (max. + min. /2):
21.1 ± 0.6, 20.5–21.8°C (N=5).
Habitat: No quantitative assessment of soil type; on deep dune sand near
Richards Bay: only sampled in undisturbed dune forest (386), none in
0–21 yr old regenerating vegetation (grassland, dense woodland or open
understorey woodland); DBRU collection records on finer-grained
soils: sandy clay loam (1), sandy loam (2) in forest (4) or woodland (1).
Food types: No quantitative assessment; sampled to composite baits of
cattle and pig dung.
Temporal activity: Flightless; diel periodicity not known; recorded in the
summer rainy season (Oct. to Feb.).
Bioregions South Africa: Scarp Forest (FOz 5), Northern Coastal Forest
(FOz 7) (Forest Biome).
Assessment rationale: EOO = 2 900 km2; occupies a small range with
an even smaller AOO due to vegetation specialisation in an area that
is subject to exploitation pressures through mining of coastal dunes
or forest resource utilisation; although protected in several reserves,
some remain pressurised by exploitation; under these circumstances,
2 mm a small AOO across a fragmented range with apparent sensitivity to
disturbance, coupled with flightlessness and low vagility, suggest that
the species warrants a threat category of at least Vulnerable (VU).
Conservation measures: Continuing protection of undisturbed scarp and
dune forest patches would be essential for the conservation of this spe-
cies; however, resource utilisation is widespread even in some officially
protected forest patches; although sampled in relative abundance in
Sokhulu Forest in 2000 and 2010, large parts of this forest fall within
a strip mining concession; protected in Hluhluwe–iMfolozi Game Re-
serve, oNgoye Forest Reserve, Mapelane Nature Reserve.

CANTHONINI
SURICATA 6 (2020) 117

Gyronotus glabrosus
Scholtz & Howden, 1987

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Magoebaskloof, N. Tvl.’ [Limpopo Province, South Afri-

ca].
Taxonomy: Accepted species.
Distribution: Primarily, upper NE edge of E escarpment, Limpopo Prov-
ince, South Africa.
Locality data (mean ± SD, range): Altitude: 1 276 ± 585, 575–2 005 m;
annual rainfall: 851 ± 76, 732–943 mm; annual temperature (max. +
min. /2): 16.7 ± 3.5, 12.5–18.9oC (N=5).
Habitat: No quantitative assessment; DBRU collection data from type
locality: alpine [montane] pasture on sandy loam; recent collections
support occurrence in grassland habitat.
Food types: No quantitative assessment; DBRU collection data for type
series: sampled to a human/cattle dung mixed bait.
Temporal activity: Flightless; diurnal activity; recorded during the sum-
mer rainy season (Dec., Feb.).
Bioregions South Africa: Vegetation units: Northern Escarpment
Quartzite Sourveld (Gm 23), Woodbush Granite Grassland (Gm 25)
(Mesic Highveld Grassland Bioregion, Grassland Biome); one record in
Tzaneen Sour Bushveld (SVl 8) (Lowveld Bioregion, Savanna Biome).
Assessment rationale: EOO = 550 km2; AOO would be smaller due to
habitat transformation; on the basis of small range < 5 000 km2 in only
four known locations would qualify for Vulnerable (VU) status partic-
ularly as it is flightless and apparently occurs in low population density
in a region subject to much transformation, primarily conversion to
tree plantations (Gm 25: 69%; Gm 23: 38%). 2 mm

Conservation measures: Assessment of conservation status would be im-

proved by a survey to determine the full EOO and AOO in montane
grasslands as well as the impact of habitat disturbance and transforma-
tion; protected in Lekgalameetse Nature Reserve.

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Gyronotus perissinottoi
Moretto, 2013

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘South Africa, KwaZulu-Natal, Umthamvuna Nature Re-

serve, Beacon Hill Section (31°00’47”S; 30°10’23”E)’.
Taxonomy: Accepted species.
Distribution: Currently known from only the type locality in Umtamvu-
na Nature Reserve, but possibly also represented in the past at nearby
Amanzimtoti (see notes with G. pumilus Boheman, 1857).
Locality data (average): Altitude: 161 m; annual rainfall: 966 mm; annu-
al temperature (max. + min. /2): 19.4°C (N=1).
Habitat: No quantitative assessment; described by Moretto and Perissinot-
to (2013) as ‘on escarpment at the edge of riverine forest, in grassland
interspersed with rocky outcrops and boulders’, amongst which most
individuals were recorded leading to speculation that this is where they
may secrete food.
Food types: No quantitative assessment; sampled to baboon dung.
Temporal activity: Flightless; diurnal activity; recorded during the sum-
mer rainy season (Jan.).
Bioregions South Africa: Vegetation unit: Pondoland-Ugu Sandstone
Coastal Sourveld (CB 4) (Indian Ocean Coastal Belt Biome).
Assessment rationale: EOO uncertain; only known from the grassland
portion of a single partially forested nature reserve covering a total of
0.324 km2; on the basis of minute range in only a single locality, would
qualify for an IUCN threat category of at least Vulnerable (VU), par-
ticularly considering the degree of transformation in the coastal zone
2 mm (CB 4 listed as VU; 29% transformed); however, currently assessed as
Data Deficient (DD).
Conservation measures: A quantitative survey of other natural grasslands
along the Pondoland and KwaZulu-Natal coastline is required to deter-
mine the EOO, AOO, ecological associations and conservation status;
protected in Umtamvuna Nature Reserve (South Africa).

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Gyronotus pumilus
(Boheman, 1857)

= Chalconotus marginatus Péringuey, 1888 (but see Taxonomy)

Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Chalconotus pumilus: ‘juxta fluvium Gariep’ [near Or-

ange River, South Africa; undoubtedly erroneous]. C. marginatus: ‘East
London, Cape Colony’ [Eastern Cape, South Africa]
Taxonomy: Accepted species, synonymy of the larger-bodied C. margi-
natus with G. pumilus now overturned (revalidated by Deschodt &
Davis (2019), see Preamble); in the N: larger-bodied specimens previ-
ously cited as G. pumilus should be compared to the recently described,
G. perissinottoi Moretto 2013.
Distribution: Endemic to coastal forest patches of Pondoland and south-
ern KwaZulu-Natal from East London to KwaDukuza; red squares
(G. pumilus), purple square (?G. perissinottoi), black squares (G. mar-
ginatus).
Locality data (mean ± SD, range): Altitude: 119 ± 125, 0–418 m; annual
rainfall: 914 ± 71, 741–977 mm; annual temperature (max. + min. /2):
19.5 ± 0.8, 18.4–20.9°C (N=18).
Habitat: No quantitative assessment; apparently centred on finer-grained
soils in lower-lying forests; forest origin is cited on 50% of locality la-
bels for 20 available localities; recorded as a rarity from one forest patch
in Vernon Crookes Nature Reserve, but not from grassland.
Food types: No quantitative assessment; attracted to a mixture of pig and
cattle dung, but dietary preference unknown.
2 mm
Temporal activity: Flightless, diurnal diel activity; recorded in the sum-
mer rainy season (Oct. to Feb.).
Bioregions South Africa: Centred on the southern half of the Scarp For-
est vegetation unit (FOz 5) (Forest Biome).
Assessment rationale: EOO = 6 350 km2; overall, 20% of FOz 5 is of-
ficially protected in reserves, but across the southern half of the unit,
uncontrolled exploitation of forest, land claims and mining proposals
are potential threats; in Umtamvuna Nature Reserve, G. pumilus occurs
in forest and G. perissinottoi in grassland, so it is presumed that forest
clearance would be detrimental to the former although only 5% of
FOz 5 is currently recorded as transformed; currently assessed as Least
Concern (LC) owing to moderate range size in more than 10 locations.
Conservation measures: Taxonomic examination of larger-bodied indi-
viduals is required to validate their identity as G. pumilus or as other
species; assessment of conservation status would be improved by quan-
titative data on ecological associations; however, continued protection
of Pondoland Scarp Forest patches along the coastline would be essen-
tial to conserve this endemic species; protection is currently offered
in various nature reserves, including, Dwesa-Cwebe, Silaka, Vernon
Crookes and Umtamvuna; proposals to protect this centre of ende-
mism by the creation of Pondoland National Park remain unfulfilled at
the time of writing.

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Gyronotus schuelei
Moretto, 2013

No synonyms
Global: DD
Endemic: Eswatini

J A S O N D J F M A M J

Type locality: ‘Swaziland, Mlilwane’ [Mlilwane Wildlife Sanctuary, Es-

watini].
Taxonomy: Accepted species.
Distribution: Currently known only by the holotype and allotype from
W Eswatini.
Locality data (mean ± SD, range): Altitude: 870 ± 28, 850–890 m; an-
nual rainfall: 922 ± 1, 921–923 mm; annual temperature (max. + min.
/2): 19.1 ± 0.1, 19.1–19.2°C (N=2).
Habitat: No quantitative assessment; described by Moretto and Perissinot-
to (2013) as: ‘in mountain grassland with pockets of sour bushveld’.
Food types: No quantitative assessment; holotype recorded on unidenti-
fied herbivore dung.
Temporal activity: Flightless; diurnal activity; recorded during the sum-
mer rainy season (Nov., Mar.).
Bioregions Eswatini: Straddles the boundaries of two vegetation units:
KaNgwane Montane Grassland (Gm 16) (Mesic Highveld Grassland
Bioregion, Grassland Biome) and Swaziland Sour Bushveld (SVl 14)
(Lowveld Bioregion, Savanna Biome).
Assessment rationale: EOO = 83 km2 (presumably underestimated);
currently known only from protected areas with a combined area of
22.6 km2; habitat transformation amounts to 21% (SVl 14) and 30%
(Gm 16), but range, precise associations and degree of threat unknown;
currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status; a survey is required
to determine the full EOO, AOO and ecological associations in W
Eswatini and (probably) across the border in South Africa; this should
also determine the level of threats from tree plantations, cultivation and
other disturbances; protected in Mlilwane Wildlife Sanctuary, Malalot-
ja Nature Reserve (Eswatini).
2 mm

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Genus Hammondantus Cambefort, 1978

Type species and designation: Hamondandantus psammophilus Cambefort, 1978, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Scholtz and Howden (1987).

Hammondantus Cambefort, 1978 is a monotypic genus that Scholtz and Howden (1978) indicate as probably misplaced
in the polyphyletic tribe, Canthonini (see placement in the phylogeny of Tarasov and Dimitrov (2016)). Most of its
known range is restricted to reserves in the dune fields of the S Namib Desert.

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Hammondantus psammophilus
Cambefort, 1978

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Sesriem Farm / Maltahöhe district’ [SW Namibia].

Taxonomy: Accepted species although classification in the tribe, Can-
thonini, has been questioned (Scholtz & Howden 1978).
Distribution: Restricted to Namibia where it is centred on the main
Namib Desert dune field; also occurs on Obib Dunes to the S.
Locality data (mean ± SD, range): Altitude: 845 ± 326, 400–1 248 m;
annual rainfall: 76 ± 39, 18–131 mm; annual temperature (max. + min.
/2): 16.1 ± 0.4, 15.6–16.7°C (N=6).
Habitat: Little precise data available, but recorded from sandy areas; ho-
lotype recorded from sand dunes; four of six known mapped localities
also coincide with sand dunes.
Food types: Not known.
Temporal activity: Flight periodicity unknown; recorded sporadically in
spring (Sept., Oct.), autumn (Mar.) and winter (June).
Ecoregions Namibia: Namib Desert (AT1315); also on Obib Dunes in
the N of the Succulent Karoo (AT1322) and the W edge of the arid
Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 13 600 km2; although poorly known eco-
logically, assessed as Least Concern (LC) on basis of sizeable range and
because all known localities but one lie within officially or privately
protected areas.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative ecological data; protected in Namib-Naukluft
and Sperrgebiet national parks and the private Namibrand Nature
Reserve.

2 mm

CANTHONINI
SURICATA 6 (2020) 123

Genus Namakwanus Scholtz & Howden, 1987

Type species and designation: Namakwanus irishi Scholtz & Howden, 1987, by original designation.
Synonyms: None.
Last review: Genus created by Scholtz and Howden, 1987; revised by Deschodt and Davis (2017);
new species (Deschodt & Davis 2018).

Namakwanus Scholtz & Howden, 1987 was created to accommodate a new flightless species from two different localities,
one in the highlands and one from the edge of the Namib Desert in Central Namibia. The single paratype from the second
locality has, subsequently, been described as a second new species (Deschodt & Davis 2017). Other species subsequently
described within Namakwanus have been transferred by Deschodt and Davis (2017) to Namaphilus Deschodt & Davis,
2017, or Versicorpus Deschodt, Davis & Scholtz, 2011. No species accounts are provided for the two new species (Deschodt
& Davis 2018) described subsequent to the completion of the book, but see Preamble.

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Namakwanus irishi
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Windhoek, S.W.A.’ [Namibia].

Taxonomy: Accepted species.
Distribution: Known by only four individuals from the inexact highland
type locality in central Namibia.
Locality data (average): Altitude: 1 815 m; annual rainfall: 353 mm;
annual temperature (max. + min. /2): 18.3°C (N=1).
Habitat: Unknown.
Food types: Unknown.
Temporal activity: Flightless; diel and seasonal periodicity unknown.
Ecoregions Namibia: Type series: NE Kalahari Xeric Savanna (AT1309).
Assessment rationale: EOO and AOO unknown; in the absence of distri-
butional and ecological data, assessed as Data Deficient (DD).
Conservation measures: A survey of the mountains around Windhoek
is required to determine the EOO, AOO and ecological associations;
only then will an assessment of conservation status be possible; not
currently known to occur in any protected area.

2 mm

CANTHONINI
SURICATA 6 (2020) 125

Namakwanus scholtzi
Deschodt & Davis, 2017

= Namakwanus irishi Scholtz & Howden, 1987 (pars – paratype from

Kuiseb River)
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘NAMIBIA, Gobabeb, Kuiseb River, Central Namib, S.

W. Afr.’.
Taxonomy: Accepted species formerly cited as a paratype of N. irishi
Scholtz & Howden, 1987.
Distribution: Inexact locality along the Kuiseb River near Gobabeb.
Locality data (average): Altitude: 400 m; annual rainfall: 18 mm; annual
temperature (max. + min. /2): 16.7°C (N=1).
Habitat: Unknown.
Food types: Unknown.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the late summer rainy season (May).
Ecoregions Namibia: Holotype: Namib Desert (AT1315).
Assessment rationale: EOO and AOO unknown; in the absence of distri-
butional and ecological data, assessed as Data Deficient (DD).
Conservation measures: A survey of the area around the Kuiseb River
near Gobabeb is required to determine the EOO, AOO and ecological
associations; only then will an assessment of conservation status be pos-
sible; protected in the Namib-Naukluft National Park.

2 mm

CANTHONINI
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Genus Namaphilus Deschodt & Davis, 2017

Type species and designation: Namakwanus endroedyi Deschodt, Davis & Scholtz, 2011, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Deschodt and Davis (2017); new species (Deschodt &
Davis 2018).

Namaphilus Deschodt & Davis, 2017, is a genus of flightless taxa. It was created to accommodate two species transferred
from Namakwanus Scholtz & Howden, 1987, plus one newly described species. All species have been recorded from rocky
areas in the arid Namibian savanna woodlands of central Namibia where one of the species has been observed to collect
dung pellets of the rock hyrax, which defecates in middens close to each colony. No accounts are provided for the two new
species (Deschodt & Davis 2018), described subsequent the the completion of the book, but see Preamble.

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SURICATA 6 (2020) 127

Namaphilus ameibensis
Deschodt & Davis, 2017

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘NAMIBIA, S.W.Afr., Erongo Mt., Farm Ameib’.

Taxonomy: Accepted species.
Distribution: Known only from the type locality in N central Namibia.
Locality data (average): Altitude: 1 197 m; annual rainfall: 210 mm;
annual temperature (max. + min. /2): 17.5°C (N=1).
Habitat: No quantitative assessment; holotype collected from under a
rock.
Food types: No quantitative assessment; unknown.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the late summer rainy season (Feb.).
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO unknown; little ecological data recorded
with the single known individual (holotype female); assessed as Data
Deficient (DD).
Conservation measures: A survey of the Erongo Mountains and nearby
highlands is required to determine the EOO, AOO, plus ecological
associations and to assess conservation status; not currently known to
be protected in any reserve.

2 mm

CANTHONINI
128 SURICATA 6 (2020)

Namaphilus davisi
(Deschodt & Scholtz, 2007)

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: As Namakwanus davisi: ‘Hardap Dam, Namibia’.

Taxonomy: Accepted species.
Distribution: Known only by the type series found adjacent to the camp-
site in the Hardap Resort, Namibia.
Locality data (average): Altitude: 1 164 m; annual rainfall: 176 mm;
average temperature (max. + min. /2): 21.0°C (N=1).
Habitat: No quantitative assessment; type series collected from sandy
loam soils at the base of a rocky outcrop in open scrub.
Food types: No quantitative assessment; observed to collect small pieces
of dung that are broken from hyrax pellets and dragged back to their
tunnels.
Temporal activity: Flightless; active during darkness, recorded during the
late summer rainy season (Mar.).
Ecoregions Namibia: S Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 25 km2 (probable underestimate based on
the protected area around Hardap Dam); AOO probably localised to
bases of rocky outcrops where rock hyrax middens provide an assured
dung supply in an arid region; probably not facing any threats, but full
extent of EOO currently unknown; ecological habits supported only by
limited observations; therefore assessed as Data Deficient (DD).
Conservation measures: A survey to determine the full EOO is required
before an assessment may be made of conservation status; localised
AOO, due to association with hyrax middens, should be confirmed by
gradsects away from several middens; protected in Hardap Game Park
flanking the Hardap Dam.

2 mm

CANTHONINI
SURICATA 6 (2020) 129

Namaphilus endroedyi
(Deschodt, Davis & Scholtz, 2011)

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: As Namakwanus endroedyi: ‘Naukluft Mountains [Namib-

ia]’.
Taxonomy: Accepted species.
Distribution: Known only from a few localities in the arid Naukluft
Mountains, S Namibia.
Locality data (mean ± SD, range): Altitude: 1 474 ± 181, 1 266–1 589 m;
annual rainfall: 155 ± 23, 128–170 mm; annual temperature (max. +
min. /2): 15.7 ± 0.3, 15.5–16.1°C (N=3).
Habitat: Collected on sandy soils in shrub/woodland on a steep rocky
hillside.
Food types: Sampled to banana bait, which is presumably a chance record.
Temporal activity: Flightless; one observation of diurnal foraging in
cool shade between boulders on a warm day; this single live individual
recorded under dry conditions during the late summer rainy season
(Jan.); only dead specimens collected in Mar.
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 280 km2; small known range extending
across a little-transformed area; but, very poorly known ecologically;
currently impossible to determine conservation status; assessed as Data
Deficient (DD).
Conservation measures: Before conservation status may be assessed, a
survey is required to assess the EOO, AOO and ecological associations;
part of the known range protected in Namib-Naukluft National Park. 2 mm

CANTHONINI
130 SURICATA 6 (2020)

Genus Nebulasilvius Deschodt & Scholtz, 2008

Type species and designation: Nebulasilvius insularis Deschodt & Scholtz, 2008, by original designation.
Synonyms: None.
Last review: Genus created by Deschodt and Scholtz (2008).

Nebulasilvius Deschodt & Scholtz, 2008, currently comprises two small-bodied, flightless species recorded from a river
bank or surface litter in different patches of Southern Mistbelt Forest, SE South Africa. Patches occur at lower altitude
than that supporting Parvuhowdenius Deschodt & Scholtz, 2008. Given the very small known ranges and limited ecolog-
ical data, both known Nebulasilvius species are currently assessed as Vulnerable (VU), pending surveys to provide more
information.

CANTHONINI
SURICATA 6 (2020) 131

Nebulasilvius insularis
Deschodt & Scholtz, 2008

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S.Africa; Natal Middld. Doreen Clarck Nat. R.’ [Doreen
Clark Nature Reserve, KwaZulu-Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Known only from a single, very small forest patch on the E
escarpment in the vicinity of Pietermaritzburg, KwaZulu-Natal, South
Africa.
Locality data (average): Altitude: 974 m; annual rainfall: 847 mm; annu-
al temperature (max. + min. /2): 16.9°C (N=1).
Habitat: No quantitative assessment; represented in museum collections
by 104 individuals recorded on a river bank in forest.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded in the
summer rainy season (Dec.).
Bioregions South Africa: N Southern Mistbelt Forest (FOz 3) (Forest
Biome).
Assessment rationale: EOO uncertain; AOO = 0.07 km2; (approximate
area of Podocarpus forest in Doreen Clark Nature Reserve); known only
from this reserve comprising an isolated forest patch on the lower E es-
carpment (Parvuhowdenius harrisoni Deschodt & Scholtz, 2008, occurs
at higher altitude along this stretch of FOz 3 forest patches); assessed
as Vulnerable (VU) on the basis of extremely small known range in a
single protected location; however, verging on Data Deficient (DD)
considering current limited state of knowledge.
Conservation measures: A survey of other nearby patches of FOz 3 forest
along the lower escarpment is required to determine if the EOO and
AOO are greater than currently known; a gradsect from grassland into
forest patches would confirm the vegetation specialisation and restrict-
ed AOO; protection of Doreen Clark Nature Reserve and any other 1 mm
occupied forest patches would be essential for the conservation of this
species.

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132 SURICATA 6 (2020)

Nebulasilvius johani
Deschodt & Scholtz, 2008

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S.Afr; S.Natal, Weza. Lower Stinkwood for.’ [S KwaZulu-

Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Known from one or two forest patches on the E escarpment
near Kokstad, KwaZulu-Natal, South Africa; validated for Weza Forest,
tentative for Bangeni Forest (possibly another species).
Locality data (mean ± SD, range): Altitude: 1 179 ± 11, 1 171–1 187 m;
annual rainfall: 815 ± 0.7, 814–815 mm; annual temperature (max. +
min. /2): 15.4 ± 0.1, 15.3–15.4°C (N=2).
Habitat: No quantitative assessment; sifted from forest litter and recorded
in ground traps.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded in the
summer rainy season (Nov.).
Bioregions South Africa: N Central patches of Southern Mistbelt Forest
(FOz 3) (Forest Biome).
Assessment rationale: EOO uncertain; validated AOO = 19 km2 (area
of Weza Forest); validated type locality in Weza–Ngele has State Forest
status and has become fragmented having been previously exploited for
timber; on the basis of small known range in few locations deserves at
least Vulnerable (VU) status although verging on Data Deficient (DD).
Conservation measures: A survey of the Eastern Mistbelt group of South-
ern Mistbelt Forest patches (FOz 3) is required to determine the full
EOO and AOO; a gradsect from grassland into forest patches would
confirm the vegetation specialisation and restricted AOO; conservation
status in Weza State Forest requires further research.

1 mm

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SURICATA 6 (2020) 133

Genus Odontoloma Boheman, 1857

Type species and designation: Odontoloma pauxillum Boheman, 1857, by monotypy.
Synonyms: = Caccobius (Diaglyptus) d’Orbigny, 1902: Type species: Caccobius metasternalis d’Or-
bigny, 1902, by monotypy (subsequently, raised to generic status as Diaglyptus
d’Orbigny, 1913, but synonymised with Odontoloma by Boucomont and Gillet
(1927)).
Last review: Entire genus reviewed by Howden and Scholtz (1987).

The long-established genus, Odontoloma Boheman, 1857, currently comprises 20 Afrotropical species with distributions
mostly, although not entirely, restricted to southern Africa. Most distributions occurring outside of South Africa, Namibia
and Botswana are biased towards the E in Zimbabwe, Democratic Republic of the Congo (DRC) or Ethiopia, with only
one species endemic to West Africa (Côte d’Ivoire). By virtue of very small body size and mostly close morphological sim-
ilarity, they are difficult to separate and require revision, particularly as the alpha taxonomy is incomplete.

A total of 12 species is currently listed as being restricted to South Africa (one also in Lesotho) with three further species
recorded from South Africa and Namibia (1), South Africa and Zimbabwe (1), or South Africa, Zimbabwe, Kenya and
Eritrea (identified as a species complex). Species accounts are provided for these 15 taxa on the understanding that iden-
tities and distribution patterns all require validation. Brief notes are provided to point out taxonomic problems associated
with each taxon.

Of the South African endemics, seven are more-or-less restricted to bimodal and/or winter rainfall regions; three to E sum-
mer rainfall and bimodal rainfall regions; two only to summer rainfall regions. However, the three other more widespread
species (see previous paragraph) also occur only in summer rainfall regions in southern Africa.

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Odontoloma apiculum
Howden & Scholtz, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. AFRICA, C.P. Olifantskop Pass betw. Port Elizabeth and
Cookhouse’ [Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Known only from the coastal Alexandria Forest and Oli-
fantskop Pass along the Sundays River Valley in the bimodal spring/
autumn rainfall region of the Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 227 ± 148, 123–332 m;
annual rainfall: 411 ± 101, 339–482 mm; annual temperature (max. +
min. /2): 18.3 ± 0.9, 17.7–18.9°C (N=2).
Habitat: No quantitative assessment; recorded from an area where the
natural vegetation is low succulent thicket or forest.
Food types: No quantitative assessment; no records.
Temporal activity: Humeral umbone reduced, but all related species
pterous; diel periodicity unknown, but probably diurnal like other
Odontoloma species; recorded during summer (Dec.) and autumn
(May).
Bioregions South Africa: Vegetation units: Sundays Noorsveld (AT 5)
(Albany Thicket Biome); Southern Coastal Forest (FOz 6) (Forest
Biome).
Assessment rationale: EOO = 905 km2 (presumably underestimated);
AOO may be much more restricted than EOO if O. apiculum is a shade
specialist; conservation status difficult to determine as range data are
severely limited and ecological data are absent; known from only two
localities although one is under protection; assessed as Data Deficient
(DD).
Conservation measures: Before conservation status may be assessed, it is
necessary to (1) determine the full EOO and AOO from a survey of the
area around Alexandria Forest and Olifantkop Pass and (2) to obtain
1 mm quantitative ecological data; protected in Alexandria Forest as part of
the Greater Addo Elephant National Park.

CANTHONINI
SURICATA 6 (2020) 135

Odontoloma dentinum
(Harold, 1868a)

= Diaglyptus serraticeps d’Orbigny, 1913

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Epirhinus dentinus: ‘Cap bon. spei’ [Cape of Good

Hope]; D. serraticeps: ‘Colonie du Cap’ [Cape Colony] [both SW
South Africa].
Taxonomy: Accepted species.
Distribution: Primarily coastal lowlands of the winter and bimodal rain-
fall regions of the Western and Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 179 ± 197, 0–681 m; annual
rainfall: 316 ± 126, 120–595 mm; annual temperature (max. + min.
/2): 17.0 ± 1.1, 14.9–18.7°C (N=21).
Habitat: In the Western Cape: well represented on deep sand to sandy
loam, and natural shrubland to pasture resulting from clearance of in-
digenous shrubs; however, on both the cooler Cape Peninsula and drier
West Coast, much more abundant in pasture (respectively, one site:
2 100; four sites: 854–1 301) than shrubland (respectively: two sites:
6–60; four sites: 2–114).
Food types: No quantitative assessment; sampled abundantly to cattle
dung baits.
Temporal activity: Diurnal flight activity; recorded during most months;
in Kikuyu pasture on Cape Peninsula (farm Bonne Attente), sampled
from winter to spring (June to Dec.) with peak numbers from Aug. to
Nov. and no activity in summer and autumn (Jan. to May).
Bioregions South Africa: Namaqualand Hardeveld (SKn), Namaqualand
Sandveld (SKs) (Succulent Karoo Biome); West Strandveld (F 11),
West Coast Renosterveld (FO 7) (Fynbos Biome); Albany Thicket
Biome (AT). 1 mm

Assessment rationale: EOO = 52 460 km2; widespread with high frequen-

cy of records; recorded at all 11 study sites during quantitative sampling
in the SW Western Cape; existing data suggest a soil generalist that
is especially abundant at sites transformed from natural shrubland to
pasture; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by standardised quantitative ecological data, but existing data
are probably adequate; protected in West Coast National Park (Western
Cape).

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136 SURICATA 6 (2020)

Odontoloma disalatum
Howden & Scholtz, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. AFRICA: Cape Mt. Matjisfontein’ [Western Cape,

South Africa].
Taxonomy: Accepted species.
Distribution: Dry often sandy areas, primarily in fynbos vegetation of the
Western Cape winter rainfall region.
Locality data (mean ± SD, range): Altitude: 289 ± 430, 0–1 415 m;
annual rainfall: 266 ± 110, 157–427 mm; annual temperature (max. +
min. /2): 16.9 ± 1.7, 12.5–18.5°C (N=11).
Habitat: No quantitative assessment; no records.
Food types: No quantitative assessment; holotype recorded on human
dung.
Temporal activity: Apterous; diel periodicity unknown, but probably
diurnal like other Odontoloma species; active during winter and spring
(June to Oct.).
Bioregions South Africa: Centred on Northwest (FO 1) and Southwest
(FO 4) Fynbos (Fynbos Biome); also Namaqualand Sandveld (SKs)
(Succulent Karoo Biome).
Assessment rationale: EOO = 43 165 km2; conservation status difficult
to assess as it is ecologically poorly known; apparently restricted to dry
areas in fynbos; also margins of S Succulent Karoo; occupies a wide
range, but could be potentially threatened by habitat transformation as
it is flightless; currently assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and improved
distribution data; possibly protected on the Cederberg.

1 mm

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SURICATA 6 (2020) 137

Odontoloma doubei
Howden & Scholtz, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘REP. SOUTH AFRICA: Natal, 75 km WSW Estcourt,

Cathedral Peaks For. Sta........ Rainbow Gorge, 1 500 m, Podocarp
Forest’ [Cathedral Peak Forest Station, KwaZulu-Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Moist edge of E escarpment possibly mainly in shaded sit-
uations: South Africa.
Locality data (mean ± SD, range): Altitude: 1 206 ± 495, 324–1 746 m;
annual rainfall: 833 ± 233, 484–1 157 mm; annual temperature (max.
+ min. /2): 14.8 ± 3.00, 9.2–17.4°C (N=6).
Habitat: No quantitative assessment; type locality in forest, but also re-
corded in coastal grassland and fynbos shrubland.
Food types: No quantitative assessment; sampled to both dung and car-
rion.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; recorded primarily during the summer
rainy season (Nov. to May).
Bioregions South Africa: Primarily E edges of Grassland Biome (Mesic
High­veld Grassland (Gm); Drakensberg Grassland (Gd); Sub-Escarpment
Grassland (Gs)), possibly mainly in forest patches; also Eastern Fyn-
bos-Renosterveld (FO 6).
Assessment rationale: EOO = 13 375 km2 (gross estimate); moderately
widespread, but poorly known ecologically; type locality may indicate
associations primarily with shaded situations although quantitative
support is required; currently assessed as Data Deficient (DD).
Conservation measures: Before conservation status may be adequately
assessed a survey is required to determine the full EOO and AOO;
quantitative data is required to establish ecological associations; pro-
tected in uKhahlamba Drakensberg Park (World Heritage Site).

1 mm

CANTHONINI
138 SURICATA 6 (2020)

Odontoloma endroedyi
Howden & Scholtz, 1987

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘SOUTH AFRICA: Swartberge, Hagas Farm, 1 050 m’

[Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on dry fynbos shrubland areas in the winter and bi-
modal rainfall regions of the Western and Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 913 ± 539, 97–1 993 m;
annual rainfall: 313 ± 70, 206–415 mm; annual temperature (max. +
min. /2): 15.0 ± 2.6, 9.6–18.7°C (N=11).
Habitat: No quantitative assessment; no records.
Food types: No quantitative assessment; attracted in abundance to human
dung.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; primarily active during spring, summer
and autumn (Sept. to May).
Bioregions South Africa: Centred on Fynbos and Renosterveld (FO 1,
FO 5, FO 6, FO 8) in various vegetation units: Sand Fynbos (FFd 8),
Sandstone Fynbos (FFs 4, FFs 21, FFs 22, FFs 23); margins of Shale
Renosterveld (FRs 6, FRs 16); marginal in Albany Thicket Biome (AT)
and Lower Karoo (NKl) (Nama Karoo Biome).
Assessment rationale: EOO = 45 810 km2; occupies a wide range with
records primarily from mountain localities where natural shrubland
vegetation still dominates (e.g. Matjiesfontein, Blesberg, Swartberg,
Gamkaberg); on the basis of abundance in collections and the wide
EOO, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
1 mm Gamkaberg Nature Reserve, Eastern Cape, South Africa.

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SURICATA 6 (2020) 139

Odontoloma louwi
Howden & Scholtz, 1987

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: ‘NAMIBIA/S.W.A. Etosha Pan, Okaukuejo’.

Taxonomy: Accepted species, but almost all of the type series from N
Namibia; most examined individuals from South Africa were excluded
from the type series due to shorter parameres and doubts on their spe-
cies identity.
Distribution: Disjunct pattern separated by deep Kalahari sands; type
series from upland savanna of N Namibia and N South Africa (red
squares); further specimens mostly recorded in grassland of the N
Highveld, South Africa, but excluded from type series (black squares).
Locality data (mean ± SD, range): Type series, mainly Namibia: altitude:
1 305 ± 373, 708–1 815 m; annual rainfall: 339 ± 92, 171–512 mm;
annual temperature (max. + min. /2): 18.6 ± 2.0, 16.5–22.2°C (N=11).
Excluded series, South Africa: altitude: 1 345 ± 245, 846–1 678 m;
annual rainfall: 675 ± 144, 433–878 mm; annual temperature (max. +
min. /2): 16.5 ± 1.5, 14.4–18.9°C (N=9).
Habitat: No quantitative assessment for type series; excluded series rep-
resented in Gauteng bushveld: strong bias to finer-grained soils (sandy
clay loam: 293; deep sand: 43) with generalist vegetation associations
(grassland: 119; open woodland: 76; shaded thicket: 141).
Food types: No quantitative assessment.
Temporal activity: Diel periodicity unknown, but probably diurnal like
other Odontoloma species; active during the summer rainy season (Oct.
to Apr.).
Ecoregions Namibia: Type series: Namibian Savanna Woodlands
(AT1316), Angolan Mopane Woodlands (AT0702); NW Kalahari
Xeric Savanna (AT1309).
Bioregions South Africa: Type series: Central Bushveld (SVcb) (Savan-
na Biome); excluded series: primarily Dry Highveld Grassland (Gh), 1 mm
Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs)
(Grassland Biome).
Assessment rationale: EOO Namibia = 131 940 km2; EOO South Africa
= 75 220 km2; unclear if O. louwi comprises one or two species; how-
ever, populations of both elements widespread, therefore, assessed as
Least Concern (LC).
Conservation measures: Accurate assessment of conservation status re-
quires further taxonomic study to establish if one or two species are
included under the name, O. louwi; further quantitative ecological
data are also required, particularly, with regard to the type series; pop-
ulations represented by the type series protected in Etosha National
Park; populations of excluded group not currently known from any
protected area.

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140 SURICATA 6 (2020)

Odontoloma obscurum
Howden & Scholtz, 1987

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type locality: ‘LESOTHO, Drakensbg. Sani Pass Valley’ [Lesotho: pre-

sumably upper Sani Pass].
Taxonomy: Accepted species.
Distribution: SE escarpment from high rainfall and high altitude in the N
to low rainfall and low altitude in the S: Lesotho, South Africa.
Locality data (mean ± SD, range): Altitude: 1 838 ± 1 310, 97–3 218 m;
annual rainfall: 688 ± 283, 389–1 001 mm; annual temperature (max.
+ min. /2): 11.4 ± 5.6, 4.8–18.7°C (N=5).
Habitat: On an altitudinal gradsect in KwaZulu-Natal, recorded only
in low numbers at high altitude (0.3 per sample at 2 400 m, 0.8 at
2 800 m) on sandy clay loam in grassland.
Food types: No quantitative assessment; attracted to cattle dung.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active primarily during the summer
rainy season (Jan. to May).
Bioregions South Africa: Drakensberg Grassland (Gd) (Grassland
Biome); Albany Thicket Biome (AT); Eastern Fynbos-Renosterveld
Bioregion (FO 6) (Fynbos Biome).
Assessment rationale: EOO = 8 020 km2 (gross underestimate); poorly
known species both biogeographically and ecologically; therefore, as-
sessed at Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative
survey is required to determine the full EOO, AOO and ecological
associations; protected in the uKhahlamba Drakensberg Park (World
1 mm Heritage Site).

CANTHONINI
SURICATA 6 (2020) 141

Odontoloma pauxillum
Boheman, 1857

= Caccobius (Diaglyptus) pluridens d’Orbigny, 1905

Global: LC

J A S O N D J F M A M J

Type localities: O. pauxillum: ‘prope fluvium Gariep’ [near Orange River,

South Africa, although female type labelled ‘Caffraria’]; C. pluridens:
‘Nyassa’ [type labelled: ‘Mozam. Nyassa’ = Mozambique].
Taxonomy: Accepted species.
Distribution: Recorded from scattered localities across southern African
savanna into the E and NE tropics: South Africa, Zimbabwe, Mozam-
bique, Kenya, Ethiopia.
Locality data (mean ± SD, range): Altitude: 924 ± 476, 64–1 383 m;
annual rainfall: 629 ± 197, 306–852 mm; annual temperature (max. +
min. /2): 20.1 ± 1.5, 18.9–22.1°C (N=5).
Habitat: No quantitative assessment; however, most of the few known
southern African localities coincide with deep sands: E coast (Sodwana
Bay); Southern African Bushveld (Nylsvlei, Boekenhoutskloof, Mma-
boela Estate); Zimbabwean Miombo woodland (Harare).
Food types: No quantitative assessment; no records.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active primarily during the summer
rainy season (Oct. to Apr.).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb) (Savanna Biome);
Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 021 280 km2 (gross estimate); wide-
spread EOO, but AOO likely smaller because of putative specialisation
to sandy soils; recorded in high frequency on deep sands at Nylsvlei;
assessed as Least Concern (LC) on basis of wide range, but essentially
Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected at
Nylsvlei Nature Reserve (South Africa).

1 mm

CANTHONINI
142 SURICATA 6 (2020)

Odontoloma peckorum
Howden & Scholtz, 1987

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘REP. SOUTH AFRICA: Natal, 75 km WSW Estcourt,

Cathedral Peaks For. Sta.’ [Cathedral Peak Forest Station, KwaZulu-
Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Moist upland or highland grassland along the edges of the
NE escarpment, South Africa.
Locality data (mean ± SD, range): Altitude: 1 544 ± 463, 974–2 497 m;
annual rainfall: 816 ± 52, 726–914 mm; annual temperature (max. +
min. /2): 14.4 ± 2.3, 9.2–16.9°C (N=12).
Habitat: On an altitudinal transect in KwaZulu-Natal, sampled in grass-
land on sandy clay loam soils primarily at 1 500 and 1 900 m (11.9
or 19.8 per trap), uncommon at 1 000 (2.4), 2 400 (0.1) and 2 800 m
(0.04).
Food types: No quantitative assessment; sampled to cattle dung baits.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active during the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: E edges of Drakensberg Grassland (Gd), Mesic
Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland
Biome).
Assessment rationale: EOO = 19 885 km2; wide range, primarily in
grassland along NE escarpment where substantial local abundance is
recorded at 1 500–1 900 m; large part of range at higher altitude under
protection; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
uKhahlamba Drakensberg Park (World Heritage Site).

1 mm

CANTHONINI
SURICATA 6 (2020) 143

Odontoloma planatum
Howden & Scholtz, 1987

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘SOUTH AFRICA: Natal, 75 km WSW Estcourt, Cathedral

Peaks For. Sta., 1400 m’ [Cathedral Peak Forest Station, KwaZulu-Natal,
South Africa].
Taxonomy: Accepted species.
Distribution: Mostly moist upland and coastal grassland along lower NE
escarpment and SE coast, South Africa.
Locality data (mean ± SD, range): Altitude: 1 096 ± 811, 114–2 497 m;
annual rainfall: 900 ± 140, 726–1 157 mm; annual temperature (max.
+ min. /2): 16.3 ± 3.5, 9.2–19.5°C (N=7).
Habitat: No quantitative assessment, no records.
Food types: No quantitative assessment, no records.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active during the summer rainy season
(Sept. to Feb.).
Bioregions South Africa: E edges of Drakensberg Grassland (Gd), Mesic
Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland
Biome); also Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 14 615 km2; ecologically poorly known,
but occupies a moderately large range so currently assessed as Least
Concern (LC) although essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative ecological data; unclear whether or not it has
been recorded from protected areas.
1 mm

CANTHONINI
144 SURICATA 6 (2020)

Odontoloma pusillum
Howden & Scholtz, 1987

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘[S. AFRICA], S.W. Cape Prov., Langebaan (12 km SE

Geelbek)’ [Farm Geelbek, now in West Coast National Park].
Taxonomy: Accepted species.
Distribution: Dry west coastal deep sands of the SW Western Cape,
South Africa.
Locality data (mean ± SD, range): Altitude: 94 ± 79, 31–199 m; annual
rainfall: 273 ± 68, 191–350 mm; annual temperature (max. + min. /2):
16.6 ± 1.0, 15.7–17.9°C (N=5).
Habitat: On the West Coast: primarily recorded at farms on W coastal
deep sands; at Geelbek: equally abundant in natural strandveld shrub­
land (550) and adjacent sparse grassland cleared of shrubs (530); at
Pampoenvlei much more abundant in shrubland on deep sand (287)
than on adjacent sandy loam (31).
Food types: No quantitative assessment; sampled to cattle dung in abun-
dance.
Temporal activity: Diurnal flight activity; seasonal activity during the
winter rainy season in autumn, winter and spring (Apr. to Oct.) with
autumn (May) and spring peaks (Aug., Sept.).
Bioregions South Africa: West Strandveld (F 11) vegetation units: Lange-
baan Dune Strandveld (FS 5), Lamberts Bay Strandveld (FS 1); also
Southwest Fynbos (FO 3), Atlantis Sand Fynbos (FFd 4) (all Fynbos
Biome).
Assessment rationale: EOO = 1 685 km2; possibly restricted to a small
natural range in an area subject to extensive habitat transformation
(FFd 4: 40%, FS 1: 25%, FS 5: 35%); on the basis of small EOO
< 5 000 km2 at only five locations; would qualify for an IUCN threat
category of Endangered (EN), but on the basis of small body size and
local abundance in a protected area with tolerance of vegetation trans-
formation, currently tentatively assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
1 mm proved by quantitative data on food type associations particularly in
West Coast National Park as cattle have been removed from the type
locality on the Farm Geelbek since the proclamation of the park; as-
sumed to be protected within West Coast National Park.

CANTHONINI
SURICATA 6 (2020) 145

Odontoloma pygidiale
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Cape Colony (Cape Town, Stellenbosch, Paarl)’ [Western

Cape, South Africa].
Taxonomy: Accepted species, but Howden and Scholtz (1987) noted
clinal differences in morphology, particularly in relative prominence of
the elytral umbone, which is associated with relative wing development
and may indicate a species complex; close to O. pusillum Howden &
Scholtz, 1987 and O. endroedyi Howden & Scholtz, 1987.
Distribution: Disjunct pattern, which may or may not indicate more
than one species; records mostly from dry to arid, lower-altitude parts
of winter and bimodal rainfall regions plus NE edge of Karoo, South
Africa.
Locality data (mean ± SD, range): Altitude: 334 ± 409, 1–1 408 m;
annual rainfall: 243 ± 91, 108–448 mm; annual temperature (max. +
min. /2): 16.8 ± 1.4, 13.2–19.2°C (N=47).
Habitat: No quantitative assessment; no records.
Food types: No quantitative assessment; no records.
Temporal activity: Diel periodicity unknown, but probably diurnal like
other Odontoloma species; recorded during all months except mid-
summer (Jan.), but most records in spring in the Western Cape and
spring plus autumn in the Eastern Cape.
Bioregions South Africa: Primarily Namaqualand Hardeveld (SKn),
Rainshadow Valley Karoo (SKv) (Succulent Karoo Biome); Northwest
Fynbos (FO 1), Southwest Fynbos (FO 2) (Fynbos Biome); NE Upper
Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 71 165 km2; widespread in arid regions
that are little transformed; probable that most populations are not
threatened, so irrespective of questions on taxonomic identity, the spe- 1 mm

cies or species complex may be assessed as Least Concern (LC).

Conservation measures: Assessment of conservation status would be im-
proved by (a) a taxonomic study to determine if O. pygidiale comprises
a single species or a complex, and (b) quantitative data on ecological
associations; currently no records from protected areas.

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Odontoloma quadridens
(d’Orbigny, 1908)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: As Caccobius (Diaglyptus) quadridens: ‘Rhodesia, Plumtree’

[Plumtree, Zimbabwe].
Taxonomy: Accepted species revalidated by Howden and Scholtz (1987)
after synonymy with O. pauxillum Boheman, 1857, by Janssens
(1938a) and Ferreira (1969).
Distribution: Bushveld of the E Kalahari and its outlier patches: South
Africa, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 040 ± 321, 708–1 349 m;
annual rainfall: 486 ± 156, 306–585 mm; annual temperature (max. +
min. /2): 20.3 ± 1.5, 19.3–22.1°C (N=3).
Habitat: No quantitative assessment; the three known localities coincide
with patches of deep sands.
Food types: No quantitative assessment; no records.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active during the summer rainy season
(Oct., Mar.).
Ecoregions Zimbabwe: Southern African Bushveld (AT0717).
Bioregions South Africa: Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 15 205 km2 (presumably underestimated);
ecologically poorly known; occupies a large EOO, but AOO may be
limited by soil specialisation, although this has not been supported
quantitatively; currently assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative ecological data and a survey to determine its full
EOO and AOO; protected in Nylsvlei Nature Reserve.

1 mm

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Odontoloma sculpturatum
(Harold, 1868a)

= Caccobius (Diaglyptus) multifidus d’Orbigny, 1908

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Epirhinus sculpturatus: ‘Cap bon. spei’ [Cape of Good

Hope, South Africa], lectotype: ‘Oranje, Burghisrd.’ [South Africa];
C. multifidus: ‘Colonie du Cap’ [Cape Colony, South Africa].
Taxonomy: Accepted species.
Distribution: Widespread along the SE seaboard of South Africa from
upland summer rainfall areas in the N to lowland winter and bimodal
rainfall areas in the S.
Locality data (mean ± SD, range): Altitude: 861 ± 808, 93–1 371 m;
annual rainfall: 597 ± 187, 392–858 mm; annual temperature (max. +
min. /2): 16.3 ± 2.7, 10.6–18.7°C (N=7).
Habitat: No quantitative assessment; no records but N distribution pri-
marily in grassland regions.
Food types: No quantitative assessment; no records.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
like other Odontoloma species; active primarily during spring and the
summer rainy season in the N (Sept. to Apr.).
Bioregions South Africa: Dry Highveld Grassland (Gh), Drakenberg
Grassland (Gd), Sub-Escarpment Grassland (Gs) (Grassland Biome);
Sub-Escarpment Savanna (SVs) (Savanna Biome); Albany Thicket
Biome (AT); Southwest and Eastern Fynbos (FO 2, FO 6).
Assessment rationale: EOO = 64 415 km2 (gross estimate); widespread
across climatically different areas, but ecologically poorly known; there-
fore assessed as Data Deficient (DD).
Conservation measures: Before conservation status may be assessed a
survey is required to improve distributional data (EOO, AOO) and
provide quantitative ecological data; not currently known from any
nature reserve.
1 mm

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Odontoloma spinicaudum
Howden & Scholtz, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. AFRICA: Namaqualand, Klein Kogel Fontein’ [farm in

Namaqualand, South Africa].
Taxonomy: Accepted species.
Distribution: Known from only the type locality on the arid West Coast
sandveld, South Africa.
Locality data (average): Altitude: 38 m; annual rainfall: 142 mm; annual
temperature (max. + min. /2): 18.2°C (N=1).
Habitat: No quantitative assessment; recorded in a area of deep sands
where the natural vegetation is coastal shrubland.
Food types: No quantitative assessment; no records.
Temporal activity: Wings obsolete; diel periodicity unknown, but prob-
ably diurnal like other Odontoloma species; recorded in spring during
the winter rainy season (Aug.).
Bioregions South Africa: Vegetation unit: Namaqualand Strandveld (SKs
7) (Namaqualand Sandveld Bioregion, Succulent Karoo Biome).
Assessment rationale: EOO unknown; known only by the single female
holotype; conservation status impossible to determine in absence of
range and ecological data; assessed as Data Deficient (DD).
Conservation measures: Before conservation status may be assessed, it is
necessary to survey the sandveld around Klein Kogel Fontein and ob-
tain quantitative ecological data; not known from any protected area.

1 mm

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Genus Outenikwanus Scholtz & Howden, 1987

Type species and designation: Outenikwanus tomentosus Scholtz & Howden, 1987, by original designation.
Synonyms: None.
Last review: Genus created by Scholtz and Howden (1987).

Outenikwanus Scholtz & Howden, 1987, is small-bodied, flightless and monotypic. It has been recorded from leaf litter
of a single Southern Afrotemperate Forest patch (FOz 1) on mountains along the S seaboard of the Eastern Cape, South
Africa. It has been assessed as Data Deficient (DD) although the combined area of upland patches of FOz 1 would prob-
ably be sufficiently small to justify an assessment of Vulnerable (VU).

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Outenikwanus tomentosus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. Cape Mts., Outenikwa Pass’ [Eastern Cape, South Af-
rica].
Taxonomy: Accepted species.
Distribution: Currently known only from an indigenous forest patch
adjoining the Outeniqua Pass, Outeniqua Mountains.
Locality data (average): Altitude: 999 m; annual rainfall: 404 mm; annu-
al temperature (max. + min. /2): 13.8°C (N=1).
Habitat: No quantitative assessment; recorded from leaf litter of indige-
nous forest.
Food types: Not known.
Temporal activity: Flightless, diel periodicity unknown; recorded during
early summer (Nov., Dec.) in the bimodal rainfall region (spring, au-
tumn peaks).
Bioregions South Africa: Southern Afrotemperate Forest (FOz 1) (Forest
Biome).
Assessment rationale: EOO uncertain; currently, only known by the type
series and a few other individuals, all recorded in forest on the Outeni-
qua Pass; nothing else known concerning EOO, AOO and ecological
habits; currently assessed as Data Deficient (DD).
Conservation measures: EOO, AOO and conservation status may only
be assessed after a survey of the many small scattered upland patches
of Afrotemperate indigenous forest between Mossel Bay and Jeffreys
Bay in the S Cape; a gradsect from grassland into forest patches would
confirm the vegetation specialisation and restricted AOO; currently
only known to be protected within Witfontein Nature Reserve on the
Outeniqua Pass.
1 mm

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SURICATA 6 (2020) 151

Genus Parvuhowdenius Deschodt & Scholtz, 2008

Type species and designation: Parvuhowdenius harrisoni Deschodt & Scholtz, 2008, by original designation.
Synonyms: None.
Last review: Genus created by Deschodt and Scholtz (2008).

Parvuhowdenius Deschodt & Scholtz, 2008, is small-bodied, flightless and monotypic. Known only from litter in a single
small patch of Southern Mistbelt Forest. The patch occurs at higher altitude than those supporting the genus, Nebulasilvius
Deschodt & Scholtz, 2008. Given the very small known range and limited ecological data, Parvuhowdenius is assessed as
Data Deficient (DD), although an alternative assessment as Vulnerable (VU) is probably justified.

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Parvuhowdenius harrisoni
Deschodt & Scholtz, 2008

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Karkloof For. 1 300 m’ [Karkloof Forest, KwaZulu-Natal,

South Africa].
Taxonomy: Accepted species.
Distribution: Monotypic genus currently known only from the type lo-
cality: Karkloof Forest.
Locality data (average): Altitude: 1 364 m; annual rainfall: 855 mm;
annual temperature (max. + min. /2): 14.7°C (N=1).
Habitat: No quantitative assessment; all known specimens sifted from
forest leaf litter; assumed to be a forest specialist.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the summer rainy season (Dec.).
Bioregions South Africa: Southern Mistbelt Forest (FOz 3) (Forest
Biome).
Assessment rationale: EOO = 16.9 km2 (size of Karkloof Forest patch
in which P. harrisoni was recorded); currently known by only three
individuals from a single privately owned forest patch that is under sig-
nificant threat; deserves at least Vulnerable (VU) status, but so poorly
known, assessed as Data Deficient (DD).
Conservation measures: Before an assessment of conservation status may
be made, it is necessary to determine its full EOO and population
density by conducting a further survey of Karkloof Forest and many
other nearby fragments of FOz 3; a gradsect from forest fragments into
grassland would confirm its limited AOO due to vegetation specialisa-
tion; protected by the Karkloof Conservancy comprising landowners,
conservation and forestry organisations.

0.5 mm

As published in Deschodt & Scholtz (2008).

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Genus Peckolus Scholtz & Howden, 1987

Type species and designation: Peckolus parvus Scholtz & Howden, 1987, by original designation.
Synonyms: None.
Last review: Genus created by Scholtz and Howden (1987); new species added by Howden and
Scholtz (1988), Deschodt and Scholtz (2008).

Peckolus Scholtz & Howden, 1987, currently comprises three species from small Northern Mistbelt or Northern Afrotem-
perate forest patches along the upper edge of the NE escarpment of South Africa. Although all three species are assessed as
Data Deficient (DD), the rarity of collection and small putative EOO and AOO would deserve IUCN threat categories
of at least Endangered (EN), pending thorough surveys of other forest patches.

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Peckolus alpinus
Howden & Scholtz, 1988

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘South Africa, Transvaal, 11 km S.E. Pilgrims Rest, 1 500 m’

[Mpumalanga, South Africa].
Taxonomy: Accepted species.
Distribution: Localised on upper edge of E escarpment within the north-
ernmost of three discrete blocks of FOz 4 forest patches forming the
Mpumalanga Mistbelt Forest Region, South Africa.
Locality data (mean ± SD, range): Altitude: 1 647 ± 218, 1 493–1 800 m;
annual rainfall: 961 ± 4, 958–963 mm; annual temperature (max. +
min. /2): 14.6 ± 1.3, 13.7–15.5°C (N=2).
Habitat: No quantitative assessment; observed in ravines supporting
patches of relict forest with well-developed leaf litter.
Food types: No quantitative assessment; sampled to carrion.
Temporal activity: Flightless; diel periodicity unknown; recorded in the
summer rainy season (Dec.).
Bioregions South Africa: Northern (Limpopo) division of Northern
Mistbelt Forest (FOz 4) (Forest Biome).
Assessment rationale: EOO = 15 km2; current known forest AOO =
4.8 km2 (both presumably underestimated); probably occurs in nu-
merous other larger, nearby forest patches that remain unsurveyed; the
FOz 4 vegetation unit is cited as poorly protected, but not currently
threatened; species assessed as Data Deficient (DD) since range and
ecological habits are poorly known; however, would currently deserve
an IUCN threat category of at least Endangered (EN) on basis of AOO
< 500 km2 at < 5 known localities.
Conservation measures: For the conservation of this species, it would
be essential to protect the block of FOz 4 forest patches found on the
escarpment edge between Sabie and Klaserie; to determine the EOO,
it would be necessary to survey the high altitude FOz 4 forest patches
1 mm along the entire NE edge of the escarpment in Mpumalanga and Lim-
popo; a gradsect from grassland into forest patches would confirm the
vegetation specialisation and restricted AOO; protected in the private
Mt Sheba Nature Reserve.

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Peckolus parvus
Scholtz & Howden, 1987

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Natal, 75 m WSW Estcourt, Cathedral Peaks For. Sta.’

[South Africa].
Taxonomy: Accepted species.
Distribution: Known only from a single high altitude locality on the E
escarpment of South Africa on the NE border of Lesotho.
Locality data (average): Altitude: 2 497 m; annual rainfall: 810 mm;
annual temperature (max. + min. /2): 9.2°C (N=1).
Habitat: No quantitative assessment; observed in high altitude (1 800 m)
podocarp forest litter containing moss and fungal hyphae; probably a
forest specialist.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the summer rainy season (Dec.).
Bioregions South Africa: Northern Afrotemperate Forest (FOz 2) (Forest
Biome).
Assessment rationale: EOO = 1.4 km2 (presumably underestimated;
comprises area of type locality, forest patch); range and ecological hab-
its poorly known; therefore, assessed as Data Deficient (DD); the small
known AOO of < 500 km2 at < 5 locations would deserve an IUCN
threat category of at least Endangered (EN); however, the single patch
of montane forest from which it is known, occurs within a protected
area containing other similar forest patches; therefore, despite minute
known range, possibly not currently under serious threat.
Conservation measures: Continuing protection of FOz 2 forest patches
would be essential for the conservation of this species; to determine
the EOO, it is necessary to conduct a survey of the many other FOz
2 forest patches that occur along the W border of KwaZulu-Natal ad-
joining Lesotho, Free State and Mpumalanga; a gradsect from grassland
into forest patches would confirm the vegetation specialisation and re- 1 mm
stricted AOO; protected in the uKhahlamba Drakensberg Park (World
Heritage Site).

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Peckolus poenskopius
Deschodt & Scholtz, 2008

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Nelshoogte, Knuckles rocks for.’ [Forest near Knuckles

Rocks, Mpumalanga, South Africa].
Taxonomy: Accepted species.
Distribution: Localised on E escarpment within the southernmost of
three discrete blocks of FOz4 forest patches forming the Mpumalanga
Mistbelt Forest Region, South Africa.
Locality data (mean ± SD, range): Altitude: 1 085 ± 224, 831–1 257 m;
annual rainfall: 1 005 ± 137, 892–1 157 mm; annual temperature
(max. + min. /2): 18.0 ± 1.4, 17.0–19.6°C (N=3).
Habitat: No quantitative assessment; recorded in wet forest litter.
Food types: Not known.
Temporal activity: Flightless; diel periodicity unknown; recorded in the
summer rainy season (Feb.).
Bioregions South Africa: Southern (Mpumalanga) division of Northern
Mistbelt Forest (FOz 4) (Forest Biome).
Assessment rationale: EOO = 85 km2 (presumably underestimated); may
occur in numerous other nearby forest patches that remain unsurveyed;
the FOz 4 vegetation unit is cited as poorly protected, but not currently
threatened; species assessed as Data Deficient (DD) since range and
ecological habits are poorly known; however, current known EOO of
< 5 000 km2 at < 5 locations would deserve an IUCN threat category
of at least Endangered (EN).
Conservation measures: For the conservation of this species, it would
be essential to protect the block of FOz 4 forest patches found on the
escarpment near the Eswatini border between Barberton and Malelane;
to determine the EOO, it would be necessary to survey the lower-
altitude FOz 4 forest patches along the entire NE edge of the escarp-
ment in Mpumalanga and Limpopo; a gradsect from grassland into for-
1 mm
est patches would confirm the vegetation specialisation and restricted
AOO; protected in Nelshoogte and Josephdal nature reserves.

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Genus Pycnopanelus Arrow, 1931

Type species and designation: Pycnopanelus rotundus Arrow, 1931, by original designation.
Synonyms: None.
Last review: Genus created by Arrow (1931); African species reviewed by Scholtz and Howden,
(1987).

Pycnopanelus Arrow, 1931, currently comprises three species with disjunct distributions in the Afrotropical (2) and W
Oriental regions (India: 1). The Indian species is represented only by the type specimen. The African species have been
recorded from arid regions in the SW and NE (Sudan: two specimens). The SW species from the arid late summer rain-
fall region of Namibia, Botswana and South Africa is known by many individuals and is assessed as Least Concern (LC).
Scholtz and Howden (1987) express their doubts that Pycnopanelus belongs with the Canthonini (see placement in the
phylogeny of Tarasov and Dimitrov (2016)).

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Pycnopanelus krikkeni
Cambefort, 1978

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Noachebeb, 27 mls NNE Grunau’ [S Namibia].

Taxonomy: Accepted species.
Distribution: Warmer parts of the arid late summer rainfall region from
South Africa across SW Botswana and W Namibia; probably also SW
Angola, but no known records.
Locality data (mean ± SD, range): Altitude: 987 ± 208, 361–1 782 m;
annual rainfall: 191 ± 65, 39–423 mm; annual temperature (max. +
min. /2): 18.8 ± 1.2, 15.8–21.3°C (N=110).
Habitat: In Northern Cape: high frequency of records in warmer, arid
areas, especially Bushmanland (5.7/trap at 91% of 67 study sites) on
sand (3.8/trap), loamy sand (6.1), sandy loam (2.3), sandy clay loam
(0.7); DBRU collection records from sand (4), sandy loam (1) in grass-
land (1), scrub/shrubland (3), open woodland (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (2), donkey (1); sampled in large numbers to composite
sheep/cattle dung baits.
Temporal activity: Diel flight periodicity unknown, possibly diurnal;
recorded during the late summer rainy season (Jan. to Apr.) and in the
winter dry season in central Namibia (June, July).
Ecoregions Namibia, Botswana: NW Nama Karoo (AT1314), SW
Kalahari Xeric Savanna (AT1309), Namibian Savanna Woodlands
(AT1316).
Bioregions South Africa: Low-altitude N Bushmanland (NKb) (Nama
Karoo Biome) with representation in the Kalahari Duneveld (SVk)
1 mm (Savanna Biome).
Assessment rationale: EOO = 264 500 km2; does not face any threats
due to widespread distribution across an arid area used primarily for
conservation or the grazing of domestic livestock whose dung it readily
colonises; transformation: close to 0% (both NKb and SVk); assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would ben-
efit from further quantitative ecological data, particulartly food asso-
ciations; protected in Kgalagadi Transfrontier Park (Botswana, South
Africa) and inland margin of the Namib-Naukluft Park (Namibia).

CANTHONINI
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Genus Silvaphilus Roets & Oberlander, 2010

Type species and designation: Silvaphilus oubosiensis Roets & Oberlander, 2010, by original designation.
Synonyms: None.
Last review: Genus created by Roets and Oberlander (2010).

Silvaphilus Roets & Oberlander, 2010, is small-bodied, flightless and monotypic. It has been recorded from dung and leaf
litter of a single Southern Afrotemperate Forest patch (FOz 1) on mountains along the S seaboard of the Western Cape,
South Africa. By virtue of extremely small known range on private land, it currently deserves an assessment of Vulnerable
(VU) or even Endangered (EN).

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Silvaphilus oubosiensis
Roets & Oberlander, 2010

No synonyms
Global: VU
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Western Cape Province, Riviersonderend, Oubos Forest’

[South Africa].
Taxonomy: Accepted species.
Distribution: Known only from Oubos Forest, Riviersonderend Moun-
tains, Western Cape, South Africa.
Locality data (average): Altitude: 389 m; annual rainfall: 367 mm; annu-
al temperature (max. + min. /2): 16.3°C (N=1).
Habitat: No quantitative assessment; recorded only within forest after a
search of herbivore dung both outside and inside forest habitat; also
found in leaf litter and in top 10 mm of soil under dung.
Food types: No quantitative assessment; recorded under fresh mammalian
dung, both herbivore pellets and an insectivore dropping; not recorded
under old dung.
Temporal activity: Flightless; diel periodicity unknown; recorded in late
spring at end of winter rainy season (Nov.).
Bioregions South Africa: Southern Afrotemperate Forest (FOz 1) (Forest
Biome).
Assessment rationale: EOO 6 km2 (estimated); AOO = 1.6 km2 (Oubos
Forest); only known from a single small, privately owned forest patch
in an isolated mountain range where only one other even smaller forest
patch occurs (0.7 km2); owing to the extremely small potential EOO
and AOO in few locations, this species would qualify for at least Vul-
nerable (VU) status, possibly even Endangered (EN).
Conservation measures: A survey is required to determine if this mono-
typic genus might occur in any of the other few patches of Southern
Afrotemperate Forest found in the Western Cape or if it is endemic
1 mm to the Riviersonderend Mountains; however, owing to its isolation, it
seems likely that protection of Oubos Forest and the only other forest
patch in this mountain range would be essential for the conservation of
S. oubosiensis; Oubos Forest is conserved by the owners.

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Genus Versicorpus Deschodt, Davis & Scholtz, 2011

Type species and designation: Versicorpus erongoensis Deschodt, Davis & Scholtz, 2011, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Deschodt and Davis (2017).

Versicorpus Deschodt, Davis & Scholtz, 2011, now comprises two species from arid mountain blocks in Namibian Sa-
vanna Woodlands, Central Namibia, after a second species was added to V. erongoensis Deschodt, Davis & Scholtz, 2011,
following transferral from Namakwanus Scholtz & Howden, 1987 (Deschodt & Davis 2017). As both species are known
by only the holotype specimens, they are assessed as Data Deficient (DD) with no further speculative assessment; both
type localities occur on private, but little-transformed, farm rangeland. One new species has been recently described from
the Brandberg, Namibia (Deschodt & Sole 2019).

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Versicorpus erongoensis
Deschodt, Davis & Scholtz, 2011

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘S.W.AFR. [Namibia], ErongoMT [Mountains]., Farm

Amieb’ [Farm Ameib, Erongo Mountains, Namibia].
Taxonomy: Accepted species.
Distribution: Known only by a single individual from the Erongo Moun-
tains, Namibia.
Locality data (average): Altitude: 1 197 m; annual rainfall: 210 mm;
annual temperature (max. + min. /2): 17.5°C (N=1).
Habitat: Unknown; holotype found under a stone.
Food types: Unknown.
Temporal activity: Flightless; diel periodicity unknown; recorded during
the late summer rainy season (Feb.).
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO unknown; little ecological data recorded
with the single known individual (holotype female); assessed as Data
Deficient (DD).
Conservation measures: To assess conservation status, a survey is required
of the Erongo Mountains and nearby highlands; this should determine
the EOO, AOO and ecological associations; not currently known to be
protected in any reserve.

2 mm

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Versicorpus streyi
(Frolov, 2005)

No synonyms
Global: DD (see IUCN Red List – DD as Namakwanus streyi)
Endemic: Namibia

J A S O N D J F M A M J

Type locality: As Namakwanus streyi: ‘Bullspoort [Bullsport 16o22’E

24o8’S], S.W.A.’ [farm in Namibia].
Taxonomy: Accepted species.
Distribution: Known only from the somewhat inexact type locality in
arid mountains of S Namibia (possibly Remhoogte Mts).
Locality data (average): Altitude: 1 621 m; annual rainfall: 179 mm;
annual temperature (max. + min. /2): 15.9°C (N=1).
Habitat: Unknown.
Food types: Unknown.
Temporal activity: Flightless; diel and seasonal periodicity unknown.
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO unknown; extremely poorly known species;
known only by a single individual (holotype male); no ecological data
available; assessed as Data Deficient (DD).
Conservation measures: A survey is required to determine the EOO,
AOO plus ecological associations and to assess conservation status;
possibly protected within the Naukluft part of Namib-Naukluft Na-
tional Park.

2 mm

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164 SURICATA 6 (2020)

TRIBE COPRINI
Leach, 1815

As Coprides: Type genus: Copris Geoffroy, 1762.

Synonyms:
= Pinotinae Kolbe, 1905; Type genus: Pinotus Erichson, 1847b [=Dichotomius Hope, 1838, senior name] (but see
Tarasov and Dimitrov 2016).
= Coptodactylini Paulian, 1933: Type genus Coptodactyla Burmeister, 1846.
= Dichotomiini Perreira, 1954: Type genus: Dichotomius Hope, 1838 (but see Tarasov and Dimitrov 2016).

Although they were listed as synonyms of the tribe Ateuchini by Bouchard et al. (2011), the recently named
tribe Dichotomiini and older Pinotinae are here listed as synonyms of the Coprini to which the type genus,
Dichotomius [= syn. Pinotus] and other Copris-like genera were transferred by the morphological phylogeny
of Montreuil (1998). However, this synonymy would be superseded by the recent partial tribal revision
of Tarasov and Dimitrov (2016), which has defined the Neotropical and Nearctic Dichotomius lineage as
Dichotomiini sensu novo.

As defined by Montreuil (1998), the tribe Coprini is represented globally within areas of tropical or warm
temperate climate. At the present time, 19 genera may be assigned to the tribe according to morpholog-
ical criteria (14 with morphological phylogenetic support). These occur in three biogeographical centres
(Afro-Eurasia: 12; Americas: 6; Australia/New Guinea: 2) with Copris Geoffroy, 1762, shared between Af-
ro-Eurasia and the Americas. However, molecular phylogenies show that the tribal membership (as defined
by Montreuil 1998) is polyphyletic with six distantly related lineages (Monaghan et al. 2007; Tarasov &
Dimitrov 2016) in just 42% (8) of the genera (Monaghan et al. 2007). As a result, Tarasov and Dimitrov
(2016) redefine a single lineage (genera: Copris, Litocopris) as Coprini sensu novo. All other coprine genera
are awarded the status of incertae sedis. Under these circumstances the old tribal classification system is used
in this book pending a full revision.

In the old sense, a total of 11 genera have been recorded for the Coprini in the Afrotropical region al-
though some of these have recently (Marchisio & Zunino 2012) been claimed to be synonyms of Copris
(see generic account for Litocopris Waterhouse, 1891). Hypothesised relationships differ between molecular
phylogenies. Monaghan et al. (2007) indicate at least three distantly related lineages: (1) the basally derived
Macroderes Westwood, 1842; (2) Copris plus Heliocopris Hope, 1837; and (3) the more terminally derived
Catharsius Hope, 1837, plus Metacatharsius Montreuil, 1998 (but see discussion on authorship under ge-
neric account for Metacatharsius).

The recent partial tribal revision of Tarasov and Dimitrov (2016) also indicates at least three to four distant-
ly related groupings of lineages, but suggests that none are basally derived. In rank order of derivation, these
are: (1) Copris plus Litocopris now termed Coprini sensu novo; (2) Catharsius and Metacatharsius; with (3)
Heliocopris in a neighbouring clade; (4) Xinidium Harold, 1869a, with Macroderes in a neighbouring clade.
Subsequent to the preparation of this book in 2017, Catharsius and Metacatharsius have been transferred to
the new tribe, Catharsiini by Takano (2018) although this transfer currently remains unpublished. Here,
these genera and all three lineages remain included as incertae sedis in the tribe Coprini (old sense).
SURICATA 6 (2020) 165

All three of these lineages are represented in Botswana, Namibia and/or South Africa where a total of seven
genera are found.
(1A) Copris Geoffroy, 1762: Widely distributed genus in the Americas and Afro-Eurasia; represented
by 33 southern African species centred primarily on winter and bimodal rainfall (6), Kalahari (3),
upland grassland (7), E forest or grassland (3) or savanna (14) regions.
(1B) Litocopris Waterhouse, 1891: Species-poor Afrotropical genus; represented in South Africa by two
species with SE or E seaboard distributions.
(2A) Catharsius Hope, 1837: Widely distributed genus in the Afrotropical and Oriental regions repre-
sented by 16 southern African species in upland grasslands (4), Kalahari (2), E forests and grasslands
(2) and savannas (8).
(2B) Metacatharsius Montreuil, 1998 (but see discussion on authorship under generic account for Meta-
catharsius): Widely distributed genus found primarily on sandy soils in the Afrotropical region;
represented by 11 validated southern African species centred mainly in the Kalahari (4), Kalahari/
Namib (1), Kalahari/Namib/savanna (3), Kalahari/savanna (2) or SW Arid (1) regions.
(3) Heliocopris Hope, 1837: Widely distributed genus in the Afrotropical and Oriental regions repre-
sented by seven southern African species primarily found in arid to moist savannas.
(4A) Macroderes Westwood, 1842: Flightless genus comprising 14 species endemic to winter and bimodal
rainfall regions, South Africa, after the revision of Frolov and Scholtz (2004). Further species were
described by Abdalla et al. (2018) after the completion of this book (no species accounts presented
here, but see Preamble).
(4B) Xinidium Harold, 1869a: Species-poor genus restricted to grassland and forest of the SE African
highlands; represented by three South African species.

In the old sense, Coprini comprise medium to large-bodied, tunnelling taxa, some of which are known
to construct brood ovoids within subterranean chambers. Most taxa do not currently face serious threats.
However, two flightless Copris species are assessed as Endangered (EN) due to extreme habitat loss on the S
coast in the winter and bimodal rainfall regions of South Africa. The poorly known flightless genus, Mac-
roderes, may also face threats due to small species ranges, particularly those with more southerly occurrence
where habitat transformation is greater.

COPRINI
166 SURICATA 6 (2020)

Genus Catharsius Hope, 1837

Type species and designation: Scarabaeus molossus Linnaeus, 1758, by original designation (Hope 1837).
Synonyms: None.
Last review: Afrotropical species reviewed by Ferreira (1960a, 1960b) with a few new species
described subsequently (Ferreira 1962a, 1963, 1971; Josso & Prévost 2009b; Josso
2011); recent review of a species group (Moretto 2008) and a discussion of Harold’s
types (Génier & Josso 2016); revision of entire genus (Takano, pending publication).

Catharsius Hope, 1837, was described as a subgenus of Copris Geoffroy, 1762 and has subsequently been raised to generic
level. There is no phylogeny for the genus and although it clearly comprises a number of lineages, there has been no formal
division into species groups. It currently comprises circa 100 valid species, most of which are Afrotropical (approx. 85)
with fewer in the Oriental region (14) and one shared between the Afrotropical and Palaearctic (Egypt) regions. Takano
(2018) has reviewed the genus and a number of name changes plus reassessments of synonyms are pending publication
(indicated in species accounts).

Catharsius species are large to very large bodied. All species are macropterous and fly in darkness. They are considered to
show similarities in behaviour to Copris in that dung is first buried at the end of a shallow tunnel and then relocated to a
deeper tunnel. However, Catharsius species construct single brood balls per chamber. These are placed in series along the
tunnel or adjacent at the tip of the tunnel. As this nesting strategy requires large amounts of dung, Catharsius are mostly
found in moister regions together with large herbivores and their large droppings.

Of 16 species with valid names found in southern Africa, 12 are restricted to the region with three showing wider dis-
tributions. Some occur primarily on sandy soils, others on finer-grained soils. There seems to be a strong bias to carrion,
omnivore and monogastric herbivore dung unlike in Copris. Three are specialists on monogastric herbivore dung.

In southern Africa, distributions of three species are centred on N Highveld Grassland (2) or on Highveld Grassland and
Upper Karoo (1). However, most species show savanna distributions. Two are centred on the deep sands of the Kalahari,
one a carrion specialist, the other biased to omnivore and monogastric herbivore dung. Two others are centred on the
deep E coastal sands of Maputaland, one in grassland, one in forest. Some are centred across the savanna belt and beyond
into the Kalahari (1) or onto the Highveld and S coast (1). Some are restricted to just savanna from E to W (1), or further
restricted by separations between E and W (1), or occurrence only in the E (4).

A total of 11 species are assessed as Least Concern (LC) because they are widespread or well protected in reserves con-
taining elephants. The remaining five species are assessed as Data Deficient (DD) because they are ecologically poorly
known and are represented from few localities. These species include the only endemic southern African monogastric
dung specialist (C. bradshawi van Lansberge, 1887), which could be facing threats.This discussion of wider distribution
and conservation status is based on the older nomenclature, which precedes the soon to be published revision of Takano.

COPRINI
SURICATA 6 (2020) 167

Catharsius aegeus
Génier, 2017

No synonyms
Global: LC (see IUCN Red List – LC; cited as C. pandion Harold,
1877)

J A S O N D J F M A M J

Type locality: ‘MOZ.: CABO DELGADO | Mareja (site 3), P.N. Qui-
rimbas’ [NE Mozambique, Quirimbas National Park].
Taxonomy: Accepted species; recently found to comprise an undescribed
species after having been wrongly identified as C. pandion Harold,
1877, which is an E coastal endemic (see species account); sexually
dimorphic (head and prothoracic disc), prominence of characters varies ♂
with body size.
Distribution: Southern African savanna: N South Africa, NE Namibia,
NE Botswana, Zimbabwe, Zambia, Malawi, Mozambique; reports
from Angola, Democratic Republic of the Congo, Tanzania, require
validation.
Locality data (mean ± SD, range): Average altitude: 792 ± 410,
55–1 477 m; average annual rainfall: 631 ± 166, 260–1 136 mm; av-
erage annual temperature (max. + min. /2): 20.7 ± 1.9, 16.9–25.2°C
(N=48).
Habitat: No quantitative assessment; in Botswana; sampled on deep sands
in open shrubland (70); few DBRU collection records on sand (2),
sandy loam (1) in forest (3) or open woodland (2).
Food types: In Botswana: only sampled from carrion (chicken livers) (70),
not sampled to any dung type, pig, elephant, cattle, sheep (all 0) (cited
as C. pandion Harold); on Gauteng bushveld: sampled only to rotting
sheep offal (4) not to any dung type, pig, horse, cattle (all 0); carrion
specialisation may account for the few DBRU collection records: mixed
human/cattle dung bait (2), gut contents of slaughtered impala (1);
also cited from rotting fish, dead millipedes and in two instances from
5 mm
banana.
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Jan.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian and Mopane
Woodlands (AT0725), Southern African Bushveld (AT0717); scattered
records in Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambe-
zian Baikiaea Woodlands (AT0726), Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Primarily in Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 812 345 km2 (underestimated); somewhat
Data Deficient (DD), but probably a vegetation generalist and carri-
on specialist showing a bias to sandy soils, which would restrict the
AOO; poor data from which to generate an assessment, but widespread
records across farm rangeland and reserves suggest few threats at the
present time; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations and population
density; protected in several national parks: Kruger (South Africa),
Chobe (Botswana).

♀ 5 mm

COPRINI
168 SURICATA 6 (2020)

Catharsius bradshawi
van Lansberge, 1887

= Catharsius insignis Péringuey, 1892

Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type localities: C. bradshawi: ‘Afrique orient. : Zambèze’ [presumably

eastwards on the Zambezi River]; C. insignis: Not stated.
Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
Distribution: Difficult to assess owing to inexact localities for older refer-
ence specimens; recorded in the 1970s along the Zambezi River in N
♂ Zimbabwe; recorded in the 19th century along the Okavango and Evari
rivers [Evari = upper reaches of Etaka River, NW of Etosha Pan] (DM,
NCI) suggesting occurrence in N Botswana and N Namibia; also cited
from Mozambique, which would, presumably, be an interpretation of
the type locality.
Locality data (mean ± SD, range): Altitude: 717 ± 234, 373–895 m;
annual rainfall: 650 ± 7, 641–659 mm; annual temperature (max. +
min. /2): 23.6 ± 1.5, 22.4–25.9°C (N=4).
Habitat: No quantitative assessment; limited DBRU collection records on
sandy loam (1), sandy clay loam (1) in grassland (2).
Food types: No quantitative assessment; limited DBRU collection records
on dung of elephant (2), buffalo (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov., Feb.).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725)
in the Zambezi Valley.
Assessment rationale: EOO = 52 950 km2 (gross estimate, N Zimbabwe
to upper Etaka River, N Namibia); not well represented in reference
collections and poorly known ecologically; AOO probably much small-
5 mm er than EOO due to possible bias to finer-grained soils in grassland on
dung of monogastric herbivores, particularly elephant, but supporting
quantitative data is required; rarity of modern reference material may
be due to range contraction of monogastric herbivores, but this also
requires testing; assessed as Data Deficient (DD), but may deserve an
IUCN threat category.
Conservation measures: Before an assessment of conservation status may
be made, a quantitative survey is required concentrating on reserves
containing elephants on the floodplains along the Okavango and Zam-
bezi river systems to determine the full EOO, AOO and ecological
associations; protected in Victoria Falls National Park (Zimbabwe).

5 mm ♀

COPRINI
SURICATA 6 (2020) 169

Catharsius calaharicus
Kolbe, 1893a

= Catharsius disseptus Péringuey, 1901 (but see Taxonomy)

Global: LC (See IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: C. calaharicus: ‘Kalahari-Wüste’ [Kalahari Desert]; C. dis-

septus: ‘Transvaal (Potchefstroom), Southern Rhodesia (between Lim-
popo and Zambesi Rivers)’ [North-West Province, South Africa and W
Zimbabwe].
Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size, removal of syn-
onym pending. ♂
Distribution: Deep Kalahari sands: Namibia, Botswana, E Zimbabwe,
central N and NW South Africa.
Locality data (mean ± SD, range): Altitude: 1 062 ± 158, 581–1 689 m;
annual rainfall: 329 ± 115, 157–659 mm; annual temperature (max. +
min. /2): 19.9 ± 1.2, 17.7–23.0°C (N=152).
Habitat: No quantitative assessment; DBRU collection records primarily
from deep sand (42), few from sandy loam (2); recorded in grassland
(10), scrub/shrubland (17) and open woodland (14).
Food types: In Botswana: primarily sampled from omnivore: pig (84),
and monogastric herbivore dung: elephant (62); few sampled from
ruminant herbivore dung: sheep (5), cattle (0) or carrion (0); DBRU
collection records from dung of elephant (3), donkey (4), horse (1),
cattle (33), buffalo (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: Centred on Kalahari Xeric
Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Kalahari Duneveld (SVkd), East Kalahari
Bushveld (SVk), NW Central Bushveld (SVcb), NW Mopane (SVmp) 5 mm
(Savanna Biome).
Assessment rationale: EOO = 639 850 km2; widespread, centred in an
area used primarily for the grazing of domestic livestock from whose
dung it has been widely recorded despite a quantitative bias to the dung
of pig and elephant; transformation across its Northern Cape, South
African range, only 0–2% (SVkd, SVk); assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations and further data on
food associations; widely represented in farm rangeland and protected
in Chobe National Park, Central Kalahari Game Reserve and Kgalagadi
Transfrontier Park (Botswana, South Africa).

♀ 5 mm

COPRINI
170 SURICATA 6 (2020)

Catharsius harpagus
Harold, 1877b (but see Taxonomy)

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: ‘Afric. Austral’ [southern Africa], lectotype: ‘Delagoa Bay’

[Maputo, S Mozambique].
Taxonomy: Accepted species, but name change pending; sexually dimor-
phic (head and prothoracic disc), prominence of characters varies with
body size.
Distribution: Deep coastal sands of Maputaland Centre of Endemism;
♂ NE KwaZulu-Natal, South Africa; SE Mozambique.
Locality data (mean ± SD, range): Altitude: 28 ± 36, 0–117 m; annual
rainfall: 801 ± 103, 661–971 mm; annual temperature (max. + min.
/2): 22.2 ± 0.6, 21.3–23.6°C (N=12).
Habitat: No quantitative assessment of soil type; only two DBRU collec-
tion records on sand, but certainly a sand specialist; on sand in Maputo
Special Reserve: restricted to grassland (192), forest (0).
Food types: No quantitative assessment; DBRU collection records on
cattle dung (2); sampled in large numbers to pig dung.
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season (Oct. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT1119).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB); also
margins of Lowveld (SVl) in adjoining Savanna Biome.
Assessment rationale: EOO = 8 250 km2; difficult to assess conservation
status in the absence of survey data to the N along the Mozambique
coast or the absence of clear quantitative support for soil specialisation
on coastal sands; assessed as Data Deficient (DD) although could equal-
ly deserve Least Concern (LC) status as part of the range is protected;
may even deserve a threat category as the known EOO is < 20 000 km2
and the AOO could be < 5 000 km2; apparently centred on grassland
5 mm
patches within a forest/woodland matrix.
Conservation measures: Assessment of conservation status would be im-
proved by a survey to determine soil and food associations, the number
of occupied grassland patches, and population density; protected in
Tembe Elephant Park and in iSimangaliso Wetland Park (World Her-
itage Site) (South Africa) plus Maputo Special Reserve (Mozambique)
where there are a number of natural grassland patches on fossil lagoons.

♀
5 mm

COPRINI
SURICATA 6 (2020) 171

Catharsius heros
Boheman, 1860

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘juxta lacum N’Gami’ [near Lake Ngami, N. Botswana].

Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
Distribution: Dry savanna from S to E Africa: South Africa, Namibia,
Botswana, Zimbabwe, Angola, Mozambique, Kenya; occurrence now
patchy due to dung type specialisation; most records outside of reserves
older than 50 yr. ♂
Locality data (mean ± SD, range): Altitude: 895 ± 368, 43–1 377 m;
annual rainfall: 538 ± 169, 155–1 016 mm; annual temperature (max.
+ min. /2): 21.3 ± 2.0, 17.1–23.2°C (N=32).
Habitat: No quantitative assessment; most DBRU collection records from
game reserves (15) or nearby localities (2); one record from a stray
donkey holding pen (Kuruman); sandy soil, woody vegetation bias:
sand (2), sandy loam (7), sandy clay loam (1) in open woodland (7),
grassland (2).
Food types: In Botswana: bias to monogastric herbivore dung compared
to other bait types: carrion (0), pig (2), elephant (18), cattle (0), sheep
(0); all DBRU collection records from dung of large monogastric herbi-
vores now found primarily in game reserves: elephant (12), rhinoceros
(1), donkey (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna Wood-
lands (AT1316), Zambezian and Mopane Woodlands (AT0725), An-
golan Mopane Woodlands (AT0702), Kalahari Acacia-Baikiaea Wood- 5 mm
lands (AT0709), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bush-
veld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 3 687 200 km2; AOO now much smaller,
largely equivalent to the area of game reserves in dry savanna contain-
ing elephants and rhinoceros between southern and East Africa; one
small-bodied individual recorded on donkey dung near Kuruman indi-
cates that C. heros may also be present in very low population density
where equids occur outside of reserves; currently assessed as Least Con-
cern (LC) owing to protection in various reserves over a wide range.
Conservation measures: Conservation status largely dependent on con-
tinuing protection of elephants and rhinoceros in game reserves and
national parks, including Kruger, Ndumu (South Africa), Chobe, Cen-
tral Kalahari (Botswana), Etosha (Namibia), Bicuar (Angola), Hwange
(Zimbabwe), Tsavo West (Kenya).

♀ 5 mm

COPRINI
172 SURICATA 6 (2020)

Catharsius laticeps
Boheman, 1857 (but see Taxonomy)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘prope fluvium Limpopo’ [close to Limpopo River, south-

ern Africa].
Taxonomy: Subsequent to completion of this book, H. Takano (pers.
comm.) pointed out that the true C. laticeps is a species of coastal for-
est on deep sands of NE KwaZulu-Natal and SE Mozambique (see
Preamble) and is roughly sympatric with C. pandion Harold, 1877b;
♂ this wrongly-named species should be compared with C. parafastidiosus
Ferreira, 1971, described from an unknown locality purported to be in
South Africa; sexually dimorphic (head and prothoracic disc), promi-
nence of characters varies with body size.
Distribution: Dry, warm, lowland savanna: S Zimbabwe, NE South Afri-
ca; also cited from Mozambique.
Locality data (mean ± SD, range): Altitude: 308 ± 209, 55–495 m; an-
nual rainfall: 513 ± 96, 395–629 mm; annual temperature (max. +
min. /2): 22.8 ± 0.2, 22.7–23.1°C (N=4).
Habitat: No quantitative assessment; one DBRU collection record on
sandy clay loam in shrubland.
Food types: No quantitative assessment; limited DBRU collection records
on the dung of elephant (1), cattle (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov., Mar.).
Ecoregions Zimbabwe: margins of Zambezian and Mopane Woodlands
(AT0725) and Southern African Bushveld (AT0717).
Bioregions South Africa: Mopane (SVmp), Lowveld (SVl) (Savanna
Biome).
Assessment rationale: EOO = 15 750 km2 (probably underestimated);
5 mm not well represented in reference collections and poorly known ecolog-
ically; AOO possibly biased to finer-grained soils in open woody vege-
tation, but quantitative support required; insufficient data to determine
if there is a bias to the dung of monogastric herbivores, which would in-
fluence conservation status; currently assessed as Data Deficient (DD).
Conservation measures: Before an assessment of conservation status can
be made, a quantitative survey is required in drier savanna along the
Limpopo River Valley and on the S Mozambique Coastal Plain to de-
termine the full EOO, AOO and ecological associations; validation of
the species name also required; protected in Kruger National Park and
Ndumo Game Reserve (South Africa).

5 mm ♀

COPRINI
SURICATA 6 (2020) 173

Catharsius longiceps
Gillet, 1907

= Catharsius melancholicus sensu Péringuey, 1901 (?pars)

Catharsius vansoni
Ferreira, 1960a (but see Taxonomy)

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: C. longiceps: ‘Transvaal: Boksburg’ [Boksburg, Gauteng, ♂

South Africa]; C. melancholicus sensu Péringuey: ‘Natal (Estcourt,
Durban), Transvaal (Boksburg, Lydenburg). Southern Rhodesia (Bu-
luwayo, Salisbury)’ [South Africa/Zimbabwe range possibly represents
more than a single species]; C. vansoni: ‘Transvaal: Potchefstroom’
[Potchefstroom. North West Province, South Africa].
Taxonomy: Both C. longiceps and C. vansoni are, currently, accepted spe-
cies; synonymy of C. vansoni pending publication; sexually dimorphic
(head and prothoracic disc), prominence of characters varies with body
size.
Distribution: Uncertain; recent samples localised near two type localities
on NW Highveld, South Africa: Boksburg (C. longiceps) (blue square),
Potchefstroom (C. vansoni) (red squares); old C. vansoni reference ma-
terial from both non-validated Highveld and coastal localities (black
squares); no material seen from Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 470 ± 68, 1 356–1 615 m;
annual rainfall: 657 ± 28, 619–714 mm; annual temperature (max. +
min. /2): 16.8 ± 0.3, 16.4–17.3°C (N=10); only for localities represent-
ed by red squares.
Habitat: In Abe Bailey Nature Reserve: sampled from sandy loam primar-
5 mm
ily in grassland (50) compared to open woodland (2).
Food types: No quantitative assessment; sampled to cattle dung and com-
posite baits of pig and cattle dung.
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Nov. to May).
Bioregions South Africa: Straddling NE and NW edges of Dry High-
veld Grassland (Gh) and Mesic Highveld Grassland (Gm) (Grassland
Biome) (red squares).
Assessment rationale: EOO = 6 130 km2 (range for red squares only,
presumably an underestimate); difficult to assess given the confused
and meagre data; very small validated range in an area of North West
Province where vegetation units are 25–50% transformed; however,
presumably more widely distributed despite the absence of recent re-
cords elsewhere in southern Africa (previous 50 yrs); currently assessed
as Data Deficient (DD).
Conservation measures: A survey is required to provide validated data on
the EOO, AOO and ecological associations, and to generate a balanced
assessment of conservation status; an assessment of the possible effects
of habitat transformation would also be useful; currently known to be
protected only in Abe Bailey Nature Reserve (South Africa).

♀
5 mm

COPRINI
174 SURICATA 6 (2020)

Catharsius marcellus
Kolbe, 1893a

No synonyms
Global: LC
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type locality: ‘In Zoutpansberg bei Mphôme in Nord-Transvaal’

[Mphome Mission Station, Haenertsburg, Limpopo Province, South
Africa].
Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
♂ Distribution: Highveld Grassland: South Africa, Eswatini.
Locality data (mean ± SD, range): Altitude: 1 533 ± 383, 1 142–3 061 m;
annual rainfall: 802 ± 127, 450–1 157 mm; annual temperature (max.
+ min. /2): 15.2 ± 2.3, 13.5–16.6°C (N=37).
Habitat: No quantitative assessment of soil type; DBRU collection re-
cords primarily from finer-grained soils: sandy clay loam (5), sandy
loam (3), sand (1), all in pasture/grassland (8); bias to cooler grassland
at Suikerbosrand Nature Reserve: montane grassland (31), Highveld
Grassland (20), open woodland (5); at Carolina, little measured re-
sponse to disturbance: natural Themeda grassland (44), fallow crop field
(10), improved Kikuyu pasture (38).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (10).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Nov. to May).
Bioregions South Africa: Centred on cooler moister vegetation units in
the E Mesic Highveld Grassland (Gm), N Drakensberg Grassland (Gd)
and N Sub-Escarpment Grassland (Gs); scattered records in W uplands
of Gm and E Dry Highveld Grassland (Gh).
Assessment rationale: EOO = 126 300 km2; widespread; readily attracted
to cattle dung on the South African Highveld where it is still frequently
recorded; available data suggest limited threats from pasture improve-
5 mm
ment and transformation, but most records appear to be from collec-
tions or quantitative studies conducted in natural grassland; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; in particular,
the effects of pasture improvement and transformation should be re-
assessed to ensure that data from disturbed habitats at Carolina were not
influenced by immigration from nearby natural grassland; protected in
various reserves including Suikerbosrand, Ithala, Royal Natal, Lotheni.

♀
5 mm

COPRINI
SURICATA 6 (2020) 175

Catharsius melancholicus
Boheman, 1860

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘prope lacum N’Gami’ [close to Lake Ngami, N Botswana].

Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc),
prominence of characters varies with body size.
Distribution: Deep sands of the Kalahari and its outliers: NW South
Africa, Botswana, Namibia; also reported from Angola, Democratic
Republic of the Congo (DRC); report from Mozambique presumably
an error. ♂
Locality data (mean ± SD, range): Altitude: 1 050 ± 108, 871–1 246 m;
annual rainfall: 338 ± 137, 91–659 mm; annual temperature (max. +
min. /2): 20.0 ± 1.3, 16.9–22.4°C (N=45).
Habitat: No quantitative assessment; sampled only from study sites on
deep sand (29) in Botswana and Northern Cape, South Africa; DBRU
collection records only on deep sand (3) in shrubland (3).
Food types: In Botswana: sampled mainly from carrion (chicken livers:
297), few on four dung types, pig (38), elephant (6), cattle (6), sheep
(1); DBRU collection records only from a dead owl (1) and rotten goat
meat (2).
Temporal activity: Flight activity during darkness in the summer rainy
season (Dec. to Mar.).
Ecoregions Namibia, Botswana: Kalahari Xeric Savanna (AT1302),
Namibian Savanna Woodlands (AT1316), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Kalahari Duneveld (SVkd), East Kalahari
Bushveld (SVk) and sand outliers in Central Bushveld (SVcb) (Savanna
Biome).
5 mm
Assessment rationale: EOO = 970 820 km2 (southern Africa only); SW
half of EOO little transformed (0–2%, SVkd, E SVk) as land usage is
dominated by grazing of domestic livestock; although, AOO would be
about 30% smaller than EOO due to sand specialisation, food would
be available for this carrion specialist throughout its range of which at
least 10% is under protection; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations; protected in
Chobe National Park, Central Kalahari Game Reserve and Kgalagadi
Transfrontier Park (Botswana, South Africa).

♀ 5 mm

COPRINI
176 SURICATA 6 (2020)

Catharsius pandion
Harold, 1877b

= Catharsius mossambicanus Ferreira, 1960a

Global: LC

J A S O N D J F M A M J

Type localities: ‘Port Natal’ [Durban, KwaZulu-Natal, South Africa], lecto-

type: ‘Natal’ [KwaZulu-Natal, South Africa]; C. mossambicanus: ‘Provín-
cia de Moçambique: Lourenço Marques’ [Maputo, Mozambique].
Taxonomy: Accepted species; C. pandion Harold was recently shown to be
the senior name for C. mossambicanus Ferreira (Génier & Josso 2016)
after having been assigned to a close, hitherto undescribed relative (now
♂ C. aegeus Génier, 2017); sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
Distribution: Coastal range in KwaZulu-Natal, South Africa, and SE Mo-
zambique, including the Maputaland Centre of Endemism.
Locality data (mean ± SD, range): Altitude: 43 ± 140, 0–991 m; annual
rainfall: 853 ± 111, 646–1 010 mm; annual temperature (max. + min.
/2): 21.9 ± 0.8, 17.7–23.3°C (N=52).
Habitat: In uMkhuze Game Reserve: entirely on sand (19), sandy clay
loam and clay (0) (cited as Catharsius near pandion); on deep sand in
Maputo Special Reserve, primarily in forest: coastal dune forest (15.8),
inland sand forest patches (large: 13.8, medium: 9.9, small: 4.5),
grassland (0.6); across a gradient of rehabilitating vegetation on coastal
dunes at Richards Bay: grassland (68), dense early sucession woodland
(1 135), open understorey later succession woodland (714), dense nat-
ural forest (1 072); largely supported by DBRU collection records: all
on sand (8) in forest (15) or grassland (6).
Food types: No quantitative assessment; DBRU collection records from
omnivore: human (4), monogastric: elephant (1) and ruminant herbi-
vore dung: cattle (3).
Temporal activity: Flight activity in darkness, primarily during the sum-
5 mm mer rainy season (Oct. to Jan.), but also cooler dry season months on
the warm coastline (Aug., May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome, adjoining
edge of Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 18 650 km2; occupies a limited coastal
range where AOO is primarily restricted to Sand Forest (FOz 8) and
Dune Forest; population density negatively influenced by small FOz 8
patch size; percentage transformation of FOz 8 unclear in South Africa,
but listed as Critically Endangered due to clearance, firewood collection
and elephant damage; Dune Forest threatened by mining of coastal
dunes at S end of range although 30% of the concession is managed for
post-mining vegetative restoration; protected across ca. 20% of known
species range, therefore, currently assessed as Least Concern (LC) rather
than Near Threatened (NT).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on food associations; as a forest specialist
on range-restricted deep coastal sands, conservation status of this spe-
cies is dependent on the continuing protection of large sand and dune
forest patches in and around the Indian Ocean Coastal Belt; protected
in iSimangaliso Wetland Park (World Heritage Site), Tembe Elephant
Park (South Africa) and Maputo Special Reserve (Mozambique).
♀
5 mm

COPRINI
SURICATA 6 (2020) 177

Catharsius philus
Kolbe, 1893a (but see Taxonomy)

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Senna in Mosambik…Sansibar.’ [Mozambique: Sena;

Tanzania: Zanzibar].
Taxonomy: Accepted species but name change and designation of a syn-
onym pending; sexually dimorphic (head and prothoracic disc), prom-
inence of characters varies with body size.
Distribution: Dry SE African savanna on finer-grained soils: N South
Africa, E Botswana, SE and N Zimbabwe, Mozambique, E Tanzania; ♂
doubtful locality (black square).
Locality data (mean ± SD, range): Altitude: 783 ± 407, 48–1 850 m;
annual rainfall: 617 ± 158, 326–1 136 mm; annual temperature (max.
+ min. /2): 20.5 ± 2.1, 13.4–25.7°C (N=94).
Habitat: On Gauteng bushveld: restricted to finer-grained soils: sandy clay
loam (37), deep sand (0) with a bias to grassland (28) rather than open
woodland (9) or shaded tickets (0); DBRU collection records indicate
a bias to finer-grained soils: sandy clay loam (9), sandy loam (24), sand
(8), and open woodland (25) as opposed to grassland/pasture (7) or
even shrubland (5).
Food types: At Phalaborwa: sampling bias to dung of cattle (57) and pig
(32), fewer on elephant (7); DBRU collection records primarily from
ruminant herbivore dung: cattle (35), buffalo (1), wildebeest (2); few
from monogastric herbivore dung: elephant (2), rhinoceros (1), horse
(2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); on Gauteng bushveld: activity commences early
summer (Oct., Nov.) peaks in mid-season (Dec., Jan.) and tails off in
late summer (Feb. to Apr.); at Phalaborwa: much more abundant on
a warm evening soon after very heavy rainfall (88) than on evenings
following warm cloudy conditions and light rainfall (4) or hot, dry 5 mm

conditions (3).
Ecoregions Botswana, Zimbabwe: Southern African Bushveld (AT0717),
S Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Loweld (SVl) (Savanna Biome).
Assessment rationale: EOO = 774 500 km2 (gross estimate); AOO prob-
ably smaller due to finer-grained soil bias in drier savanna; possible
effect of woodland clearance difficult to assess owing to conflicting re-
sults; however, widespread occurrence on cattle dung in farm rangeland
suggests few current threats; transformation in South African range,
0–58% (SVl), 2–49% (SVcb), 0–20% (SVmp); assessed as Least Con-
cern (LC).
Conservation measures: Quantitative data on effects of woodland clear-
ance on finer-grained soils would improve the assessment of conser-
vation status; protected in Kruger National Park, Hluhluwe–iMfolozi
Game Reserve, Tswaing Nature Reserve (South Africa).

♀
5 mm

COPRINI
178 SURICATA 6 (2020)

Catharsius platycerus
(Klug, 1855) (but see Taxonomy)

= Catharsius obtusicornis Boheman, 1857

= Catharsius gibbicollis Gerstaecker, 1884
= Catharsius princeps Kolbe, 1893a
Global: DD

J A S O N D J F M A M J

Type localities: As Copris platycera: ‘Sena’ [Mozambique]; C. obtusicornis:

‘Caffraria interiore’ [interior of SE South Africa]; C. gibbicollis: ‘Massai’
[East Africa]; C. princeps: ‘Stanley Pool’ [Democratic Republic of the
Congo (DRC)].
♂ Taxonomy: Accepted species, but change to validity of names and removal
of some synonyms pending; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
Distribution: Dry to moist savanna from southern to East Africa, but
very patchy occurrence due to dung type specialisation: South Africa,
Zimbabwe, Mozambique, Democratic Republic of the Congo (DRC),
Kenya; also reported from Namibia, Rwanda, Ethiopia.
Locality data (mean ± SD, range): Altitude: 568 ± 435, 59–1 106 m;
annual rainfall: 704 ± 192, 529–1 136 mm; annual temperature (max.
+ min. /2): 23.2 ± 2.4, 19.8–25.9°C (N=8).
Habitat: No quantitative assessment; limited DBRU collection records
(6) inconclusive: primarily from game reserves (5) on sand (1), sandy
loam (1), sandy clay loam (2) in grassland (1), open woodland (3),
forest (1).
Food types: No adequate quantitative assessment; at Phalaborwa: one
specimen trapped to elephant rather than pig or cattle dung; limited
DBRU collection records suggest bias to monogastric herbivore dung:
rhinoceros (2), elephant (2), cattle (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.); attracted to light.
5 mm Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725),
Zambezian Baikiaea Woodlands (AT0726), Southern Miombo Wood-
lands (AT0719).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 3 012 810 km2 (gross estimate); widespread,
but rarely recorded; ecological data severely limited, but recent records
primarily from game reserves of S and E Africa either on monogastric
herbivore dung or attracted to light; does not qualify for threat status
owing to large range and occurrence in many game reserves, but pop-
ulation density apparently very low; assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and population
density; observations suggest that protection of elephant and rhinocer-
os in reserves is essential for the conservation of this species; protected
in reserves from S to E Africa, including Meru, Masai Mara, Tsavo East
(Kenya), Hwange (Zimbabwe), Gorongosa (Mozambique), Kruger
(South Africa).

5 mm
♀

COPRINI
SURICATA 6 (2020) 179

Catharsius sesostris
Waterhouse, 1888 (but see Taxonomy)

= Copris pithecius Olivier, 1790

= Catharsius pylades Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: C. sesostris: ‘Egypt.’; C. pithecius: ‘Sénégal’ [Senegal; also

reported from Egypt and East Indies by Olivier!]; C. pylades: ‘Natal
(Durban, Frere), Southern Rhodesia (Salisbury)’ [KwaZulu-Natal,
South Africa; Harare, Zimbabwe].
Taxonomy: Accepted species, but revalidation of one synonym and re-
moval of all other species names pending; sexually dimorphic (head ♂
and prothoracic disc), prominence of characters varies with body size.
Distribution: Range reported to extend across West Africa (Senegal, Gam-
bia, Cameroon), along E seaboard from South Africa to Egypt and into
Middle East (Palestine, Syria, Jordan, Arabia); mapped taxon validated
for NE South Africa, Zimbabwe, Zambia, Mozambique, Kenya; report
from Namibia (Okahandja) requires validation.
Locality data (mean ± SD, range): Altitude: 993 ± 449, 1–1 958 m; an-
nual rainfall: 757 ± 180, 383–1 415 mm; annual temperature (max. +
min. /2): 19.3 ± 2.3, 14.3–25.9°C (N=96).
Habitat: On Gauteng bushveld: extreme sample bias to sand and strong
bias to shaded situations, deep sand (48), sandy clay loam (0), grass-
land (1), open woodland (10), shaded thickets (37); DBRU collection
records differ: sandy clay loam (5), sandy loam (10), sand (3) in grass-
land/pasture (9), shrub/woodland (3), forest (5).
Food types: On Gauteng bushveld: very strong sample bias to carrion
(15) and omnivore dung pig (52), few on dung of horse (4), cattle (1);
DBRU collection records differ: cattle (16), rhinoceros (3), horse (1),
baboon (1), human/ruminant dung mixture (6).
Temporal activity: Flight activity in darkness during the summer rainy 5 mm

season (Oct. to Apr.); on Gauteng bushveld activity spread evenly

throughout summer.
Ecoregions Zimbabwe: Southern African Bushveld (AT0717), Southern
Miombo Woodlands (AT0719); marginal in Zambezian and Mopane
Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl)
(Savanna Biome); Indian Ocean Coastal Belt Biome (CB); marginal
occurrence in Mopane (SVmp), E Mesic Highveld Grassland (Gm).
Assessment rationale: EOO = 5 293 770 km2 (C. sesostris underestimat-
ed); EOO = 1 586 240 km2 (mapped taxon); difficult to assess given
questions of taxon identity, range and conflicting ecological data, but
probably biased to omnivore dung in shady moist savanna, possibly on
sandier soils; transformation in South Africa: 0–58% (SVl), 2–49%
(SVcb), but not threatened at present due to widespread occurrence
from S to E Africa and protection in many reserves; assessed as Least
Concern (LC).
Conservation measures: Taxonomic issues concerning species identity
need to be resolved; quantitative data suggest that protection of dense,
moist, savanna woodland on sandy soils would be advantageous for
the conservation of the taxon treated, here, as C. sesostris; protected
in various reserves including, Kruger (South Africa), Lake Mutirikwe
(Zimbabwe), Meru, Marsabit (Kenya).
♀
5 mm

COPRINI
180 SURICATA 6 (2020)

Catharsius tricornutus
(DeGeer, 1778)

= Scarabaeus nemestrinus Fabricius, 1781

= Catharsius areolatus Boheman, 1857
= Catharsius dubius Péringuey, 1901 (but see Taxonomy)

Global: LC

J A S O N D J F M A M J

Type localities: As Scarabaeus tricornutus: Not stated; S. nemestrinus:

‘Cap bon sp.’ [Cape of Good Hope, South Africa]; C. areolatus: ‘juxta
fluvium Gariep’ [near Orange River, South Africa]; C. dubius: ‘Trans-
♂ vaal (Rustenburg), Southern Rhodesia (Zambesi River), Ovampoland
(Okovango River)’ [South Africa, Zimbabwe, Namibia].
Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size; removal of one
synonym pending.
Distribution: Moist savanna and grasslands of southern Africa: S coastal
to NE South Africa, Zimbabwe, Mozambique, N Botswana, N Namib-
ia, S Angola; reports from Democratic Republic of the Congo (DRC),
Zambia, Tanzania and Kenya require validation.
Locality data (mean ± SD, range): Altitude: 873 ± 549, 0–1 958 m; an-
nual rainfall: 701 ± 204, 201–1 415 mm; annual temperature (max. +
min. /2): 18.9 ± 2.4, 13.1–24.8°C (N=205).
Habitat: On Gauteng bushveld: extreme association with deep sand, but
relatively generalist vegetation associations, deep sand (778), sandy
clay loam (1), grassland (246), open woodland (353), shaded thickets
(180); similar soil association in uMkhuze Game Reserve: sand (31),
sandy clay loam/clay (0); partly supported by DBRU collection records:
coarse-grained soil bias, but vegetation bias to grassland, clay (3), sandy
clay loam (13), sandy loam (33), sand (43) in grassland/pasture (55),
shrubland (8), open woodland (17); also crop fields (7).
Food types: On Gauteng bushveld: bias to dung of omnivores compared
to monogastric and ruminant herbivores: pig (60), horse (24), cattle
5 mm (13); but DBRU collection records primarily on cattle dung (96); also
buffalo (1), elephant (4), horse (2), zebra (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to May); on Gauteng bushveld: seasonal peak in early
summer (Oct. to Jan.), tailing off to late summer (Feb. to Apr.); in
mid-summer, flight activity peak at dusk (18:00–20:00) declining
thereafter, but may fly sporadically throughout the night, particularly
after showers; most Western and Eastern Cape records (11 out of 15) in
spring (Oct.) or autumn (Mar. to May); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Maputa-
land Coastal Forest Mosaic (AT0119), Southern Zanzibar-Inhambane
Coastal Forest Mosaic (AT0128), Southern African Bushveld (AT0717),
Southern Miombo Woodlands (AT0719), Zambezian Baikiaea Wood-
lands (AT0726); Angolan Mopane Woodlands (AT0702).
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bushveld
(SVcb), Lowveld (SVl) (Savanna Biome); moister, warmer, N and E
Grassveld Biome: N Dry Highveld Grassland (Gh), E Mesic Highveld
Grassland (Gm), Sub-Escarpment Grassland (Gs); also Indian Ocean
Coastal Belt, Albany Thicket and Fynbos Biomes.

COPRINI
SURICATA 6 (2020) 181

Assessment rationale: EOO = 1 064 120 km2 (possibly underestimated);

AOO smaller due to soil type bias; nevertheless widespread in open
vegetation and readily attracted to cattle dung in moist farm range-
land despite a bias to omnivore dung association; apparently adaptable ♀
to habitat disturbance as all 11 collection records in the Western and
Eastern Cape were either in fallow crop fields (4) or grassland (7), sug-
gesting range expansion from the summer rainfall region in response to
clearance of natural shrubland; assessed as Least Concern (LC).
Conservation measures: None required owing to adaptability across a
widespread range in farmland as well as reserves; protected in Hluhlu-
we–iMfolozi Game Reserve and various nature reserves, including
Tswaing, Leeuwfontein, Roodeplaat (South Africa).

5 mm

COPRINI
182 SURICATA 6 (2020)

Catharsius ulysses
Boheman, 1857

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Caffraria tota’ [all SE South Africa – undoubtedly inexact

usage of ‘Caffraria’].
Taxonomy: Accepted species; sexually dimorphic (head and prothoracic
disc), prominence of characters varies with body size.
Distribution: Disjunct; centred on dry upland savanna or grassland: NW
♂ Highveld, South Africa, SE Botswana; N Namibian Plateau; report
from Uganda presumably an error.
Locality data (mean ± SD, range): Altitude: 1 307 ± 253, 757–1 928 m;
annual rainfall: 454 ± 130, 179–738 mm; annual temperature (max. +
min. /2): 18.9 ± 1.5, 15.8–21.8°C (N=70).
Habitat: No quantitative assessment; DBRU collection records suggest a
bias to sandier soils in open woody vegetation; clay (2), sandy clay loam
(4), sandy loam (10), sand (21) in grassland (7), open shrubland (11),
open woodland (17).
Food types: No quantitative assessment; DBRU collection records pri-
marily from ruminant herbivore dung: cattle (35); monogastric herbi-
vore dung: donkey (1).
Temporal activity: Flight activity during darkness, primarily in the sum-
mer rainy season (Nov. to May); attracted to light.
Ecoregions Namibia, Botswana: NW & SE Kalahari Xeric Savanna
(AT1302), margins of three peripheral ecoregions.
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bush-
veld (SVcb) (Savanna Biome), N Dry Highveld Grassland (Gh) (Grass-
land Biome).
Assessment rationale: EOO = 367 200 km2; widespread; in South Afri-
ca: E extreme of range little transformed (0–2%, E SVk), remainder
5 mm variously transformed (13–42%, W SVk; 10–63%, N Gd; 14–49%,
upland SVcb); however, readily attracted to cattle dung in fairly open
vegetation in farm rangeland of both South Africa and Namibia, al-
though few records from nature reserves; somewhat Data Deficient
(DD) but assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and population
density in nature reserves versus adjoining farm rangeland; only known
to be protected in Rustenburg, Benfontein and Dronfield nature re-
serves (South Africa).

♀
5 mm

COPRINI
SURICATA 6 (2020) 183

Catharsius vitulus
Boheman, 1857 (but see Taxonomy)

= Catharsius camillus Harold, 1877b

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: C. vitulus: ‘Caffraria interiore’ [interior of SE South Afri-

ca]; C. camillus: ‘Port Natal’ [Durban, South Africa, but probably a lo-
cality error], lectotype: ‘cap b. sp.’ [Cape of Good Hope, South Africa].
Taxonomy: Accepted species, but changes to validity of name and syn-
onym pending; sexually dimorphic (head and prothoracic disc), prom-
inence of characters varies with body size. ♂
Distribution: Dry to arid, W Highveld and Upper Karoo, South Africa;
Zimbabwe record requires validation; those from Democratic Republic
of the Congo, Kenya and E coast of South Africa (black square: C. ca-
millus) presumably errors.
Locality data (mean ± SD, range): Altitude: 1 385 ± 103, 1 151–1 632 m;
annual rainfall: 404 ± 139, 266–676 mm; annual temperature (max. +
min. /2): 15.3 ± 1.0, 13.4–18.3°C (N=40). Durban (black square) not
included: 0.6 m, 955 mm, 20.8°C.
Habitat: No quantitative assessment; Upper Karoo samples from sand
(14) or sandy loam (11) in grassland (14) or Karoo shrubland (13);
limited DBRU collection records inconclusive: clay (1), sandy loam
(3), sand (1) in pasture (3) and open woodland (1).
Food types: No quantitative assessment; DBRU collection records only
from cattle dung (6).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May).
Bioregions South Africa: Primarily E Upper Karoo (NKu) (Nama Karoo
Biome); Dry Highveld Grassland (Gh) (Grassland Biome); marginal
records in Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland
5 mm
(Gs) (Grassland Biome).
Assessment rationale: EOO = 92 325 km2 (minus Durban); patchy
distribution possibly a collecting artefact, but may be related to some
unidentified specialisation; SW part of range coincides with areas that
have been little transformed; NE part of range 4–63% transformed
(Gh); somewhat Data Deficient (DD), but assessed as Least Concern
(LC) on basis of widespread range, 30% of which coincides with little
modified regions in the Upper Karoo.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and the possible
effects of modification of natural vegetation by pasture improvement;
no available localities occurred in protected areas.

♀ 5 mm

COPRINI
184 SURICATA 6 (2020)

Genus Copris Geoffroy, 1762

Type species and designation: Scarabaeus lunaris Linnaeus, 1758, by subsequent designation (Latreille 1810).
Synonyms: None.
Last review: Selected Afrotropical groups revised by Nguyen-Phung and Cambefort (1986a, 1986b,
1987), Nguyen-Phung (1987, 1988a, 1988b, 1988c, 1989) and Cambefort (1992a,
1992b); phylogeny of entire Afrotropical Copris fauna (Cambefort & Nguyen-Phung
1996); partial reappraisal of Copris and allied genera (Marchisio & Zunino 2012); new
species (Deschodt et al. 2015a).

The long-stablished genus, Copris Geoffroy, 1762, is widely distributed across climatically suitable areas. It is represent-
ed by a total of 346 species, some classified under subgeneric names that are variously considered to have generic status
by some authors, or, to be invalid by others (Copris Geoffroy, 1762; Paracopris Balthasar, 1939a; Microcopris Balthasar,
1958; Sinocopris Ochi, Kon & Bai, 2009). Copris species have been recorded from the Afrotropical (108), Palaearctic
(13), Oriental (99), Palaearctic + Oriental (1) and Nearctic (19) regions as well as the extreme NW of the Neotropical
region (16).

The Afrotropical members of the genus have been divided into 48 species groups with 24 represented in the area encom-
passing South Africa, Botswana and Namibia. A total of 33 currently valid species is found in one or more of these three
countries, although one is probably a synonym (see Copris misellus Péringuey under C. obesus Boheman). Another is rep-
resented by Kalahari and E coast subspecies. At least one undescribed species is also known.

Copris species are relatively large bodied although different groups show trends to either larger or smaller body size. Dung
is buried at the ends of tunnels leading from under droppings. A number of species have been recorded to first sequester
dung at the end of a shallow tunnels and then remove it to a deeper tunnel where separate brood ovoids are cut from a
dung cake within a chamber. This nest architecture requires fairly large amounts of dung. Therefore, they are found pri-
marily in moister regions that support large herbivores dropping large amounts of dung.

Copris species are mostly macropterous, flying during darkness, although a few species from cooler highland or southern
regions may be diurnal fliers (Groups 17–18) or even apterous and flightless (Group 16). Of 32 valid species in southern
Africa, 26 are restricted to the region with only six showing larger, more widespread ranges into the tropics.

Owing to patterns of relationships, it has been suggested that the history of the genus in Africa is concentrated on savanna
with occasional local radiations into mountains and forest. However, in southern Africa, less than half of the species show
ranges centred on northern savanna, either in the east (8), the west (3) or both (1). Others are centred on the Kalahari
sands (2 + a subsp.) or across Highveld Grassland (3), possibly with a bias to the N (1) or the E escarpment (3). A number
are southern endemics in the winter and bimodal regions, either along the W coast (1), S coast (4) or both (1). Others
occur along the summer rainfall E coastal zone of South Africa (1), in Maputaland (1 + a subsp.), or northwards and up
the Zambezi Valley (1). One shows a composite range along the S and E coasts and E escarpment. None are found in SW
Arid regions.

Most Copris species are assessed as Least Concern (LC), since they are widespread in farmland and reserves (23) or are cur-
rently well protected in reserves along with elephants (1), although relatively few species appear to be coarse or omnivore
dung specialists. However, several are assessed as Data Deficient (DD: 5), Near Threatened (NT: 1) or even Endangered
(EN: 2). These eight species are poorly known ecologically and seven have been recorded at few localities. The three
threatened species are endemic to the southern Cape where there has been extreme transformation of the natural fynbos
or Renosterveld shrubland at lower altitude.

The system of groups suggested by Cambefort and Nguyen-Phung (1996) is used here, as it covers the entire Afrotropi-
cal fauna and is based on a morphological phylogeny. The patterns seem to describe serial radiations in different groups,
some representing radiation into the same regions. There seems to be a bias to occurrence on herbivore dung, both that
of monogastrics and ruminants.

COPRINI
SURICATA 6 (2020) 185

In southern Africa:
Groups 1–3: Three species with basal phylogenetic derivation; two with eastern savanna associations in southern
Africa, one primarily found in shady forest situations along the southern and E coasts as well as the E
escarpment of South Africa.
Group 4: One monogastric herbivore dung specialist, with basal phylogenetic derivation, distributed in savannas
from southern to E Africa.
Group 11: One species centred on deep sands of the southern Kalahari.

Major clade, mostly larger-bodied, more terminally derived.

Group 12: One species with two subspecies; both centred on deep sands in the southern Kalahari or Maputaland
(NE coast South Africa).
Group 16: Two diurnally active, apterous, flightless species from the southern coast of South Africa; both assessed
as at least EN owing to rarity of records from a highly transformed region.
Groups 17–18: Four species centred on cool moist or dry areas of the Highveld (3 spp.) or the highlands bordering the
S coast, South Africa (1 sp.); possibly all showing diurnal flight; three species assessed as DD.
Groups 19–20: Three species centred on upland savanna in N Namibia (1 sp.) or Highveld Grassland (1 sp.) and S coast
(1 sp.), South Africa.
Groups 21–22: Two species with distributions centred on the winter or winter and bimodal rainfall regions of South
Africa.
Group 23: One savanna species widespread to East Africa.
Group 26: Two species centred on Highveld Grassland of South Africa or savanna in SE Africa.

Major clade, mostly smaller-bodied, more terminally derived.

Groups 30–31: Three species centred on Highveld Grassland, South Africa (1 sp.), upland savanna, N Namibia (1 sp.)
or N Highveld Grassland and eastern savanna to East Africa (1 sp.).
Group 38: One moist E coastal subspecies of a species found in moist savannas across tropical Africa.
Groups 39-40: Three species; two centred on dry savanna in N Namibia or in the east from South to East Africa; one
with a distribution from the E coast up the Zambezi Valley.
Group 41: Two species centred on southern African savannas; one on coarse-grained soils in drier savanna; one on
finer-grained soils in moister savanna.
Group 42: One species centred on SE African savanna.
Groups 47–48: Two species with distributions centred on deep sands of the southern Kalahari or Maputaland (NE coast
South Africa).

COPRINI
186 SURICATA 6 (2020)

Copris amyntor
Klug, 1855

= Copris confusus Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type localities: C. amyntor: ‘Sena’ [central Mozambique]; as C. confusa:

‘Caffraria tota’ [inexact, all SE South Africa].
Taxonomy: Accepted Group 42 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Widespread in drier, lower-altitude, eastern savanna wood-
lands from N South Africa across E Botswana, Zimbabwe Malawi and
♂ Mozambique; Kenyan records are undoubtedly confused with the
closely related C. harrisi Waterhouse, 1891.
Locality data (mean ± SD, range): Altitude: 739 ± 421, 24–1 694 m;
annual rainfall: 631 ± 149, 303–1 659 mm; annual temperature (max.
+ min. /2): 20.7 ± 1.9, 15.1–25.9°C (N=255).
Habitat: On Gauteng bushveld: restricted to finer-grained soils: sand (0),
sandy clay loam (31), primarily in open woodland (24), grassland (3),
shaded thickets (4); largely supported by uMkhuze Game Reserve data:
sand (1), clay (11) and DBRU collection records: clay (7), sandy clay
loam (25), sandy loam (42), sand (27) in shrub/woodland (73), pas-
ture/grassland (16), crop fields (4).
Food types: At Phalaborwa: bias to dung of monogastric herbivores
compared to ruminant herbivores and omnivores: elephant (26), cattle
(3), pig (4); at SA Wildlife College: reduced population density where
elephants are absent on finer-grained, gabbro-derived soils in farm
rangeland, (1.7) compared to the Kruger National Park (33.0); DBRU
collection records: elephant (35), rhinoceros (13), zebra (2); cattle (73),
buffalo (9), wildebeest (2); baboon (1).
Temporal activity: Flight activity during darkness primarily in the early
2 mm summer rainy season (Nov. to Jan.) tailing off into late summer (Feb.
to Apr.) (Gauteng); in the warmer lowlands, recorded during the cool
winter dry season (July) in elephant dung (Kruger National Park); at-
tracted to light.
Ecoregions Botswana, Zimbabwe: Primarily Southern African Bushveld
(AT0717), Zambezian and Mopane Woodlands (AT0725), Southern
Miombo Woodlands (AT0719).
Bioregions South Africa: Mainly Central Bushveld (SVcb), Mopane
(SVmp) and Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 016 860 km2; ecological data suggest
that loss of elephants and clearance of woodland would impact on
population density although collection data suggest that C. amyntor is
sufficiently adaptable to remain extant over a wide area where transfor-
mation varies from 0 to 58% (SVl), 2–49% (SVcb), 0–20% (SVmp);
assessed as Least Concern (LC).
Conservation measures: Quantitative data suggest that conservation
status of C. amyntor will be enhanced by continued protection of ele-
phants and woodland on finer-grained soils; protected in some lowland
national parks: Kruger (South Africa), Gorongosa (Mozambique); also
uMkhuze and Hluhluwe–iMfolozi game reserves (South Africa).

2 mm
♀

COPRINI
SURICATA 6 (2020) 187

Copris anceus
(Olivier, 1789)

= Copris minator Harold, 1880

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus anceus: Not stated. C. minator: ‘Südafrika’

[South Africa].
Taxonomy: Accepted Group 21 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Centred on the SW coast of South Africa (red squares) with
tentative records from Namaqualand and S coastline (black squares); S ♂
coast record probably accurate.
Locality data (mean ± SD, range): Altitude: 77 ± 79, 0–340 m; annual
rainfall: 270 ± 84, 95–475 mm; annual temperature (max. + min. /2):
16.7 ± 1.0, 14.8–18.5°C (N=35).
Habitat: No quantitative assessment of soil association, but DBRU collec-
tion records show a strong bias to deep coastal sands (18) compared to
sandy loam (1); on deep sand in West Coast National Park: strong bias
to natural shrubland (72) compared to adjacent sparse pasture (5) (now
restored as shrubland); largely supported by DBRU collection records:
scrub/shrubland (12), pasture/grassland (3), fallow crop fields (4).
Food types: No quantitative assessment; DBRU collection records from
cattle (17) and horse (1) dung.
Temporal activity: Flight activity in darkness during warmer parts of the
winter rainy season; on SW coast: primarily in spring (Aug., Sept.) tail-
ing off into Oct. with few records in summer (Nov. to Mar.) and winter
(June, July), but slightly more in autumn (Apr., May); evening flight
varying in time with the onset of darkness.
Bioregions South Africa: Primarily centred on the W coastal extremes
of the Fynbos Biome, particularly in vegetation units of the Western
Strandveld (FS 1, 3, 5) and Sand Fynbos (FFd 3, 4, 5).
2 mm
Assessment rationale: EOO = 7 625 km2; although C. anceus has been
recorded in fallow crop fields, clearance of natural vegetation drastical-
ly reduces population density; protected in several reserves although
transformation due to urban development and cultivation varies from
25 to 80% across the main range (red squares); despite this small range
centred on a highly transformed region, currently assessed as Least
Concern, although monitoring of status would be useful.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on soil and food associations; protected in
West Coast National Park, Cape of Good Hope Nature Reserve and the
private Grotto Bay Nature Reserve.

♀
5 mm

COPRINI
188 SURICATA 6 (2020)

Copris antares
Ferreira, 1958a

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Orange’ [Free State, South Africa].

Taxonomy: Accepted Group 30 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Widespread at higher altitude on the Highveld and at lower
altitude along the SE coast of South Africa.
♂ Locality data (mean ± SD, range): Altitude: 1 136 ± 540, 1–1 860 m;
annual rainfall: 628 ± 152, 327–961 mm; annual temperature (max. +
min. /2): 16.5 ± 1.7, 13.0–20.8°C (N=61).
Habitat: No quantitative assessment; DBRU collection records from clay
(1), sandy clay loam (8), sandy loam (14), sand (4) primarily in grass-
land/pasture (19), rarely in open woodland (1) and fallow crop fields
(2).
Food types: No quantitative assessment; DBRU collection records from
ruminant herbivore dung: cattle (33), buffalo (2), sheep (1).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to May).
Bioregions South Africa: Centred on Dry Highveld Grassland (Gh),
Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs)
(Grassland Biome), and the lower-altitude Albany Thicket (AT) and
Indian Ocean Coastal Belt (CB) biomes.
Assessment rationale: EOO = 269 985 km2; widespread on the South
African Highveld; no quantitative data available to support soil gen-
eralisation or determine if modification of natural grasslands would
be detrimental to population density; extensive transformation due to
cultivation occurs over much of its Grassland Biome range (2–50%
2 mm Gh, 6–69% Gm, 2–40% Gs); currently assessed as Least Concern on
basis of wide EOO.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
possible effects of natural grassland modification to improved pastures;
nevertheless widespread in farmland; protected in Abe Bailey Nature
Reserve and Addo Elephant National Park.

♀
2 mm

COPRINI
SURICATA 6 (2020) 189

Copris bootes
Klug, 1855

= Copris excavata Klug, 1855

= Copris bornemisszai Ferreira, 1972
Global: LC

J A S O N D J F M A M J

Type localities: C. bootes: ‘Inhambane’; C. excavata: ‘Inhambane’ [SE

Mozambique coast]; C. bornemisszai: ‘Umfolozi (Natal)’ [Hluhluwe–
iMfolozi Game Reserve, KwaZulu-Natal, South Africa].
Taxonomy: Accepted Group 4 species; sexually dimorphic (head, protho-
racic disc), prominence of characters varies with body size.
Distribution: Currently patchily distributed in moist, hot, lowland sa-
♂
vannas from SE to E Africa: NE South Africa, N Botswana, Zimbab­
we, Mozambique, Tanzania, Kenya; most available southern African
records (19 out of 21) from game reserves or national parks.
Locality data (mean ± SD, range): Altitude: 497 ± 395, 0–1 106 m; an-
nual rainfall: 740 ± 174, 511–1 136 mm; annual temperature (max. +
min. /2): 22.4 ± 1.4, 20.5–25.9°C (N=19).
Habitat: No quantitative assessment: DBRU collection data from clay (1),
sandy clay loam (2), sandy loam (8), deep sand (4), exclusively from
wooded savanna: shrubland (2) or woodland (12).
Food types: In Botswana: sampled exclusively to elephant dung (17),
none on dung of pig, cattle, sheep or on carrion (0); DBRU collec-
tion records dominated by monogastric herbivore dung: elephant (7),
middens of hooked-lipped (black) or square-lipped (white) rhinoceros
(10), horse (1), few on ruminant herbivore dung: buffalo (1), cattle (1).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Aug. to May).
Ecoregions Botswana, Zimbabwe: Mostly in reserves bordering and
within Zambezian Baikiaea Woodlands (AT1726).
Bioregions South Africa: Only in reserves in the south of the Lowveld 2 mm
(SVl) (Savanna Biome).
Assessment rationale: EOO = 959 800 km2; AOO much smaller and
almost exclusively restricted to game reserves in recent decades as re-
flected by patchy distribution; extreme association with elephant or
rhinoceros dung, but protected over a very wide area in many reserves
containing woodland; therefore, currently assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations; dung and probable
vegetation specialisation suggest conservation status is dependent on
continuing protection of savanna woodlands and large monogastric
herbivores in reserves; currently protected in various game reserves or
national parks: uMkhuze, Hluhluwe–iMfolozi (South Africa), Hwange
(Zimbabwe), Gorongosa (Mozambique), Chobe (Botswana), Manyara
(Tanzania), Meru, Samburu, Amboseli (Kenya).

♀
2 mm

COPRINI
190 SURICATA 6 (2020)

Copris caelatus
(Fabricius, 1794)

= Copris contracta Boheman, 1857

Global: LC
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type localities: As Scarabaeus caelatus: ‘Cap. Bon. Spei’ [Cape of Good

Hope, South Africa]; C. contracta: ‘juxta fluvium Limpopo’ [near Lim-
popo River, southern Africa].
Taxonomy: Accepted Group 17 species, but name requires validation;
type(s) of C. caelatus need to be compared with those of S Cape species:
♂ C. victorini Boheman, C. sexdentatus Thunberg and C. sphaeropterus
Harold; type locality of C. contractus (= C. contracta) inaccurate, but the
species occurs nearby on the Soutpansberg; sexually dimorphic (head
and prothoracic disc), prominence of characters varies with body size.
Distribution: Primarily moist to wet upper edges of E escarpment in both
N (Limpopo, Mpumalanga, KwaZulu-Natal) and S (Eastern Cape);
also descending to lower-altitude sub-escarpment levels in S; restricted
to mid-summer rainfall region, South Africa.
Locality data (mean ± SD, range): Average altitude: 1 480 ± 473,
483–2 296 m; average annual rainfall: 792 ± 94, 626–1 047 mm; av-
erage annual temperature (max. + min. /2): 14.9 ± 2.5, 10.3–20.7°C
(N=40).
Habitat: No quantitative assessment; DBRU collection records all from
finer-grained soils: sandy loam (9), sandy clay loam (9) in pasture/
grassland (17).
Food types: No quantitative assessment; DBRU collection records all
from dung of cattle (22).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Sept. to May).
2 mm Bioregions South Africa: Primarily E edges of Mesic Highveld Grass-
land (Gm) and Drakensberg Grassland (Gd) plus lower-altitude Sub-
Escarpment Grassland (Gs) (Grassland Biome).
Assessment rationale: EOO = 91 950 km2; widespread along E escarp-
ment, but no quantitative ecological data; however, habitat transforma-
tion low in areas showing greatest frequency of records along Lesotho
and Eswatini borders; currently assessed as LC.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
data on effects of habitat transformation and pasture improvement;
protected in the uKhahlamba Drakensberg Park (World Heritage Site)
of the central Drakensberg.

♀
2 mm

COPRINI
SURICATA 6 (2020) 191

Copris cambeforti
Nguyen-Phung, 1988a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal, P. N. Kruger’ [Kruger National Park, South

Africa].
Taxonomy: Accepted Group 2 species; sexually dimorphic (head, protho-
racic disc), prominence of characters varies with body size.
Distribution: Known from only two spot localities in N South Africa;
Blouberg (may be inexact) and Punda Milia area (Lowveld). ♂
Locality data (mean ± SD, range): Altitude: 952 ± 694, 461–1 443 m;
annual rainfall: 578 ± 77, 523–632 mm; annual temperature (max. +
min. /2): 20.0 ± 4.0, 17.2–22.9°C (N=2).
Habitat: A single DBRU observation on sand in open woodland.
Food types: A single DBRU observation from elephant dung.
Temporal activity: Flight activity in darkness; recorded in the summer
rainy season (Mar.).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome).
Assessment rationale: EOO = 540 km2 (underestimated); known from
few, somewhat inexact localities, therefore, assessed as Data Deficient
(DD).
Conservation measures: A survey is required to determine its full EOO,
AOO and ecological associations before an assessment may be made of
its conservation status; protected in the Kruger National Park.

2 mm

♀
2 mm

COPRINI
192 SURICATA 6 (2020)

Copris capensis
Waterhouse, 1891

No synonyms
Global: LC (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘South Africa’.

Taxonomy: Accepted Group 22 species; sexually dimorphic (head,
prothoracic disc); prominence of characters varies with body size.
Distribution: Within the winter rainfall region of South Africa on the W
coast and SW hinterland; also a single record from the Eastern Cape in
♂ the E of the bimodal rainfall region.
Locality data (mean ± SD, range): Altitude: 175 ± 213, 0–1 007 m; an-
nual rainfall: 205 ± 95, 54–430 mm; annual temperature (max. + min.
/2): 17.1 ± 1.1, 15.0–18.6°C (N=43).
Habitat: No quantitative assessment; DBRU collection records biased to
coarse-grained soils: sand (17), sandy loam (6), sandy clay loam (1),
with a possible bias to more open vegetation: scrub/shrubland (6),
grassland/pasture replacing natural shrubland (10) and tolerance of
disturbance: fallow crop fields (6).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (22) with two from horse dung.
Temporal activity: Flight activity in darkness; primarily during the winter
rainy season across its main W range (Apr. to Oct.).
Bioregions South Africa: In N: Centred on Namaqualand Sandveld (Sks)
(Succulent Karoo Biome); In S: Scattered across vegetation units of
Western Strandveld (FS), Sand Fynbos (FFd), Sandstone Fynbos (FFs),
Granite Renosterveld (FRg) and Shale Renosterveld (FRs) (Fynbos
Biome).
Assessment rationale: EOO = 45 395 km2; apparently some tolerance to
vegetation transformation (dry pastures, crop fields); possibly owing to
5 mm high frequency occurrence in low profile, natural scrub and shrubland
of arid areas; IUCN Red List as Data Deficient (DD) now re-assessed
as Least Concern (LC) on basis of widespread occurrence in the mostly
little-transformed vegetation of Namaqualand (3–10%); however, only
recorded in low population density in the more extensively transformed
S of its range (17–90%, higher on finer-grained soils).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
soil type associations and the effects on population density of clearance
of natural scrub or shrubland; protected in two national parks: West
Coast, coastal part of Namaqua.

♀
5 mm

COPRINI
SURICATA 6 (2020) 193

Copris cassius
Péringuey, 1901

= Copris cassius Péringuey, 1901, subsp. angolensis Ferreira, 1962a

Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: C. cassius: ‘British Bechuanaland (Kanye)’ [Kanye, Bo-

tswana]. subsp. angolensis: ‘Acampamento da Reserva de Caça da Cam-
eia’ [Angola: Cameia National Park, campsite].
Taxonomy: Accepted Group 47 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size; as-
suming that the bias in locality records to the SW Kalahari is not a ♂
collection artefact (see habitat notes); the synonymy of subsp. angolensis
should be re-assessed.
Distribution: Centred on the SW Kalahari with occurrence in deep sand
outliers at the eastern edge of the range.
Locality data (mean ± SD, range): Altitude: 1 161 ± 157, 839–1 512 m;
annual rainfall: 372 ± 106, 191–689 mm; annual temperature (max. +
min. /2): 19.2 ± 1.0, 15.7–21.3°C (N=92).
Habitat: On Gauteng bushveld: restricted to deep sands (179), sandy
clay loam (0) and biased to open vegetation, grassland (172), open
woodland (6), shaded thickets (1); on deep sands in Botswana: only in
open vegetation (grassland, shrubland) of the arid SW (28) not moister,
wooded NE (0); largely supported by DBRU collection records: sand
(47), sandy loam (3), in grassland (22), shrubland (13) or open wood-
land (12).
Food types: In Botswana: poorly attracted to dung of ruminant herbivores
(cattle: 2) or omnivores (pig: 2), instead biased to dung of monogastric
herbivores, (elephant: 24) even though elephants are absent from the
arid SW; absent from carrion (chicken livers) and sheep pellets (0);
DBRU collection records primarily from dung of cattle (42), also 2 mm
horse/donkey (5).
Temporal activity: Flight activity during darkness, primarily during the
summer rainy season (Oct. to May); on Gauteng bushveld: primarily
active in early summer (Oct. to Jan.) tailing off to late summer (Feb. to
Apr.) and autumn (May).
Ecoregions Namibia, Botswana: Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Primarily centred on the Eastern Kalahari Bush-
veld Bioregion (SVk) with records in the Kalahari Duneveld (SVkd) and
deep sand outliers of the Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 547 290 km2 (southern range); SW Kala-
hari range coincides with areas of deep sand used primarily for grazing
of livestock; transformation (cultivation) limited although this increas-
es to the east (2–25%, SVk; 4–24%, sandy units in SVcb); however,
this may not be detrimental to population density owing to recorded
association with grassland; assessed as Least Concern (LC).
Conservation measures: Owing to conflicting results, assessment of
conservation status would be improved by further quantitative data
on food associations; protected in Khutse Game Reserve (Botswana),
Tswalu Kalahari Reserve and Nylsvlei Nature Reserve (South Africa).

♀ 2 mm

COPRINI
194 SURICATA 6 (2020)

Copris cornifrons
Boheman, 1860

= Copris curvicornis Boheman, 1860

= Copris achates Péringuey, 1901
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type localities: C. cornifrons: ‘juxta lacum N’Gami’ [near Lake Ngami,

Botswana]; C. curvicornis: ‘juxta lacum N’Gami’; C. achates: ‘? N’Gami-
land’ [N Botswana].
Taxonomy: Accepted Group 11 species; sexually dimorphic (head,
♂ prothoracic disc), prominence of characters varies with body size.
Distribution: Restricted to the dry to arid southwest Kalahari.
Locality data (mean ± SD, range): Altitude: 1 111 ± 153, 830–1 367 m;
annual rainfall: 312 ± 90, 157–491 mm; annual temperature (max. +
min. /2): 19.4 ± 1.1, 17.5–22.4°C (N=38).
Habitat: No quantitative assessment; DBRU collection records almost
exclusively from deep sand (20), sandy loam (1) in open vegetation:
grassland (8), open shrubland (10), open woodland (3).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (16), also horse/donkey dung (3).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Sept. to Apr.).
Ecoregions Botswana: E Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Centred on the Kalahari Duneveld (SVkd) and
Eastern Kalahari Bushveld (SVk) (Savanna Biome).
Assessment rationale: EOO = 286 300 km2; EOO ~equivalent to wide-
spread AOO across an area of deep sands used primarily for the grazing
of farm livestock on whose dung it has been recorded in low population
density; transformation 0–2% over most of RSA range (SVkd, W of
SVk); assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
2 mm
improved by quantitative data on ecological associations, particularly
further investigation to determine if population density is naturally
low or an artefact of available data; a recent quantitative gradsect study
from NE to SW across the Botswana Kalahari recorded only a single in-
dividual attracted to elephant dung bait; protected in Kgalagadi Trans-
frontier Park and Tswalu Kalahari Reserve (Botswana, South Africa).

♀
2 mm

COPRINI
SURICATA 6 (2020) 195

Copris corniger
Sahlberg, 1823

= Copris lunarioides Waterhouse, 1891 (pars)

= Copris laticornis Péringuey, 1901 (pars)
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As C. cornigera: Not stated; C. lunarioides pars: ‘S. Africa’

[South Africa]; C. laticornis pars: ‘Cape Colony (Albany, Alexandria)’
[Eastern Cape, South Africa].
Taxonomy: Accepted Group 26 sexually dimorphic (head, prothoracic
disc), prominence of characters varies with body size. ♂
Distribution: Highveld Grassland to the N and Eastern Cape lowlands
to the SE; on a 500–2 800 m gradsect from coastal hills to Sani Pass,
KwaZulu-Natal, recorded only at 1 000–1 500 m.
Locality data (mean ± SD, range): Altitude: 1 356 ± 429, 0–2 274 m;
annual rainfall: 718 ± 127, 371–1 121 mm; annual temperature (max.
+ min. /2): 15.5 ± 1.7, 10.6–20.5°C (N=90).
Habitat: At Carolina: sampled from improved Kikuyu grass pasture (5)
and fallow crop fields (1), but not natural Themeda grassland (0);
DBRU collection records suggest a bias to finer-grained soils: clay (1),
sandy clay loam (22), sandy loam (15), sand (1) mainly in pasture/
grassland (33), rather than open shrub/woodland (3).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (44), one from horse dung.
Temporal activity: Evening flight activity in darkness during the summer
rainy season (Oct. to Apr.).
Bioregions South Africa: Centred on Dry Highveld Grassland (Gh),
Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs) and
margins of the Drakensberg Grassland (Gd) (Grassland Biome) with
marginal occurrence in three coastal bioregions.
Assessment rationale: EOO = 318 935 km2; widespread across the farm- 5 mm
land on Highveld grasslands where it is readily attracted to the dung of
domestic livestock, particularly that of cattle; data from Carolina may
indicate that C. corniger is fairly adaptable to grassland modification,
although extensive transformation due to cultivation occurs over much
of the Grassland Biome (4–60% Gh, 6–69% Gm, 2–50% Gs); assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative ecological data; protected in Golden Gate
National Park.

♀
5 mm

COPRINI
196 SURICATA 6 (2020)

Copris crassus
Deschodt & Davis, 2015 (see Deschodt et al. 2015a)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Suikerbosrand Nat. Res., Gauteng, South Africa, 28°28-

32’S 28°10-17’E’.
Taxonomy: Accepted Group 3 species; sexually dimorphic (head, protho-
racic disc), prominence of characters varies with body size.
Distribution: Moist South African highlands; N edge of Highveld and E
♂ escarpment edge from Mpumalanga and KwaZulu-Natal to the Eastern
Cape; occurs at lower altitude at the S range limits.
Locality data (mean ± SD, range): Altitude: 1 437 ± 569, 324–1 948 m;
annual rainfall: 721 ± 134, 484–900 mm; annual temperature (max. +
min. /2): 15.0 ± 2.2, 12.2–18.0°C (N=8).
Habitat: No adequate quantitative assessment; in Suikerbosrand Na-
ture Reserve: primarily sampled in cool S-facing montane grassland
(> 1 880 m), not highveld grasslands (1 630–1 690 m); sampling
records and DBRU collection records (2) only from sandy loam and
sandy clay loam in grassland (1) and open woodland (1), but probably
biased to grassland.
Food types: No quantitative assessment; sampled and collected by DBRU
only from cattle dung.
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Sept. to Mar.); attracted to light.
Bioregions South Africa: Centred along the highland margins of the
Mesic Highveld Grassland (Gm), Drakensberg Grassland (Gd) and
Sub-Escarpment Grassland (Gs) (Grassland Biome); also disturbed
5 mm Sandstone Fynbos (FFs) (Fynbos Biome).
Assessment rationale: EOO = 41 240 km2; AOO probably smaller re-
flected by patchy records, mostly in moist highland grassland units of
the Grassland Biome in the summer rainfall region; transformation
in vegetation units around known localities: Gm, 15%; Gd, 5%; Gs,
10–20%; somewhat Data Deficient (DD), but assessed as Least Con-
cern (LC) on basis of wide range.
Conservation measures: Assessment of conservation status would be
improved by a survey to determine additional locations, ecological
associations, and possible effects of grassland modification; protected
in Suikerbosrand Nature Reserve and uKhahlamba Drakensberg Park
(World Heritage Site).

♀
5 mm

COPRINI
SURICATA 6 (2020) 197

Copris denticulatus
Nguyen-Phung, 1988c

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Shilouvane, près Leydsdorp’ [Shiluvane near Leydsdorp,

Limpopo Province, South Africa].
Taxonomy: Accepted Group 26 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Widespread in moist savannas of SE Africa: N South Africa,
E Botswana, Zimbabwe, Mozambique, Tanzania; in Tanzania, range
overlaps with the closely related C. fallaciosus Gillet, 1907, whose range ♂
extends to Uganda, Kenya, Somalia, Ethiopia; black squares are ques-
tionable outlier localities.
Locality data (mean ± SD, range): Altitude: 927 ± 437, 9–1 850 m; an-
nual rainfall: 690 ± 151, 383–1 177 mm; annual temperature (max. +
min. /2): 19.6 ± 1.9, 13.4–25.2°C (N=182).
Habitat: On Gauteng bushveld: strong bias to finer-grained soils, sandy
clay loam (30), deep sand (3) in a range of vegetation types, grass-
land (10), open woodland (4), shaded thickets (19); not supported by
DBRU collection records: clay (7), sandy clay loam (8), sandy loam
(19), sand (15) in grassland/pasture (21), shrubland (8), open wood-
land (8), forest (2), also crop fields (2).
Food types: No quantitative data; DBRU collection records from rumi-
nant herbivore dung: cattle (61), buffalo (3) far outweigh those from
monogastric herbivore dung: elephant (4), square-lipped (white) rhi-
noceros (1), warthog (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to May); on Gauteng bushveld: seasonal peak in early
summer (Nov., Dec.); attracted to light; bias to colonisation of older
dung (18%, 1–2 day old dung; 63%, 2–4 day). 5 mm
Ecoregions Botswana, Zimbabwe: Centred on Southern African Bush-
veld (AT0717), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Mainly in the Central Bushveld (SVcb), Mo-
pane (SVmp) and Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 940 275 km2; widely distributed in farm-
land where it is readily attracted to cattle dung, often in open vegeta-
tion; therefore, perhaps not seriously threatened in the South African
part of its range where transformation through cultivation varies from
0 to 58% (SVl), 2–49% (SVcb), 0–20% (SVmp); assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative ecological data to resolve apparent
conflicts between quantitative and qualitative results; protected in the
Kruger National Park (South Africa).

♀ 5 mm

COPRINI
198 SURICATA 6 (2020)

Copris elphenor
Klug, 1855

= Copris laticornis Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type localities: C. elphenor: ‘Sena’ [Central Mozambique]; C. laticornis:

‘Caffraria interiori’ [interior of SE South Africa].
Taxonomy: Accepted Group 23 species; sexually dimorphic (head, protho-
racic disc), prominence of characters varies with body size.
Distribution: Widespread in dry savanna from S to E Africa: N South Af-
rica, N Namibia, Botswana, Zimbabwe, Malawi, Mozambique, S An-
♂ gola, S Democratic Republic of the Congo (DRC), Zambia, Tanzania,
S Kenya, possibly Somalia; West African records are certainly errors.
Locality data (mean ± SD, range): Altitude: 972 ± 425, 0–1 998 m; an-
nual rainfall: 581 ± 188, 214–1 415 mm; annual temperature (max. +
min. /2): 20.0 ± 2.1, 14.5–26.1°C (N=420).
Habitat: On Gauteng bushveld: strong bias to coarse-grained soils in rel-
atively open vegetation, sand (247), sandy clay loam (44), grassland
(127), open woodland (148), shaded thickets (16); similar patterns
at Bushlands Halt: sand (182), clay loam (23), grassland (199), dense
woodland (6); also supported by DBRU collection records: clay (5), san-
dy clay loam (25), sandy loam (56), sand (72) in pasture/grassland (53),
shrubland (34), open woodland (50), forest (2), also crop fields (3).
Food types: On Gauteng bushveld: strong bias to dung of ruminant her-
bivores (cattle: 32) rather than that of monogastric herbivores (horse:
10), omnivores (pig: 3) or carrion (0); similar at Phalaborwa: dung of
cattle (72), elephant (14), pig (1); DBRU collection records, ruminant
herbivore dung: cattle (153), buffalo (6), wildebeest (3), monogastric
herbivore dung: elephant (18), rhinoceros (6), donkey (1), horse (3);
also baboon (2).
5 mm Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Aug, Oct. to May); on Gauteng bushveld: seasonal
peak, early to mid-summer (Oct. to Jan.) tailing off into late summer
(Feb. to Apr.); flight activity peak 1–2 hours after nightfall (20:00–
22:00); attracted to light; at Phalaborwa: much greater activity on a
mild evening following heavy rainfall (78) than on a cool evening fol-
lowing light rain (3) or a hot evening under dry conditions (4); biased
to colonisation of fresh dung (0–1 day: 56%, 1–3 days: 32%,).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna
(AT1309), Southern African Bushveld (AT0717), Zambezian and Mo-
pane Woodlands (AT0725), Southern Miombo Woodlands (AT0719),
Angolan Mopane Woodlands (AT0702), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Centred on the Eastern Kalahari Bushveld
(SVk), Central Bushveld (SVcb), Mopane (SVmp) and Lowveld (SVl)
(Savanna Biome); some records in Dry Highveld Grassland (Gh) (W
Grassland Biome); marginal occurrence in five other bioregions.
Assessment rationale: EOO = 3 968 635 km2; widespread in farmland where
it is occurs in both grassland and open woody vegetation with association
biased to cattle dung; tolerance of open vegetation may reduce impact of
habitat transformation over NE part of range in South Africa: 0–58%
(SVl), 2–49% (SVcb), 0–20% (SVmp); assessed as Least Concern (LC).
Conservation measures: None required owing to wide occurrence, tol-
5 mm erance of open vegetation and ecological suitability for farm livestock;
♀ protected in Kruger National Park (South Africa).

COPRINI
SURICATA 6 (2020) 199

Copris evanidus
Klug, 1855

= Copris modestus Boheman, 1857

= Copris orphanus Guérin-Méneville, 1847, sensu Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As C. evanida: ‘Sena’ [Central Mozambique]; As

C. modesta: ‘Caffraria tota’ [all SE South Africa]; C. orphanus sensu
Péringuey, 1901: ‘Southern Rhodesia (Victoria Falls), Mozambique
(Lourenço-Marquez)’ [Zimbabwe, S Mozambique].
Taxonomy: Accepted Group 40 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size. ♂
Distribution: Widespread in dry savanna along the E seaboard of Afri-
ca: N South Africa, E Botswana, Zimbabwe, Mozambique, Tanzania,
Kenya; also cited from Uganda, Ethiopia.
Locality data (mean ± SD, range): Altitude: 802 ± 401, 55–1 762 m;
annual rainfall: 564 ± 130, 304–909 mm; annual temperature (max. +
min. /2): 20.6 ± 1.8, 15.9–25.7°C (N=217).
Habitat: In uMkhuze Game Reserve: extreme bias to finer-grained soils,
sandy clay loam /clay (20), deep sand (0); on Gauteng bushveld: bias
less extreme, sandy clay loam (22), deep sand (11) with a slight bias to
open vegetation, grassland (20), open wood/thicket (13); partly sup-
ported by DBRU collection records: clay (2), sandy clay loam (12),
sandy loam (37), sand (25) in grassland pasture (21), shrubland (10),
open woodland (39).
Food types: At Phalaborwa: bias to elephant dung (11), few on cattle
(1) and pig dung (0); in Botswana: bias to elephant dung (8), few on
sheep (1), pig, cattle dung and carrion (0); DBRU collection records
on ruminant herbivore dung: cattle (84), buffalo (3), wildebeest (2),
monogastric herbivore dung: elephant (17), rhinoceros (2), zebra (1),
also on baboon dung (1).
Temporal activity: Flight activity in darkness, primarily in the summer 2 mm
rainy season (Aug., Oct. to Apr.); on Gauteng bushveld: seasonal peak
early to mid-summer (Oct. to Jan.) tailing off in late summer (Feb. to
Apr.); attracted to light.
Ecoregions Botswana, Zimbabwe: Centred on Southern African Bush-
veld (AT0717), Zambezian and Mopane Woodlands (AT0725); also E
margins of Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Centred on the Eastern Kalahari Bushveld
(SVk), Central Bushveld (SVcb), Mopane (SVmp) and Lowveld (SVl)
(Savanna Biome), with some records in the N of the Dry Highveld
Grassland (Gh) (W Grassland Biome).
Assessment rationale: EOO = 2 357 905 km2; possible bias to elephant
dung may result in lower population densities over much of the species
range; but remains extant using other dung types; apparently shows
similar frequency in both grassland and open woodland whereas results
for possible soil bias are inconclusive; widespread in both reserves and
farmland; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by conclusive quantitative data on soil and food associations;
protected in various reserves including the Kruger National Park and
uMkhuze Game Reserve (South Africa).

♀ 2 mm

COPRINI
200 SURICATA 6 (2020)

Copris fidius
(Olivier, 1789)

= Scarabaeus plutus Fabricius, 1794

= Copris (Paracopris) bihamatus Balthasar, 1965a
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Scarabaeus fidius: ‘l’Amérique méridionale’ [erroneous-

ly ascribed to Central America]; S. plutus: ‘India orientali’ [erroneously
ascribed to eastern India]; C. bihamatus: ‘Südafrika, Kapland (ohne
nähere fudortangabe)’ [South Africa, Cape (without more detailed
place of origin)].
♂ Taxonomy: Accepted Group 3 species; sexually dimorphic (head, protho-
racic disc), prominence of characters varies with body size.
Distribution: Dual distribution pattern; one along entire S and E coast-
line, South Africa to S Mozambique; the other parallel along lower
part of mountain escarpment diverging inland in NE South Africa;
also in Cederberg (Western Cape) and N edge of Highveld; Namibia
and Democratic Republic of the Congo (DRC) citations are certainly
errors.
Locality data (mean ± SD, range): Altitude: 650 ± 571, 0–2 194 m; an-
nual rainfall: 755 ± 222, 264–1 171 mm; annual temperature (max. +
min. /2): 17.6 ± 2.3, 11.0–22.4°C (N=110).
Habitat: No quantitative assessment of soil type association; in the Wolk-
berg: extreme bias to shaded forest (55.6), grassland (0.6); similar in
Ithala Game Reserve: Ntshondwe forest (32), open woodland (2),
grassland (0); similar in Vernon Crookes Nature Reserve: forest (16),
grassland (0); in the Wolkberg: favours lower altitudes, 900–1 000 m
(15.6), 1 500–1 550 m (0.9); DBRU collection records on both finer
and coarse-grained soils, sand (3), sandy clay loam (4) in grassland/
2 mm pasture (4), woodland/forest (6).
Food types: In the Wolkberg: equally attracted to dung of ruminant her-
bivores (cattle: 67.1) and omnivores (pig: 50.6); DBRU collection re-
cords: primarily from dung of cattle (12), also monogastric herbivores,
elephant (1), human/cattle dung mix (2).
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season in the NE part of the range (Oct. to May); spring activity
(Sept.) in the winter rainfall region; attracted to light.
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Primarily associated with scattered forest patch-
es throughout most of the Afrotemperate, Subtropical and Azonal For-
ests Biome (FO); rare in Northern Afrotemperate Forest (FOz 2); none
in Dune (FOz 8) and Azonal Forest (FOa).
Assessment rationale: EOO = 116 200 km2; widespread along the entire
E seaboard of South Africa where it is associated with forest patches;
although forest clearance would radically reduce population density,
known transformation of forest patches is low (5–6%), even though
the original extent of many is unknown; currently assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil associations; protected in many
game, forest and nature reserves including Lekgalameetse, Ithala,
Hluhluwe–iMfolozi, Ngome, Vernon Crookes and Tsitsikamma
(South Africa).
♀
2 mm

COPRINI
SURICATA 6 (2020) 201

Copris gracilis
Waterhouse, 1891

= Copris orphanus Guérin-Meneville, 1847, sensu Ferreira, 1958b

Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type localities: C. gracilis: ‘Caffraria’ [?southern Africa, see Taxonomy];

C. orphanus sensu Ferreira: ‘Okahandja’ [Namibia].
Taxonomy: Accepted Group 39 species, but status requires re-investigation;
N Namibian distribution is at odds with ‘Caffraria’ type locality unless
the term was used as a generality for southern Africa rather than in a
more restricted sense for SE South Africa; differences to the closely ♂
related, SE to E African, C. evanidus Klug require validation; sexually
dimorphic (head, prothoracic disc), prominence of characters varies
with body size.
Distribution: Dry to arid savanna: northern plateau of Namibia.
Locality data (mean ± SD, range): Altitude: 1 394 ± 243, 799–1 998 m;
annual rainfall: 340 ± 88, 125–549 mm; annual temperature (max. +
min. /2): 19.3 ± 1.6, 16.9–23.0°C (N=54).
Habitat: No quantitative assessment; DBRU collection records: sandy
clay loam (6), sandy loam (11), sand (10), mainly in woody vegetation:
open woodland (15), shrubland (10), grassland (2).
Food types: No quantitative assessment; DBRU collection records: mostly
on cattle dung (25) with a few records from horse (2), zebra (1) and
elephant (1) dung.
Temporal activity: Flight activity in darkness in the late summer rainy
season (Dec. to Apr.).
Ecoregions Namibia: Centred on S Angolan Mopane Woodlands
(AT0702), N Kalahari Xeric Savanna (AT1309) and central Namibian
Savanna Woodlands (AT1316).
Assessment rationale: EOO = 168 350 km2; probably not under threat 2 mm
due to widespread occurrence in woodland savannas on farmland
where it colonises cattle dung; assessed as Least Concern (LC) rather
than Data Deficient (DD).
Conservation measures: Taxonomic issues need to be resolved; assess-
ment of conservation status would be improved by quantitative data
on ecological associations; currently unknown if clearance of woodland
would be detrimental to population density or if it might be influenced
by particular dung type associations; protected in Etosha National Park
(Namibia).

♀ 2 mm

COPRINI
202 SURICATA 6 (2020)

Copris inhalatus
Quedenfeldt, 1884

subsp. inhalatus (Quedenfeldt, 1884)

subsp. perturbator (Péringuey, 1901)
subsp. grandis Nguyen-Phung & Cambefort, 1986b
subsp. sanctaeluciae Nguyen-Phung & Cambefort, 1986b
subsp. ngotto Moretto, 2011
No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

♂ Type localities: C. inhalatus: ‘Malange’ [Angola]; lectotype: Quango [An-

gola]; as C. perturbator: ‘Ovampoland (Evari River), Southern Rhode-
sia (no exact locality)’ [N Namibia; Zimbabwe]; subsp. grandis: ‘N.W.
Rhodesia; N. of Broken Hill’ [N of Kabwe, Zambia]; subsp. sanctaelu-
ciae: ‘St Lucia Estuary, Natal’ [KwaZulu-Natal, South Africa]; subsp.
ngotto: ‘Centrafrique, Ngotto’ [Ngotto, Central African Republic].
Taxonomy: Accepted Group 12 species comprising five subspecies; all are
sexually dimorphic (head, prothoracic disc), prominence of characters
varies with body size.
Distribution: Mega-Kalahari deep sands and E outliers; three subspp. in
Central Zambia; S Tanzania; or N Angola, W Democratic Republic
of the Congo (DRC) and Congo (Brazzaville); two subspp. in S; sanc-
taeluciae: Maputaland Centre of Endemism on coast of NE South Af-
rica and SE Mozambique; perturbator: N Namibia, Botswana, N South
Africa, Zimbabwe, SW Zambia.
Locality data (mean ± SD, range): subsp. perturbator (red squares):
Altitude: 1 147 ± 169, 924–1 444 m; annual rainfall: 492 ± 142,
346–761 mm; annual temperature (max. + min. /2): 19.9 ± 1.7,
17.5–22.4°C (N=14). subsp. sanctaeluciae (black squares): Altitude: 34
± 54, 0–273 m; annual rainfall: 831 ± 107, 682–1 004 mm; annual
temperature (max. + min. /2): 22.0 ± 0.5, 20.5–22.5°C (N=27).
Habitat: No quantitative assessment of soil bias; DBRU collection records
2 mm exclusively from sand in both subspp. (11); subsp sanctaeluciae: in Ma-
puto Special Reserve: primarily grassland (18.6) not forest (0.5); similar
across Richards Bay, post-mining, restoration gradient: grassland (42),
subsp. sanctaluciae woodland/forest (0); subsp. perturbator: no quantitative assessment;
DBRU collection records only shrubland/open woodland (5).
Food types: No quantitative assessment; DBRU collection records from
cattle (5) and elephant (1) dung.
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Feb.) extending into the cooler season for the coastal
subspp. (May to June).
Ecoregions Namibia, Botswana Zimbabwe, Mozambique: subsp. sanc-
taeluciae: Maputaland Coastal Forest Mosaic (AT0119); subsp. per-
turbator: Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726);
deep sand outliers in Southern Miombo Woodlands (AT0719).
Bioregions South Africa: subsp. sanctaeluciae: Centred on N Indian
Ocean Coastal Belt Biome (CB); subsp. perturbator: Centred on the
Eastern Kalahari Bushveld (SVk) and sand outliers in Central Bushveld
(SVcb) (Savanna Biome).
Assessment rationale: subsp. perturbator: EOO = 614 150 km2; wide-
spread, but uncommon, not recorded in a cross-Kalahari gradsect using

COPRINI
SURICATA 6 (2020) 203

five different bait types, although most records have been made on cat- ♀
tle dung; much of its SW range used for grazing of domestic livestock;
subsp. sanctaeluciae: EOO = 8 520 km2; occupies a relatively small, but
mostly protected range; both currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; reasons for low
population density in subsp. perturbator should be investigated, par-
ticularly as most known records do not coincide with protected areas
other than one in Hwange National Park (Zimbabwe); coastal subsp.
sanctaeluciae protected in Maputo Special Reserve (Mozambique) and
iSimangaliso Wetland Park (World Heritage Site) (South Africa).

5 mm

subsp. sanctaluciae

COPRINI
204 SURICATA 6 (2020)

Copris jacchoides
Nguyen-Phung & Cambefort, 1987

No synonyms
Global: LC
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type locality: ‘Transvaal’ [N South Africa].

Taxonomy: Accepted Group 19 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Highveld and E coastal regions: South Africa.
Locality data (mean ± SD, range): Altitude: 1 430 ± 357, 117–1 983 m;
♂ annual rainfall: 724 ± 109, 355–958 mm; annual temperature (max. +
min. /2): 15.3 ± 1.8, 12.1–21.3°C (N=52).
Habitat: No quantitative assessment; DBRU collection records primarily
on finer-grained soils: clay (1), sandy clay loam (16), sandy loam (9),
sand (1), in pasture/grassland (25); also fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (33).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Mar.).
Bioregions South Africa: Moister E of Grassland Biome: Mesic Highveld
Grassland (Gm), Sub-Escarpment Grassland (Gs); marginal occurrence
in W Dry Highveld Grassland (Gh) and four other bioregions.
Assessment rationale: EOO = 258 970 km2; primarily centred in moist,
cool, Highveld grasslands; at Carolina: recorded from improved Kikuyu
pasture and fallow crop fields as well as natural Themeda grasslands,
but numbers too limited to determine any habitat bias; transformation
varies from 6 to 69% (Gm), 2–40% (Gs) across main range; assessed as
Least Concern (LC) on basis of widespread occurrence.
5 mm Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
possible effects of grassland modification even though some records are
from disturbed habitat; protected in Golden Gate National Park.

5 mm
♀

COPRINI
SURICATA 6 (2020) 205

Copris jacchus
(Fabricius, 1775)

= Scarabaeus oedipus Fabricius, 1775

= Copris laticornis Wallengren, 1881
Global: NT
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus jacchus: ‘Cap B. S.’, lectotype, same type

locality; S. oedipus: ‘Cap B. S.’, lectotype same type locality [all Cape of
Good Hope, South Africa]; C. laticornis: Not stated.
Taxonomy: Accepted Group 19 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size. ♂
Distribution: Low-altitude range along the drier parts of the S coast of
South Africa in the winter and bimodal, spring/autumn, rainfall re-
gions where natural vegetation is dominated by shrubland.
Locality data (mean ± SD, range): Altitude: 244 ± 178, 0–541 m; annual
rainfall: 462 ± 140, 366–741 mm; annual temperature (max. + min.
/2): 17.5 ± 1.1, 15.8–19.2°C (N=6).
Habitat: No quantitative assessment; only a single DBRU habitat obser-
vation on clay in open shrub/grassland.
Food types: No quantitative assessment; DBRU collection records from
cattle (2) and buffalo (1) dung.
Temporal activity: Flight activity in darkness coinciding with warmer
spring and autumn temperatures (Oct., May).
Bioregions South Africa: Vegetation units in Shale Renosterveld (FRs)
(Fynbos Biome); Albany Thicket Biome (AT).
Assessment rationale: EOO = 9 990 km2; known range < 20 000 km2
with few known recent localities; natural lowland shrubland vegetation
units are highly transformed in the W: FRs (Swartland, 90%; Mossel
Bay, 58%), but less so in the E: AT (Sundays Thicket, 6%, but degraded
by overgrazing; Great Fish Thicket, 3%); rather than Data Deficient 5 mm

(DD), tentatively assessed as possibly Near Threatened (NT) on basis

of limited records from an often highly transformed region.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, particularly an
appraisal of its response to clearance of natural shrubland; without soil
and vegetation association data it is difficult to determine if the AOO
and population density would justify Least Concern (LC) status, the
current tentative assessment as NT, or even Vulnerable (VU); protected
in Addo Elephant National Park.

♀ 5 mm

COPRINI
206 SURICATA 6 (2020)

Copris laioides
Boucomont, 1932

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type localities: ‘Afrique de sud-ouest, Damaraland: Okanjande ou Oka-

handya sur le feuve Swakop’ [Okanjande or Okahandja on the River
Swakop, Namibia], lectotype, ‘Okanjande, S.W. Afrika’ [Mt Okon-
yande, Namibia].
Taxonomy: Accepted Group 20 species; sexually dimorphic (head,
♂ prothoracic disc), prominence of characters varies with body size.
Distribution: Dry savanna on the N Namibian plateau; reports from Zim-
babwe, Democratic Republic of the Congo (DRC) require validation.
Locality data (mean ± SD, range): Altitude: 1 429 ± 151, 1 097–1 627 m;
annual rainfall: 446 ± 64, 322–560 mm; annual temperature (max. +
min. /2): 20.6 ± 1.2, 18.9–23.0°C (N=20).
Habitat: No quantitative assessment; DBRU collection records from var-
ious soil types: clay (2), sandy clay loam (4), sandy loam (2), sand (4),
entirely in shrubland (5) or open woodland (8).
Food types: No quantitative assessment; DBRU records primarily on cat-
tle dung (11), also horse (1) and donkey (1) dung.
Temporal activity: Flight activity in darkness in the late summer rainfall
season (Dec. to Apr.).
Ecoregions Namibia: S Angolan Mopane Woodlands (AT0702), N Kala-
hari Xeric Savanna (AT1309).
Assessment rationale: EOO = 49 485 km2; potential threats currently
limited as it is widespread on various soil types in farmland savanna
where it colonises the dung of domestic livestock; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; possible effects
of clearance of natural shrubland and woodland need to be investigated
2 mm as no known localities coincide with protected areas.

2 mm
♀

COPRINI
SURICATA 6 (2020) 207

Copris macer
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Southern Rhodesia (Salisbury)’ [Harare, Zimbabwe].

Taxonomy: Accepted Group 41 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Patchy occurrence on finer-grained soils in moist areas of
SE Africa supporting open vegetation: N South Africa, Eswatini, Zim-
babwe; also cited from Mozambique; report from Tanzania requires
validation; that from Namibia possibly an error. ♂
Locality data (mean ± SD, range): Altitude: 1 153 ± 361, 433–1 890 m;
annual rainfall: 846 ± 150, 549–1 203 mm; annual temperature (max.
+ min. /2): 17.9 ± 2.1, 13.1–22.0°C (N=39).
Habitat: No quantitative assessment; DBRU collection records show bias
to finer-grained soils: clay (1), sandy clay loam (7), sandy loam (7),
sand (1), primarily in open vegetation: pasture/grassland (11), shrub/
woodland (2).
Food types: No quantitative assessment; DBRU collection records entire-
ly from ruminant dung: cattle (19), buffalo (1).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to May).
Ecoregions Zimbabwe: E Southern Miombo Woodlands (AT0719),
Eastern Zimbabwe Montane Forest Grassland Mosaic (AT1006).
Bioregions South Africa: Moist NE escarpment straddling the boundaries
of N Mesic Highveld Grassland (Gm) (Grassland Biome) and Lowveld
(SVl) (Savanna Biome); also moist patches in Central Bushveld (SVcb)
(Savanna Biome).
Assessment rationale: EOO = 65 380 km2 (underestimated); widespread 2 mm

in open vegetation of farmland, especially in moist grassland regions

where it readily colonises cattle dung; assessed as Least Concern (LC)
although transformation by cultivation and plantations in E Gm is
6–40%.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; possible effects
of grassland modification to improved pastures should also be investi-
gated; protected in Lake Mutirikwe Game Reserve (Zimbabwe).

♀
2 mm

COPRINI
208 SURICATA 6 (2020)

Copris mesacanthus
Harold, 1878

subsp. mesacanthus (Harold, 1878)

= Copris similis Balthasar, 1939a
subsp. tranvaalensis Nguyen-Phung, 1988
Global: LC
Endemic: RSA (subsp. tranvaalensis)

J A S O N D J F M A M J

Type localities: C. mesacanthus: ‘N’Yassa’, also lectotype: ‘Nyassa’ [inexact,

Lake Malawi region]; as Copris (Paracopris) similis: ‘Rhodesia, Monze,
Chikuni Mission’ [Zambia]; subsp. tranvaalensis: ‘Transvaal, Shilou-
♂ vane, près Leydsdorp’ [Shiluvane near Leydsdorp, Limpopo Province,
South Africa].
Taxonomy: Accepted Group 1 species treated as two subspecies, both
showing sexual dimorphism (head, prothoracic disc); prominence of
characters varies with body size.
Distribution: Essentially, restricted to SE Africa; subsp. tranvaalensis (red
squares): N South African savannas; subsp. mesacanthus (black squares):
savannas of N Botswana, Zimbabwe, Zambia, Malawi, Mozambique.
Locality data (mean ± SD, range): subsp. tranvaalensis: Altitude: 944 ±
449, 46–1 899 m; annual rainfall: 670 ± 137, 304–1 040 mm; annual
temperature (max. + min. /2): 19.4 ± 2.2, 13.1–24.5°C (N=133) | sub-
sp. mesacanthus: Altitude: 894 ± 443, 0–1 640 m; annual rainfall: 791
± 258, 427–1 623 mm; annual temperature (max. + min. /2): 20.9 ±
2.5, 17.1–26.1°C (N=34)
Habitat: subsp. tranvaalensis: On Gauteng bushveld: associated with
finer-grained soils, sandy clay loam (95), deep sand (2) in more open
vegetation, grassland (38), open woodland (51), shaded thickets (8);
supported by limited DBRU collection records: sandy clay loam (6),
sandy loam (4), sand (2) in grassland/pasture (4), shrub/woodland (8).
Food types: subsp. tranvaalensis: No adequate quantitative, data; DBRU
collection records on various dung types: monogastric herbivore: el-
2 mm ephant (4), rhinoceros (2), horse (1), ruminant herbivore: cattle (9),
buffalo (2).
Temporal activity: subsp. tranvaalensis: Flight activity during darkness,
primarily in the summer rainy season (Oct. to May); on Gauteng
bushveld; activity commences early summer (Oct., Nov.) with a mid-
summer peak (Dec. to Feb.) tailing off into late summer (Mar., Apr.);
attracted to light.
Ecoregions Botswana, Zimbabwe: subsp. mesacanthus: N Zambezian
and Mopane Woodlands (AT0725), Southern Miombo Woodlands
(AT0719); marginal in three other ecoregions.
Bioregions South Africa: subsp. tranvaalensis: Centred on Central Bush-
veld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome) with
some records overlapping into the margins of three bioregions of the
Grassland Biome.
Assessment rationale: subsp. mesacanthus: EOO = 378 400 km2 | subsp.
tranvaalensis: EOO = 223 300 km2; AOO perhaps limited by finer-
grained soil association, but widespread in both reserves and farmland
where it is found in both unshaded grassland and open woodland veg-
etation on the dung of farm livestock; assessed as Least Concern (LC).
Conservation measures: subsp. tranvaalensis: Assessment of conservation
status would be improved by quantitative data on food associations; pro-
tected in uMkhuze Game Reserve and Kruger National Park (South Af-
2 mm ♀ rica).

COPRINI
SURICATA 6 (2020) 209

Copris obesus
Boheman, 1857

= Copris contractus Péringuey, 1901 (? pars)

= Copris misellus Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As C. obesa: ‘prope fluvium Limpopo’ [close to Limpo-

po River, southern Africa]; C. contractus: ‘Cape Colony (Uitenhage,
Graham’s Town, Humansdorp, East London), Transvaal (Boksburg),
Southern Rhodesia (Enkeldoorn, Buluwayo)’ [South Africa; Zimbab­
we]; C. misellus: ‘Natal (Durban, Estcourt), Southern Rhodesia (Bulu-
♂
wayo, Salisbury), Transvaal (Waterberg, Potchefstroom)’ [South Africa,
Zimbabwe].
Taxonomy: Accepted Group 31 species but C. contractus: ‘Cape Colo-
ny’ material probably misidentified; secondary type of C. misellus from
Durban (currently a valid species) in Iziko South African Museum ap-
pears to be a minor female specimen of C. obesus; sexually dimorphic
(head, prothoracic disc), prominence of characters varies with body
size.
Distribution: Moist grassland and savanna: SE to East Africa: Lesotho,
South Africa, Zimbabwe, Mozambique, Zambia, Tanzania, Kenya;
black squares are questionable localities.
Locality data (mean ± SD, range): Altitude: 1 237 ± 348, 273–2 392 m;
annual rainfall: 752 ± 139, 377–1 160 mm; annual temperature (max.
+ min. /2): 17.7 ± 2.1, 9.9–23.3°C (N=131).
Habitat: No adequate quantitative assessment; in Abe Bailey Nature Re-
serve: grassland (52), open woodland (34); at Carolina: natural grass-
land (19) fallow crop field (7), improved Kikuyu grass pasture (4);
DBRU collection records indicate bias to coarse-grained soils: clay (1),
sandy clay loam (5), sandy loam (13), sand 13), and open vegetation:
pasture/grassland (21), shrubland (4), open woodland (5).
2 mm
Food types: No quantitative assessment; DBRU collection records pri-
marily from ruminant herbivore dung: cattle (51), buffalo (1), few
from monogastric herbivore dung: rhinoceros (2), zebra (1).
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season (Oct. to May).
Ecoregions Zimbabwe: Centred on Eastern Zimbabwean Montane
Forest Grassland Mosaic (AT1006), Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Centred on warmer and moister vegetation units
in N Mesic Highveld Grassland (Gm) and Sub-Escarpment Grassland
(Gs) (Grassland Biome); also W Lowveld (SVl) and S Central Bushveld
(SVcb) plus N Sub-Escarpment Savanna (SVs) (Savanna Biome).
Assessment rationale: EOO = 1 252 590 km2 (gross estimate); wide-
spread in farmland where it is attracted to the dung of domestic live-
stock in partially shaded and unshaded vegetation; habitat modification
in grassland may reduce population density; transformation extensive
across South African range (Gs, 3–50%; Gm, 6–67%; 0–58% (SVl),
2–49% (SVcb)), but assessed as Least Concern (LC) on basis of large
range.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; effects of grass-
land modification should be further investigated; protected in various
nature reserves: Abe Bailey, Suikerbosrand, Rustenburg (South Africa). ♀
2 mm

COPRINI
210 SURICATA 6 (2020)

Copris orion
Klug, 1835

subsp. orion (Klug, 1835)

subsp. centralis Gillet, 1908
subsp. caffer Gillet, 1908
subsp. overlaeti (Balthasar, 1961b)
No synonyms
Global: LC
Endemic: RSA (subsp. caffer)

J A S O N D J F M A M J

♂ Type localities: C. orion: ‘Senegal’, neotype, same locality; as C. orion var.

centralis: ‘Congo ou d’Afrique orientale’ [Democratic Republic of the
Congo (DRC) or East Africa]; as C. orion var. caffer: ‘Afrique austral’
[South Africa]; C. overlaeti: ‘Lulua: Muteba’ [Democratic Republic of
the Congo (DRC)].
Taxonomy: Accepted Group 38 species comprising four described sub-
spp.; sexually dimorphic (head, prothoracic disc); prominence of char-
acters varies with body size.
Distribution: Three subspp. recorded from West (orion) or central Africa
(overlaeti, centralis); subsp. caffer (red squares): moist, coastal, lowland
distribution from N KwaZulu-Natal to E Eastern Cape, South Africa;
unidentified further subsp. from E Zimbabwe (black squares).
Locality data (mean ± SD, range): subsp. caffer: Altitude: 189 ± 250,
0–984 m; annual rainfall: 806 ± 181, 430–985 mm; annual tempera-
ture (max. + min. /2): 19.4 ± 1.8, 15.7–22.4°C (N=23).
Habitat: subsp. caffer: No quantitative assessment; limited DBRU collec-
tion records on sand (1), sandy loam (2) sandy clay loam (1), entirely
from open grassland (4) or scrub (1) vegetation.
2 mm Food types: subsp. caffer: No quantitative assessment; all DBRU collec-
tion records from cattle dung (5).
Diel activity: subsp. caffer: Flight activity in darkness in the summer rainy
season (Nov. to Mar.).
Bioregions South Africa: subsp. caffer: Centred on Indian Ocean Coastal
Belt across Albany Thicket to E edge of Fynbos Biomes.
Assessment rationale: EOO = 31 800 km2 (subsp. caffer); fairly wide-
spread, but largely restricted to the narrow coastal belt; apparent asso-
ciation with open vegetation and ready colonisation of cattle dung sug-
gests limited threats due to a certain degree of adaptability to farmland
conditions; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
vegetation associations and possible effects of grassland modification;
protected in Vernon Crookes Nature Reserve.

2 mm
♀

subsp. caffer

COPRINI
SURICATA 6 (2020) 211

Copris puncticollis
Boheman, 1857

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Caffraria interiori’ [interior of SE South Africa].

Taxonomy: Accepted Group 40 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Grassland on SE coast deep sands: NE KwaZulu-Natal
(South Africa) at least as far N as Beira (central Mozambique); also
up Zambezi Valley following Zimbabwe/Zambia border to floodplains
of Okavango (N Botswana), Kafue (Zambia) and Ovamboland (N ♂
Namibia).
Locality data (mean ± SD, range): Altitude: 245 ± 384, 0–1 094 m; an-
nual rainfall: 778 ± 186, 428–1 275 mm; annual temperature (max. +
min. /2): 22.3 ± 0.8, 20.4–24.6°C (N=43).
Habitat: At Bushlands Halt: strong bias to deep sands in grassland, sand
(223), clay loam (8), grassland (217), dense woodland (14); on deep
sands in Maputo Special Reserve: natural grassland (62), forest (2); on
sand dunes at Richards Bay: early succession post-mining grassland
(22), late succession woodland / natural dune forest (0); supported
by DBRU collection records: sand (9), finer-grained soils (3), pasture/
grassland (9), woody vegetation (4).
Food types: No quantitative assessment; DBRU collection records pri-
marily on cattle dung (9), elephant dung (2).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to Apr.), but also in the cooler season along the
warmer coast; bias to colonisation of fresh dung (63%, 0–2 day old).
Ecoregions Namibia, Botswana, Mozambique: Centred on grasslands:
Maputaland Coastal Forest Mosaic (AT0119), Southern Zanzibar-
Inhambane Coastal Forest Mosaic (AT0128); also in four isolated ele-
ments of Zambezian Flooded Grasslands (AT0907), Angolan Mopane
2 mm
Woodlands (AT0702).
Bioregions South Africa: Primarily in N Indian Ocean Coastal Belt
Biome (CB) with marginal records in the adjoining S Lowveld (SVl)
(Savanna Biome).
Assessment rationale: EOO = 211 185 km2; widespread, but with AOO
apparently centred on deep sands primarily in open grassland vegeta-
tion; occurrence on floodplains and in post-mining, early succession
grassland suggests an adaptable species; much of the coastal range in
South Africa and SE Mozambique is protected; assessed as Least Con-
cern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on food associations; protected in Tembe
Elephant Park, Sileza Nature Reserve and iSimangaliso Wetland Park
(World Heritage Site) (South Africa); Maputo Special Reserve (Mo-
zambique); and Kafue National Park (Zambia).

♀
2 mm

COPRINI
212 SURICATA 6 (2020)

Copris ritsemae
Harold, 1875

= Copris victorini Boheman, 1857, sensu Péringuey, 1901 (pars)

Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type localities: C. ritsemae: ‘Basutoland (Afric. austr.)’ [Lesotho]; C. vic-

torini sensu Péringuey (pars) (see Taxonomy): ‘Transvaal (Pretoria,
Johannesburg), Natal (Durban), Orange Free State (Parys)’ [Gauteng,
KwaZulu-Natal, Free State; South Africa], also ‘Cape Colony (Knysna,
Seymour)’ [Eastern Cape].
♂ Taxonomy: Accepted Group 18 species, but often confused with close
relatives; type series of synonym probably includes two species plus
C. victorini; Gauteng and Free State localities probably C. ritsemae Har-
old; KwaZulu-Natal locality may be C. vrydaghi Ferreira or C. caelatus
(Fabricius); Eastern Cape localities probably true C. victorini Boheman;
sexually dimorphic (head and prothoracic disc), prominence of charac-
ters varies with body size.
Distribution: Primarily recorded in moist grassland along N edge of
Highveld: South Africa; disjunctions between N, NE and S records
may be real or a collection artefact; S record close to Lesotho type lo-
cality.
Locality data (mean ± SD, range): Average altitude: 1 356 ± 362,
671–1 890 m; average annual rainfall: 724 ± 110, 619–996 mm; av-
erage annual temperature (max. + min. /2): 17.3 ± 2.3, 13.1–20.6°C
(N=10).
Habitat: No quantitative assessment; limited DBRU collection records on
sandy loam (1), sandy clay loam (1) in grassland/pasture (2).
Food types: No quantitative assessment; limited DBRU collection records
on cattle dung (2).
Temporal activity: Diel periodicity unknown, but possibly diurnal like its
close relative, C. caelatus (Fabricius); active primarily during the sum-
2 mm mer rainy season (Oct. to May).
Bioregions South Africa: Straddling N and E edges of Mesic Highveld
Grassland (Gm) (Grassland Biome) and Savanna Biome (grassy areas
in Central Bushveld (SVcb) and Lowveld (SVl)).
Assessment rationale: EOO = 18 840 km2; long series recorded in Johan-
nesburg (1905) and NW border of Lesotho (1970); more recent records
suggest very low population density, perhaps, since much of known
range coincides with highly transformed areas; fairly widespread, but
ecologically poorly known; currently assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
possible effects of habitat transformation; also, a quantitative survey
of the N Highveld is required to determine the full EOO and AOO;
protected in Telperion and Rustenburg nature reserves.

♀
2 mm

COPRINI
SURICATA 6 (2020) 213

Copris sexdentatus
Thunberg, 1818

= Copris lineatus Wiedemann, 1823

= Copris globulipennis Waterhouse, 1891
Global: EN
Endemic: RSA

J A S O N D J F M A M J

Type localities: C. sexdentatus: ‘Majorem horum partem in Capite bonae

spei collegi’ [amongst material mostly collected in the Cape of Good
Hope]; C. lineata: ‘Prom. bon. sp.’ [Cape of Good Hope]; C. globuli-
pennis: ‘Cape of Good Hope’ [South Africa].
Taxonomy: Accepted Group 16 species; sexually dimorphic (head, ♂
prothoracic disc), prominence of characters varies with body size.
Distribution: Only three spot records available, S coastal lowlands of
South Africa; bimodal rainfall region from E Western Cape to W East-
ern Cape.
Locality data (mean ± SD, range): Altitude: 88 ± 84, 16–181 m; annual
rainfall: 427 ± 67, 358–492 mm; annual temperature (max. + min. /2):
17.6 ± 0.5, 17.3–18.2oC (N=3).
Habitat: No quantitative assessment; presumably finer-grained soils in
shrubland vegetation based on a recent record from Renosterveld.
Food types: No quantitative assessment; unknown.
Temporal activity: Flightless and apterous; diel periodicity probably
diurnal; observed walking on the soil surface during daytime; recent
rediscovery during late winter / early spring, end of July.
Bioregions South Africa: W records: critically endangered Eastern Rûens
Shale Fynbos (FRs 13) (Renosterveld Bioregion, Fynbos Biome); E
record possibly inexact: Kowie Thicket (AT 8) (Albany Thicket Biome).
Assessment rationale: EOO = 3 945 km2; represented by few specimens
in collections; not recorded in the past 45 years by collectors from
research groups or Gauteng museums, despite serial collecting in dis-
turbed habitat within the species range; the recent rediscovery (2016)
of one individual in undisturbed Renosterveld (FRs 13) may be the the
only validated spot locality and the first record of the species for over a
century; two other available old records possibly inaccurate, requiring 2 mm

validation; 80% transformation in FRs 13, mostly by arable farming;

7% in AT 8, but species occurrence requires validation; a range of
< 5 000 km2 at < 5 locations with only a single recent record justifies
an EN category, possibly even CR, considering its flightless condition.
Conservation measures: To confirm threat level and conservation status
of this species, a survey of surviving natural shrubland vegetation in
lowland Renosterveld is essential; this should determine the current
EOO, AOO as well as ecological associations; a survey of Albany
Thicket is also necessary to validate or invalidate its occurrence in the
Eastern Cape; now protected in Haarwegskloof Renosterveld Reserve;
status unknown in patches of FRs 13 that are conserved in De Hoop
Nature Reserve and Bontebok National Park.

COPRINI
214 SURICATA 6 (2020)

Copris sphaeropterus
Harold, 1877b

No synonyms
Global: EN
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cap bon. spei.’ [Cape of Good Hope, South Africa].
Taxonomy: Accepted Group 16 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Only one spot record available; S coastal lowlands of South
Africa, winter rainfall region, Western Cape; published records further
♂ eastwards into the bimodal region, E Eastern Cape, remain unvalidated.
Locality data (average): Altitude: 152 m; annual rainfall: 422 mm; annu-
al temperature (max. + min. /2): 16.4°C (N=1).
Habitat: Unknown, but presumably ecologically different from its close
relative C. sexdentatus Thunberg, 1818, which has been recorded from
a validated locality on finer-grained soils in shrubland of the Eastern
Rûens Shale Renosterveld (FRs 13).
Food types: Unknown.
Temporal activity: Flightless and apterous; diel periodicity unknown, but
possibly diurnal like the closely related C. sexdentatus; no dated material
available.
Bioregions South Africa: Critically endangered Central Rûens Shale
Fynbos (FRs 12) (Renosterveld, Fynbos Biome).
Assessment rationale: No EOO estimate possible; represented in collec-
tions by few specimens; only one available relatively exact spot locality
on an old label; none recorded in the past 45 years by collectors from
Gauteng museums or research groups despite serial collecting within
the species range; transformation: FRs 12, 87% by cultivation; a range
of < 5 000 km2 at < 5 locations with no recent records justifies at least
an assessment of Endangered (EN).
2 mm
Conservation measures: A survey of surviving natural shrubland vege-
tation in lowland Renosterveld is essential to determine the ecological
associations, level of threat and conservation status of this species; not
known to be protected within any reserve, but a small amount of FRs
12 is conserved in Agulhas National Park.

2 mm
♀

COPRINI
SURICATA 6 (2020) 215

Copris subsidens
Péringuey, 1901

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Damaraland’ [N Namibia].

Taxonomy: Accepted Group 31 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Arid to dry savanna of the N Namibian Plateau with an
isolated record from the Caprivi Strip.
Locality data (mean ± SD, range): Altitude: 1 366 ± 323, 608–1 998 m; ♂
annual rainfall: 340 ± 72, 171–540 mm; annual temperature (max. +
min. /2): 19.1 ± 2.0, 15.7–23.0°C (N=58).
Habitat: No quantitative assessment; DBRU collection records on var-
ious soils: clay (1), sandy clay loam (13), sandy loam (14), sand (9),
primarily in shrubland (16) and open woodland (17) with only two
observations in grassland.
Food types: No quantitative assessment; DBRU collection records in
farmland on cattle (23) horse (1) and donkey dung (1); also various
dung types in Etosha NP: wildebeest (2), elephant (3), zebra (3).
Temporal activity: Flight activity in darkness in the late summer rainy
season (Dec. to Apr.); attracted to light.
Ecoregions Namibia: S Angolan Mopane Woodlands (AT0702), N Ka-
lahari Xeric Savanna (AT1309), central Namibian Savanna Woodlands
(AT1316).
Assessment rationale: EOO = 105 070 km2; widespread in farmland
where it colonises the dung of domestic livestock; primarily recorded
in natural shrubland and woodland although there is no quantitative
support for such habitat bias; currently assessed as Least Concern (LC)
on basis of large EOO. 2 mm
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; it is necessary to
investigate possible effects on population density of clearance of woody
vegetation; protected in Etosha National Park.

♀ 2 mm

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Copris urus
Boheman, 1857

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘prope fluvium Limpopo’ [close to Limpopo River, south-

ern Africa; ?near the Indian Ocean river mouth].
Taxonomy: Accepted Group 48 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Known only from the deep coastal sands of the Maputaland
Centre of Endemism, particularly in NE KwaZulu-Natal, South Africa,
♂ less frequently in drier SE Mozambique.
Locality data (mean ± SD, range): Altitude: 42 ± 56, 0–273 m; annual
rainfall: 847 ± 116, 682–1 006 mm; annual temperature (max. + min.
/2): 21.9 ± 0.5, 20.5–22.9°C (N=25).
Habitat: At Bushlands Halt: restricted to deep sands in both grassland and
woodland, sand (77), clay loam (0), grassland (43), dense woodland
(34); across a restoration gradient on coastal dunes at Richards Bay:
mainly in early successional grassland (19) and open late successional
woodland (5); rare in dense, early succession woodland (0) and natural
dune forest (1); DBRU collection records only on sand (4).
Food types: No quantitative assessment; DBRU collection records on
cattle (4) and elephant (1) dung.
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season (Oct. to Jan.), but also the cooler season (June, Aug.)
owing to its warm coastal range; bias to colonisation of older dung
(24%, 0–2 day old; 48%, 2–4 day).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 8 690 km2; relatively small EOO in a re-
5 mm gion comprising a mosaic of soil and vegetation types; AOO would be
smaller due to soil specialisation; possibly influenced by a bias to open
vegetation at the cooler coast compared to a more generalist distribu-
tion between open and dense vegetation types in warmer inland situa-
tions; however, assessed as Least Concern (LC) rather than Vulnerable
(VU) owing to the level of protection over much of its range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on food associations; much of the
EOO protected in the iSimangaliso Wetland Park (World Heritage
Site) (South Africa); however, as regards the AOO, a more accurate
assessment of the conservation status of this species might result from
an investigation of the relative importance of vegetation types with
increasing proximity to the coastline, particularly that of grassland on
fossil lagoons.

♀
5 mm

COPRINI
SURICATA 6 (2020) 217

Copris victorini
Boheman, 1857

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Caffraria meridionali….et ad Promont. Bonae Spei’

[southern SE South Africa to Cape of Good Hope].
Taxonomy: Accepted Group 17 species; sexually dimorphic (head and
prothoracic disc), prominence of characters varies with body size.
Distribution: Restricted to a small known range along the S coast and S
coastal mountains of South Africa in the winter and bimodal rainfall ♂
regions.
Locality data (mean ± SD, range): Altitude: 437 ± 350, 0–822 m; annual
rainfall: 425 ± 47, 363–465 mm; annual temperature (max. + min. /2):
16.1 ± 1.5, 14.4–18.0°C (N=4).
Habitat: No quantitative assessment; no soil type data, but known mainly
from sandy bioregions; recorded in fynbos shrubland (top of Dutoits­
kloof Pass near Paarl).
Food types: No quantitative assessment; recorded from a klipspringer
midden (top of Dutoitskloof Pass near Paarl).
Temporal activity: Unknown, possibly diurnal like one other member of
its species group; recorded in early summer (Nov., Dec.).
Bioregions South Africa: Vegetation units of Sandstone Fynbos (FFs),
Granite Fynbos (FFg), and Shale Fynbos (FFr).
Assessment rationale: EOO = 2 100 km2; currently assessed as Data
Deficient (DD) due to limited data, but on the basis of a known
range of < 5 000 km2 and records from less than five locations, this
species currently deserves Endangered (EN) status; however, there is
low transformation around two data points in the extensively protected
(78–86%) W vegetation units: Hawequas Sandstone Fynbos (FFs 10,
3%), South Langeberg Sandstone Fynbos (FFs 16, 4%); whereas there
2 mm
is high transformation in the E: Garden Route Granite Fynbos (FFg 5,
70%), Garden Route Shale Fynbos (FFh 9, >50%).
Conservation measures: To determine the conservation status of this
species, a survey of S mountain and S coastal habitats is necessary to as-
certain the full extent of its EOO and AOO, its ecological associations,
and the number of occupied locations; protected in Grootvadersbosch
Nature Reserve.

2 mm
♀

COPRINI
218 SURICATA 6 (2020)

Copris vilhenai
Ferreira, 1962a

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Província de Angola: n.o ANG.: 10609, Marco de Canav-

ezes (Membassoco), proximo da Central Hidroelectrica da Companhia
de Acucar de Angola’ [Membassoco, Angola].
Taxonomy: Accepted Group 41 species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size.
Distribution: Patchy occurrence across extensive areas and smaller pock-
♂ ets of deep sands in N South Africa, N Botswana, S Angola, Zimbabwe,
Mozambique.
Locality data (mean ± SD, range): Altitude: 867 ± 420, 55–1 516 m;
annual rainfall: 630 ± 121, 430–1 136 mm; annual temperature (max.
+ min. /2): 20.8 ± 1.6, 17.2–25.2°C (N=43).
Habitat: In Gauteng (cited as C. macer): extreme association with coarse-
grained soils, deep sand (19), sandy clay loam (0) in woody vegetation,
grassland (1), open woodland (4), shaded thickets (14); partly support-
ed by DBRU collection records: sand (12), sandy loam (7), clay (2) in
grassland (2), shrub/woodland (17), forest (2).
Food types: In Botswana: bias to elephant (9) rather than cattle (2), pig,
sheep dung or carrion (all 0); DBRU collection records shared between
monogastric herbivore: elephant (7), rhinoceros (1), and ruminant her-
bivore dung: cattle (9), buffalo (2), wildebeest (1).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to May) with additional cool season activity
along the warmer coastline (July, Aug.); on Gauteng bushveld: seasonal
peak during early summer (Oct. to Dec.).
Ecoregions Botswana, Zimbabwe: Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea
Woodlands (AT0726); margins of two other ecoregions.
2 mm Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome).
Assessment rationale: EOO = 677 310 km2; AOO probably smaller due
to sand specialisation; bias to monogastric dung presumably results in
lower population density in farmland although it does colonise cattle
dung; association with woody vegetation suggest its clearance would
also reduce population density; transformation across South African
range, 0–58% (SVl), 2–49% (SVcb); currently assessed as Least Con-
cern (LC) on basis of widespread range.
Conservation measures: Assessment of conservation status would be
improved by more comprehensive data on ecological associations,
particularly responses of population density to loss of elephants and
clearance of woodland on sandy soils; protected in Kruger National
Park, Leeuwfontein and Tswaing nature reserves (South Africa); Chobe
National Park (Botswana).

2 mm
♀

COPRINI
SURICATA 6 (2020) 219

Copris vrydaghi
Ferreira, 1962b

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Estcourt (N.)’ [Estcourt, KwaZulu-Natal, South Africa].

Taxonomy: Accepted Group 18 species, but often confused with close
relatives; two female individuals from non-type localities require val-
idation (N and S limits of range on map); sexually dimorphic (head
and prothoracic disc), prominence of characters varies with body size.
Distribution: Moist grassland below central E escarpment: KwaZulu- ♀
Natal, South Africa.
Locality data (mean ± SD, range): Average altitude: 1 070 ± 183,
897–1 307 m; average annual rainfall: 758 ± 28, 729–794 mm; average
annual temperature (max. + min. /2): 16.9 ± 1.4, 15.0–18.0°C (N=4).
Habitat: No quantitative assessment; not known.
Food types: No quantitative assessment; not known.
Temporal activity: Diel periodicity unknown, but possibly diurnal like its
close relative, C. caelatus (Fabricius); active during the summer rainy
season (Nov.).
Bioregions South Africa: Centred on Sub-Escarpment Grassland (Gs)
(Grassland Biome).
Assessment rationale: EOO = 5 085 km2 (probably underestimated);
known by very few individuals in reference collections with some re-
quiring validation; ecologically extremely poorly known, but appears
to occupy a grassland range at lower altitude parallel to that of the
central range of its close relative, C. caelatus (Fabricius, 1794); at pres-
ent, would qualify for an IUCN threat category owing to small known
range at few localities; however, paucity of data is likely a collection
artefact, so currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative sur- 5 mm

vey is required starting in the N Sub-Escarpment Grassland spreading

to the S and to higher and lower altitude; this should determine the
EOO, AOO, ecological associations and possible effects of habitat
transformation; possibly protected at the iSandlwana Heritage Reserve
(KwaZulu-Natal).

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220 SURICATA 6 (2020)

Genus Heliocopris Hope, 1837

Type species and designation: Scarabaeus gigas Linnaeus, 1758, by subsequent designation (Löbl 2006).
Synonyms: None.
Last review: Entire genus reviewed by Pokorný et al. (2009).

The long-established genus, Heliocopris Hope, 1837, was created to accommodate four previously described species. It is
currently represented by 54 valid species occurring in the Afrotropical (50) and Oriental regions (4) with a fossil known
from the Palaearctic region (Japan).

The genus is divided into five species groups (Pokorný et al. 2009), four of which are represented by seven species in South
Africa, Botswana and Namibia. Validation is required concerning records for other species. In South Africa, records for
H. hamifer Harold, 1878 may represent rare occurrence; records for H. colossus Bates, 1868, may represent past occurrence
or misidentification of a rare close relative. In Namibia, records for H. biimpressus Kolbe, 1893b, may represent misden-
tification.

Heliocopris comprises very large-bodied species, whose nesting behaviour requires large amounts of dung. Even so, some
taxa occur in dry or arid areas where large herbivores also occur. A relatively shallow tunnel is constructed from under
a dropping into which dung is rapidly buried. In some species the dung is relocated into a deeper tunnel. A number of
broods is constructed from the buried dung cake within a chamber at the end of the tunnel.

Heliocopris species are macropterous, flying during darkness. Three of the species are restricted to southern Africa whereas
four species show distributions extending into tropical Africa. All are essentially centred on arid to moist savanna with
three primarily recorded in the east on sandy or finer-grained soils. The others occur in both the east and the west with
two rare or absent from the intervening Kalahari deep sands and two also present in the Kalahari. Despite their large body
size, all species are assessed as Least Concern (LC) as they are widespread on farms and in reserves.

In southern Africa
Group 1 (neptunus group): One species centred on finer-grained soils in the east.
Group 2 (hamadryas group): Four mostly widespread species, three in east and west savanna with two also in the Kala-
hari, one primarily centred in the east.
Group 4 (bucephalus group): One species centred on finer-grained soils in the east.
Group 5 (gigas group): One widespread species associated with monogastric herbivore dung mostly in arid to dry
savanna.

COPRINI
SURICATA 6 (2020) 221

Heliocopris andersoni
Bates, 1868

= Heliocopris isidis Boheman, 1857

= Heliocopris coriaceus Shipp, 1897a
Global: LC

J A S O N D J F M A M J

Type localities: ‘Prope Lacum N’Gami’ [close to Lake Ngami, N Botswa-

na]; H. isidis: ‘Caffraria tota’ [all southern/South Africa]; H. coriaceus:
‘Amarato to Lake Stephanie’ [untraced locality of ‘Amarato’ to Lake
Chew Bahir, S Ethiopia].
Taxonomy: Accepted gigas group species; sexually dimorphic (head,
prothoracic disc), prominence of characters varies with body size. ♂
Distribution: Mostly dry savanna from S to NE Africa; South Africa,
Namibia, Botswana, Zimbabwe, Kenya; also reported from Angola,
Malawi, Mozambique, Democratic Republic of the Congo (DRC),
Tanzania, Ethiopia, Eritrea.
Locality data (mean ± SD, range): Altitude: 1 057 ± 440 2–1 815 m;
annual rainfall: 567 ± 192, 48–988 mm; annual temperature (max. +
min. /2): 19.6 ± 2.0, 15.0–23.6°C (N=103).
Habitat: No quantitative assessment; DBRU collection records from
sand (5), sandy loam (8), sandy clay loam (6), clay (1) in grassland (2),
scrub/shrubland (4), open woodland (11).
Food types: No adequate quantitative assessment; at Phalaborwa: limited
numbers sampled to dung of elephant (3) and pig (1), none to cattle
(0); DBRU collection records from dung of elephant (5), rhinoceros
(3), donkey (2), cattle (10), buffalo (1), wildebeest (1).
Temporal activity: Flight activity during darkness in the mid and late
summer rainy seasons (Nov. to Apr.);
Ecoregions Namibia, Botswana, Zimbabwe: Mainly Namibian Savanna
5 mm
Woodlands (AT1316), Angolan Mopane Woodlands (AT0702), NE
Kalahari Xeric Woodlands (AT1309), Zambezian and Mopane Wood-
lands (AT0725), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Mainly Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp) (Savanna Biome); also scattered in Eastern
Kalahari Bushveld (SVk) (Savanna Biome), N Dry Highveld Grassland
(Gh) (Grassland Biome); Albany Thicket Biome (AT).
Assessment rationale: EOO = 3 715 845 km2 (gross estimate); limit-
ed data may point to association with monogastric herbivore dung
although H. andersoni is widespread in both reserves and farmland;
recorded primarily on monogastric herbivore dung in reserves, from
cattle and equid dung on farms; assessed as Least Concern (LC) on
basis of large range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
better support for association with monogastric herbivore dung and ef-
fects on population density outside of reserves; protected in many game
reserves and national parks including Kruger (South Africa), Etosha
(Namibia), Lake Mutirikwe (Zimbabwe), Masai Mara (Kenya).

♀ 5 mm

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222 SURICATA 6 (2020)

Heliocopris atropos
Boheman, 1860

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘prope lacum N’Gami’ [close to Lake Ngami, N Botswana].

Taxonomy: Accepted hamadryas group species, but NE range edge uncer-
tain due to possible confusion with close relatives in E Africa; sexually
dimorphic (head, prothoracic disc), prominence of characters varies
with body size.
Distribution: Dry Southern African savanna with a range that may extend
♂ to E Africa: South Africa, Namibia, Botswana, Zimbabwe, Angola; also
reported from Malawi, Mozambique, Zambia; reports from Tanzania,
Kenya require validation.
Locality data (mean ± SD, range): Altitude: 1 113 ± 323, 55–1 628 m;
annual rainfall: 493 ± 149, 171–813 mm; annual temperature (max. +
min. /2): 20.0 ± 1.7, 16.3–22.5°C (N=44).
Habitat: No adequate quantitative assessment; in open woodland of
Leeuw­fontein Nature Reserve: sand (3), sandy loam (5); DBRU col-
lection records from sand (8), sandy loam (3) in grassland (5). open
shrub/woodland (6).
Food types: No adequate quantitative assessment; on deep sands in Bot­
swana: weak bias to monogastric herbivore dung: pig (1), elephant (7),
cattle (1), sheep (1); DBRU collection records from dung of cattle (8),
wildebeest (1), elephant (4).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May).
Ecoregions Namibia, Botswana, Zimbabwe: Primarily Kalahari Xeric
Savanna (AT1309), Southern African Bushveld (AT0717), Zambezian
and Mopane Woodlands (AT0725); also Kalahari Acacia-Baikiaea
5 mm Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 602 870 km2 (gross estimate; range to E
Africa?); AOO would be smaller if bias to sandy soils and monogastric
herbivore dung is supported by quantitative data; however, large range
across both farmland and reserves would justify an assessment as Least
Concern (LC), although verging on Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by more comprehensive quantitative data on ecological as-
sociations and a survey to more accurately determine NE range limits;
protected in uMkhuze Game Reserve (South Africa), Chobe, Hwange
and Bicuar national parks (Botswana, Zimbabwe, Angola).

2 mm
♀

COPRINI
SURICATA 6 (2020) 223

Heliocopris faunus
Boheman, 1857

= Heliocopris satyrus Boheman, 1860

= Heliocopris operosus Waterhouse, 1891
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: ‘prope fluvium Gariep’ [close to Orange River, South

Africa]; H. satyrus: ‘prope lacum N’Gami’ [close to Lake Ngami, N
Botswana]; H. operosus: ‘Africa’.
Taxonomy: Accepted hamadryas group species; sexually dimorphic (head,
prothoracic disc); prominence of characters varies with body size.
Distribution: Somewhat patchy recorded pattern circumscribing the Ka- ♂
lahari to W, S and E: Namibia, E Botswana, S Zimbabwe, South Africa;
unclear if this pattern is truly disjunct or a collecting artefact; higher
population density on dry W uplands (N Namibian plateau, W es-
carpment edge) and South African Upper Karoo than in lower-lying E
savanna where it occurs as a rarity; also reported from Mozambique; no
recent Lake Ngami record (20.500S 22.667E), so not plotted on map.
Locality data (mean ± SD, range): Altitude: 1 177 ± 349, 55–1 928 m;
annual rainfall: 376 ± 162, 103–900 mm; annual temperature (max. +
min. /2): 18.4 ± 2.6, 13.3–23.0°C (N=69).
Habitat: No quantitative assessment; DBRU collection records from sand
(4), sandy loam (5), sandy clay loam (2), clay (1) in pasture (1), scrub/
shrubland (5), open woodland (6).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (9), zebra (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Mostly Namibian Savanna 5 mm
Woodlands (AT1316), NW Kalahari Xeric Woodlands (AT1309), An-
golan Mopane Woodlands (AT0702); uncommon in Southern African
Bushveld (AT0717), S Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Most records from Central Bushveld (SVcb),
Lowveld (SVl) (Savanna Biome); Upper Karoo (NKu) (Nama Karoo
Biome).
Assessment rationale: EOO = 391 450 km2; possible bias to sandy soils
and low or open woody vegetation would reduce AOO; unclear if clear-
ance of woody vegetation would influence population density; patchy,
but widespread occurrence in farmland and reserves; readily attracted
to cattle dung; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations and a survey
to determine if patchiness is real or a collecting artefact; protected in
Etosha National Park (Namibia).

5 mm

COPRINI
224 SURICATA 6 (2020)

Heliocopris hamadryas
(Fabricius, 1775)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Scarabaeus hamadryas: ‘Cap. B. S.’ [Cape of Good Hope,

South Africa].
Taxonomy: Accepted hamadryas group species; sexually dimorphic (head,
prothoracic disc); prominence of characters varies with body size.
Distribution: Primarily moist upland grassland and savanna in S, E and
W central Africa, but also coastal in S Africa: South Africa: Zimbabwe,
♂ Malawi, Angola, Mozambique, Zambia, Kenya, SE Nigeria; outliers
in Botswana, Namibia; also reported from Democratic Republic of
the Congo (DRC), Rwanda, Tanzania, Uganda; records from Eritrea,
Ethiopia, Somalia require validation.
Locality data (mean ± SD, range): Altitude: 1 068 ± 405, 0–1 800 m;
annual rainfall: 678 ± 162, 326–1 232 mm; annual temperature (max.
+ min. /2): 18.6 ± 1.8, 13.7–23.0°C (N=155).
Habitat: In Gauteng bushveld: bias to deep sand (71) rather than sandy
clay loam (14) and to shaded thickets (41) rather than open woodland
(29), grassland (15); DBRU collection records across a wider range
suggest soil generalisation: sand (9), sandy loam (9), sandy clay loam
(11), clay (1) mainly in open vegetation: grassland/pasture (19), open
shrub/woodland (4).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (32), also elephant (1), rhinoceros (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May); attracted to light.
5 mm Ecoregions Namibia, Botswana, Zimbabwe: Primarily Southern Miom-
bo Woodlands, also Zambezian and Mopane Woodlands (AT0725);
marginal in two other ecoregions.
Bioregions South Africa: Dry Highveld Grassland (Gh), Sub-Escarpment
Grassland (Gs) (Grassland Biome); Eastern Kalahari Bushveld (SVk),
Central Bushveld (SVcb), Lowveld (SVl) (Savanna Biome); Indian
Ocean Coastal Belt Biome (CB); transformed habitats of Albany
Thicket and E Fynbos Biomes.
Assessment rationale: EOO = 3 416 220 km2; (gross, probable under-
estimate); AOO much smaller mostly coinciding with moist uplands
and plateaux in the tropics; conflicting data for habitat associations,
but widespread in reserves and farmland across a large range; readily
attracted to cattle dung; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations,
particularly an investigation to determine if vegetation bias to shade
in Gauteng bushveld might reflect seasonal heat in summer activity
period; protected in various reserves including Leeuwfontein (South
Africa), Lake Mutirikwe (Zimbabwe), Masai Mara (Kenya).

5 mm

COPRINI
SURICATA 6 (2020) 225

Heliocopris japetus
Klug, 1855

= Heliocopris bicarinulatus Boheman, 1860

Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Copris Japetus: ‘Sena’ [Sena, Central Mozambique]; as

H. bicarinulata: ‘juxta lacum N’Gami’ [near Lake Ngami, N Botswa-
na].
Taxonomy: Accepted hamadryas group species; sexually dimorphic (head,
prothoracic disc); prominence of characters varies with body size.
Distribution: Dry sandy savanna from S to E Africa: South Africa, Na-
mibia, Botswana, Zimbabwe, Mozambique, Angola, Zambia, Kenya; ♂
also reported from Democratic Republic of the Congo (DRC), Tanza-
nia; Rwanda and some DRC reports possibly represent confusion with
close relative, H. antenor (Olivier, 1789).
Locality data (mean ± SD, range): Altitude: 1 020 ± 361 11–1 757 m;
annual rainfall: 569 ± 178, 208–1 247 mm; annual temperature (max.
+ min. /2): 20.5 ± 1.7, 16.7–25.2°C (N=137).
Habitat: On Gauteng bushveld: extreme bias to deep sand (26) as op-
posed to sandy clay loam (0) with a slight bias to open woodland (14)
as opposed to grassland (8) or shaded thickets (4); DBRU collection
records support a bias to sand (23) as opposed to sandy loam (4), sandy
clay loam (1), clay (2) in grassland/pasture (10), shrubland (3), open
woodland (15).
Food types: On deep sand in Botswana: sampled only to dung with a
bias to that of omnivores and monogastric herbivores: carrion (chicken
livers – 0), dung of pig (28), elephant (27), cattle (11), sheep (10);
DBRU collection records reflect commonly sampled dung types: cattle
(29), elephant (5).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna 5 mm

(AT1309), Angolan Mopane Woodlands (AT0702), Southern African

Bushveld (AT0717), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726);
margins of two other ecoregions.
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome);
margins of two other bioregions.
Assessment rationale: EOO = 3 260 895 km2 (gross estimate); AOO
would be restricted by sand specialisation; however, widespread in
southern Africa where deep sands cover large areas; frequently recorded
in both open and woody vegetation on farms and in reserves, especially
at the fringes of the Kalahari; assessed as Least Concern (LC).
Conservation measures: None recommended; protected in various na-
tional parks, including Bicuar, Chobe, Hwange, Gorongosa, Kruger
(respectively, Angola, Botswana, Zimbabwe, Mozambique, South Af-
rica).

♀ 5 mm

COPRINI
226 SURICATA 6 (2020)

Heliocopris neptunus
Boheman, 1857

= Heliocopris eryx Fabricius, 1801, sensu Péringuey, 1901

Global: LC

J A S O N D J F M A M J

Type localities: ‘juxta fluvium Limpopo’ [near Limpopo River, southern

Africa]; H. eryx sensu Péringuey: ‘Transvaal (Lydenburg), Southern
Rhodesia (Mazoe, Buluwayo)’ [Mashishing, Mpumalanga, South Af-
rica; Mazoe and Bulawayo, Zimbabwe].
Taxonomy: Accepted neptunus group species; sexually dimorphic (head,
prothoracic disc); prominence of characters varies with body size.
♂ Distribution: Moist southern African savanna: South Africa, Eswatini,
NE Botswana, Zimbabwe, Mozambique; one outlier record in Namib-
ia; also reported from Zambia; record from Ethiopia would represent
confusion with a close relative.
Locality data (mean ± SD, range): Altitude: 824 ± 448, 46–1 746 m;
annual rainfall: 653 ± 128, 229–1 013 mm; annual temperature (max.
+ min. /2): 19.9 ± 1.9, 14.0–24.5°C (N=83).
Habitat: No quantitative assessment; DBRU collection records only from
finer-grained soils: sandy loam (4), sandy clay loam (3), clay (6) in
grassland/pasture (4), open shrub/woodland (5).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (15).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: Southern African Bushveld
(AT0717), S Zambezian and Mopane Woodlands (AT0725), South-
ern Miombo Woodlands (AT0719); scattered records in three other
ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome).
5 mm Assessment rationale: EOO = 328 720 km2; (presumably underesti-
mated); AOO would be restricted by soil specialisation, but requires
quantitative support; however, finer-grained soils widespread in both
reserves and farmland where it is readily attracted to cattle dung; as-
sessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by a quantitative survey to confirm its full EOO, AOO and
ecological associations; protected in Kruger National Park, Ithala Game
Reserve (South Afrca).

5 mm
♀

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SURICATA 6 (2020) 227

Heliocopris pirmal
(Fabricius, 1798)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Copris pirmal: ‘India orientali’ [ascribed to East India

in error].
Taxonomy: Accepted bucephalus group species; identity of type requires
validation; sexually dimorphic (head, prothoracic disc); prominence of
characters varies with body size.
Distribution: Moist southern African savanna: South Africa, Eswatini,
Zimbabwe; quite possibly Zambia; reports from Angola, Namibia, Bot­ ♂
swana, Mozambique require validation; SW outlier localities require
further support.
Locality data (mean ± SD, range): Altitude: 1 085 ± 362, 143–1 558 m;
annual rainfall: 702 ± 113, 433–1 010 mm; annual temperature (max.
+ min. /2): 18.7 ± 1.6, 15.7–23.2°C (N=64).
Habitat: No quantitative assessment; very limited DBRU collection re-
cords from sandy clay loam (3) in grassland (1) or woodland (3).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (1), buffalo (1), rhinoceros (4).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to May); attracted to light.
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719), Zam-
bezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Particularly, moister parts of Central Bushveld
(SVcb) and Lowveld (SVl) (Savanna Biome) along the Waterberg and
edge of Highveld; scattered records in four other bioregions.
Assessment rationale: EOO = 236 025 km2 (probably underestimated);
AOO may be smaller due to finer-grained soil and woodland bias, but
requires quantitative support; relatively poorly known species ecolog-
ically; assessed as Least Concern (LC) due to widespread distribution, 5 mm
but verging on Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, particularly if
population density might be influenced by clearance of woodland on
finer-grained soils; improved survey data is also required to confirm its
EOO and AOO; protected in Lake Mutirikwe Game Reserve (Zimba-
bwe), Hluhluwe–iMfolozi Game Reserve (South Africa).

♀ 5 mm

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228 SURICATA 6 (2020)

Genus Litocopris Waterhouse, 1891

Type species and designation: Litocopris punctiventris Waterhouse, 1891, by subsequent designation (Ferreira 1960a).
Synonyms: None.
Last review: All species listed by Ferreira (1972).

Litocopris Waterhouse, 1891, was created as a subgenus of Copris to accommodate the type species and two others pre-
viously described under Copris. One further species was added at a later date. Although recently raised to generic level,
this status has been questioned. After having being treated as a full genus (Branco 2011), it was synonymised with Copris
(Marchisio & Zunino 2012), but has, thereafter, been listed as a genus by Catalogue of Life (Schoolmeesters 2017). Here
we follow Branco (2011), although the taxon may show affinities to Copris Group 9, which is one of the more basally
derived elements in the genus Copris.

Litocopris are macropterous with similar body size to the smaller-bodied group of Copris. They construct tunnels from un-
der droppings, but nesting behaviour is essentially unknown. One species has often been observed in older dung. The two
species represented in South Africa comprise an endemic centred on the SE coast and SE escarpment and another centred
on moist savanna of NE KwaZulu-Natal, which is distributed northwards into the E tropics.

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Litocopris muticus
(Boheman, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Copris mutica: ‘Caffraria interiori’ [interior of SE South

Africa].
Taxonomy: Accepted species, but see notes on generic status in genus
section.
Distribution: Moist savanna on E seaboard of Africa: NE South Africa,
Zimbabwe, Malawi, Mozambique, Kenya; also reported from Demo-
cratic Republic of the Congo (DRC), Rwanda, Burundi, Uganda, plus
Côte d’Ivoire in W Africa.
Locality data (mean ± SD, range): Altitude: 257 ± 240, 59–808 m; an-
nual rainfall: 912 ± 144, 697–1 177 mm; annual temperature (max. +
min. /2): 22.2 ± 1.8, 20.5–25.2°C (N=11).
Habitat: No quantitative assessment; DBRU collection records biased to
finer-grained soils: clay (3), sandy clay loam (3) sandy loam (2), sand
(1); in shaded or partially shaded vegetation: forest (2), savanna wood-
land (10), grassland (1).
Food types: No quantitative assessment; DBRU collection records biased
to monogastric herbivore dung: elephant (8), rhinoceros (4) with one
record from human/cattle dung bait.
Temporal activity: Unknown; possibly flies during darkness, primarily
during the summer rainy season (Oct. to May).
Ecoregions Zimbabwe: N Southern Miombo Woodlands (AT0719), wet
areas in Zambezian and Mopane Woodlands (AT0725) (N Zimbabwe,
not shown on map).
Bioregions South Africa: Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 793 435 km2 (underestimated); AOO
very likely much smaller as all DBRU collection records are from game
reserves or protected areas (15); apparently somewhat specialised as
regards soil, vegetation and dung type association, but assessed as Least
Concern (LC) on the basis of widespread range in various reserves.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, particularly as its 2 mm
status in South Africa would be dependent on continuing protection of
both moist woodland vegetation on finer-grained soils and monogastric
herbivores, particularly elephant and rhinoceros; protected in Hluhluwe–
iMfolozi Game Reserve (South Africa), Gorongosa National Park (Mo-
zambique), Masai Mara Game Reserve (Kenya).

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Litocopris simplex
(Harold, 1868)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Copris simplex: ‘Caffraria’ [SE South Africa].

Taxonomy: Accepted species, but see notes on generic status in genus
section.
Distribution: Moist SE Highveld and S to SE coastal regions of South
Africa.
Locality data (mean ± SD, range): Altitude: 695 ± 625, 0–1 782 m;
annual rainfall: 684 ± 164, 429–965 mm; annual temperature (max. +
min. /2): 16.7 ± 2.1, 13.2–21.7°C (N=45).
Habitat: No quantitative assessment; DBRU collection records show a
bias to finer-grained soils: clay (1), sandy clay loam (8), sandy loam
(9); in grassland/pasture (14), shrub/woodland (2); also crop fields (2).
Food types: No quantitative assessment; all DBRU collection records
from cattle dung (23).
Temporal activity: Unknown; possibly flies during darkness in the sum-
mer rainy season on the Highveld (Oct. to Mar.) with some activity
in autumn and spring (Aug., Sept., May) along the warmer coastline.
Bioregions South Africa: SE Mesic Highveld Grassland (Gm) across
Sub-Escarpment Grassland (Gs) to S outlier of Drakensberg Grasslands
(Gd) (Grassland Biome); also Indian Ocean Coastal Belt Biome (CB)
to NE margin of Albany Thicket Biome (AT); outlier cluster at W ex-
treme of range coincides with high rainfall units of Sandstone Fynbos
(FFs) or Southern Afrotemperate Forest (FOz 1).
Assessment rationale: EOO = 85 760 km2; except for two woody local-
ities at margins of Albany Thicket, DBRU records entirely from moist
grassland or pasture on cattle dung whereas a number of non-DBRU
records are from forest stations, pasture (3) or a fallow crop field (1);
records for the outlier cluster in FFs and FOz 1 may merely infer tol-
erance of habitat modification, or it may infer range expansion into
the Eastern Cape in response to clearance of shrubland and forest;
2 mm
nevertheless, currently assessed as Least Concern (LC) on the basis of
widespread range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; particularly,
a survey to examine habitat bias between forest and grassland in wet
areas partly cleared of natural vegetation in the Eastern Cape; protected
in Coleford Nature Reserve, but possibly poorly conserved over most
of its range.

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SURICATA 6 (2020) 231

Genus Macroderes Westwood, 1842

Type species and designation: Onthophagus greeni Kirby, 1818, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Frolov and Scholtz (2004) and Abdalla et al. (2018) with
synonymies and descriptions of new species.

The long-established genus, Macroderes Westwood, 1842, was created for a single species previously described under
Onthophagus. After the revision of Frolov & Scholtz (2004), a total of 14 valid species was known with a further two names
whose status could not be determined. More recent work has added a further seven new taxa and a further, now validated,
old species name. Descriptions of these species were published too late to be included in the current species accounts, but
see Preamble. Macroderes is endemic to the winter and bimodal rainfall regions of SW Africa where all but one species oc-
cupy relatively small, mostly allopatric ranges (Frolov & Scholtz 2004) in the winter rainfall region. Currently, all validated
records occur entirely within the borders of South Africa.

Although there is no phylogeny (at the time of writing), morphological relationships between 13 species were indicated
in the revision of Frolov and Scholtz (2004). These suggest four species groups showing E–W or N–S patterns of distri-
bution. Two patterns extend along W coastal dunes comprising two and four species each. Another group of three species
ranges from W to SE lowlands across the Cederberg. The remaining group of four species shows a disjunct pattern on
cooler uplands with centres in Namaqualand, Cape Peninsula and the S Cape. Field observations suggest species-specific
associations with either coarse or finer-grained soils.

All Macroderes species lack a humeral umbone, which is indicative of aptery and flightlessness and may account for the
small species ranges. They are assumed to show tunnelling habits, but nesting behaviour is unknown. Some species have
been sampled to dung-baited traps set in the late afternoon and retrieved the following morning. Although they were,
thus, assumed to forage primarily in darkness, diurnal activity has been observed under particularly cold conditions. Thus,
the dominant diel periodicity is uncertain.

Reference material and ecological information is comparatively limited, leading to assessments of Data Deficient (DD)
for 12 species and Near Threatened (NT) for the remaining two. Observations made during field sampling suggest a bias
to dense vegetation. Thus, habitat degradation, fragmentation and transformation would strongly influence conservation
status, particularly considering their flightless condition. Overgrazing and mining are threats to species with N distribu-
tions although transformation is limited. Those in the more highly transformed SW and S would face greater threats.
Irrespective of distributional bias, all but one species may deserve Vulnerable (VU) or even Endangered (EN) status on the
basis of aptery and small known EOO averaging 2 325 km2 and ranging from 53 to 7 940 km2.

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Macroderes amplior
Frolov & Scholtz, 2004

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘5 km W of Rietpoort’ [Namaqualand, Northern Cape,

South Africa].
Taxonomy: Accepted species.
Distribution: Lower edge, W escarpment, arid NW South Africa.
Locality data (mean ± SD, range): Altitude: 388 ± 85, 308–476 m; an-
nual rainfall: 134 ± 27, 103–154 mm; annual temperature (max. +
min. /2): 17.6 ± 0.6, 17.0–18.2°C (N=3).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam in mountain Karoo shrubland.
Food types: No quantitative assessment; no recorded data.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug., Sept.), which may limit distri-
bution to warmer, lower areas.
Bioregions South Africa: W Namaqualand Klipkoppe Shrubland (SKn
1) (Namaqualand Hardeveld Bioregion, Succulent Karoo Biome).
Assessment rationale: EOO = 1 170 km2; known from only three local-
ities in the SKn 1 vegetation unit where there is little habitat transfor-
mation as it is characterised by steep rocky habitats; however, although
it covers about 9 900 km2, much lies above 500 m and falls within the
range of M. mutilans Kolbe, 1908; M. amplior is virtually unknown
ecologically and is assessed as Data Deficient (DD), but, on the basis
of flightlessness and very small known range, it would qualify for an
IUCN threat category of at least Vulnerable (VU).
Conservation measures: To assess conservation status, a quantitative
2 mm
survey is required to determine the full EOO, AOO and ecological as-
sociations in SKn 1 and adjacent vegetation units; not currently known
from any protected area.

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SURICATA 6 (2020) 233

Macroderes arrowi
Janssens, 1939a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Rhynsdorp’ [Vanrhynsdorp, Western Cape, South Africa].

Taxonomy: Accepted species.
Distribution: Recorded only on arid red sands in the vicinity of the type
locality at Vanrhynsdorp, W coastal plain, South Africa.
Locality data (mean ± SD, range): Altitude: 218 ± 93, 152–283 m; an-
nual rainfall: 161 ± 11, 153–169 mm; annual temperature (max. +
min. /2): 18.7 ± 0.5, 18.4–19.1°C (N=2).
Habitat: No quantitative assessment; but one exact locality on deep red
sands in scrub.
Food types: No quantitative assessment; no recorded data.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (July to Sept.).
Bioregions South Africa: Red sands of Namaqualand Spinescent Grass-
land (SKs 12) (Namaqualand Sandveld Bioregion, Succulent Karoo
Biome).
Assessment rationale: EOO = 52.5 km2; currently known from only two
localities with observations that suggest associations restricted to red
sands of SKs 12 that cover a total of 521 km2 in four natural patches;
however, virtually unknown ecologically; assessed as Data Deficient
(DD), but, on the basis of flightlessness and extremely small known
range, it would qualify for an IUCN threat category of at least Vulnera-
ble (VU), although little of SKs 12 has been transformed (4%).
Conservation measures: To assess conservation status, a quantitative sur-
vey of SKs 12 and adjacent vegetation units is required to determine
the full EOO, AOO and precise ecological associations; not currently 2 mm

known from any protected area.

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Macroderes bias
(Olivier, 1789)

= Macroderes pilula Sharp, 1880

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus bias: ‘Cap de Bonne-Espérance’ [Cape

of Good Hope, South Africa], neotype: ‘Graham’s Town’ [Grahams­
town, Eastern Cape, South Africa]; M. pilula: ‘Grahamstown, Cap de
Bonne-Espérance’ [Grahamstown, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Dry, grassy to woody vegetation units in E bimodal rainfall
region and edge of summer rainfall region, Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 839 ± 458, 147–1 522 m;
annual rainfall: 432 ± 77, 361–595 mm; annual temperature (max. +
min. /2): 15.5 ± 2.0, 11.1–18.0°C (N=9).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam in scrub; all but one of the known localities coincide with
natural shrubland vegetation units.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring, summer and autumn in the bimodal spring/autumn rainfall
region (Sept. to May).
Bioregions South Africa: Vegetation units: In E Fynbos Biome: Shale
Renosterveld (FRs 16), Sandstone Fynbos (FFs 28); in Albany Thicket
Biome (AT 4, AT 6, AT 11); in Savanna Biome: Sub-Escarpment Sa-
vanna (SVs 7); in S Grassland Biome: wooded units of Sub-Escarpment
Grassland (Gs 16, Gs 17), Dry Highveld Grassland (Gh 1).
Assessment rationale: EOO = 40 815 km2; range across various grassy
to woody vegetation units comparatively large; not known if AOO is
influenced by soil and vegetation associations or habitat transforma-
tion, which is 2–20% in cited vegetation units; species little known, so
assessed as Data Deficient (DD), but on basis of wide range, potentially
Least Concern (LC) at present.
Conservation measures: To assess conservation status, quantitative data
on ecological associations are required, particularly a comparison of
population density data from grassy to more densely wooded vegeta-
tion units; protected in Addo Elephant National Park.
2 mm

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SURICATA 6 (2020) 235

Macroderes cornutus
Frolov & Scholtz, 2004

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Namaqualand, Sand Kop’ [farm on W coast, South Africa].

Taxonomy: Accepted species.
Distribution: Arid deep W coastal sands, Namaqualand, Northern Cape,
South Africa.
Locality data (mean ± SD, range): Altitude: 73 ± 54, 0–124 m; annual
rainfall: 115 ± 33, 79–159 mm; annual temperature (max. + min. /2):
17.4 ± 1.0, 16.0–18.2°C (N=4).
Habitat: No quantitative assessment; all three known localities on deep
sands supporting shrubland; two of the records from dunes.
Food types: No quantitative assessment; one record from a bush rat nest.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug., Sept.).
Bioregions South Africa: Namaqualand Coastal Duneveld (SKs 8), Nam-
aqualand Strandveld (SKs 7) (Namaqualand Sandveld, Bioregion, Suc-
culent Karoo Biome); also Namaqualand Sand Fynbos (FFd 1) (Sand
Fynbos, Fynbos Biome).
Assessment rationale: EOO = 2 605 km2; EOO presumably underesti-
mated as SKs 7 and SKs 8 cover a combined area of ± 3 950 km2 along
the Namaqualand coast, some of which is protected in Namaqua Na-
tional Park; FFd 1 covers ± 770 km2; transformation 2–10%, especially
by coastal mining and some inland cultivation, with extensive grazing
pressure in SKs 7 and FFd 1; qualifies for an IUCN threat category of
Vulnerable (VU) on the basis of small range and few known localities,
but currently poorly known, so assessed as Data Deficient (DD).
Conservation measures: A quantitative survey of SKs 7, SKs 8 and FFd
1 is required to adequately assess conservation status; this needs to
include assessment of ecological associations and effects of both hab-
2 mm
itat transformation and land usage; not currently recorded from any
protected area.

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Macroderes endroedyi
Frolov & Scholtz, 2004

No synonyms
Global: NT (see IUCN Red List – NT)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘SW Cape, Nortier Farm’ [W Coast, Western Cape, South
Africa].
Taxonomy: Accepted species.
Distribution: Dry W coast deep sands, Western Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 162 ± 102, 37–357 m; an-
nual rainfall: 204 ± 28, 170–260 mm; annual temperature (max. +
min. /2): 17.7 ± 0.5, 16.9–18.3°C (N=9).
Habitat: No quantitative assessment; no available data.
Food types: No quantitative assessment; sampled in ground traps baited
with human dung and carrion.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug., Sept.).
Bioregions South Africa: Centred on deep sands of Lamberts Bay Strand-
veld (FS 1) (Western Strandveld, Fynbos Biome); also S tip Namaqua-
land Strandveld (SKs 7) (Namaqualand Sandveld Bioregion, Succulent
Karoo Biome) plus NE edges of Graafwater Sandstone Fynbos (FFs 2),
Cederberg Sandstone Fynbos (FFs 4) and Piketberg Sandstone Fynbos
(FFs 6) (Sandstone Fynbos, Fynbos Biome).
Assessment rationale: EOO = 2 775 km2; small range centred on FS 1,
a deep sand and shrubland vegetation unit that has been 25% trans-
formed by cultivation with additional pressures from long-term graz-
ing; other records only at edges of some adjoining units; considering
flightless habits and land utilisation pressures assessed as Near Threat-
ened (NT) in line with the IUCN Red List although essentially Data
2 mm Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
responses to habitat transformation and grazing pressures; absence of
records from Leipoldtville Sand Fynbos (FFd 2: 55% transformed)
and presence at edges of adjoining Sandstone Fynbos vegetation units
should be investigated as a possible habitat transformation effect; no
current records known from protected areas.

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Macroderes fornicatus
Sharp, 1880

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: Not stated.

Taxonomy: Accepted species.
Distribution: Currently known only from Cape Peninsula, SW Western
Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 235 ± 332, 0–469 m; annual
rainfall: 454 ± 37, 428–480 mm; annual temperature (max. + min. /2):
15.2 ± 1.3, 14.3–16.2°C (N=2).
Habitat: No adequate quantitative assessment; on deep sands of Cape
Peninsula: sampled only from dense low growth of Restio-dominated
fynbos (burnt 2–3 yrs previously), not in open ericoid/proteoid fynbos
(burnt 9–10 yrs previously) or in dense, low growth, Kikuyu pasture
replacing natural shrubland (Farm Bonne Attente).
Food types: No quantitative assessment; sampled in low numbers to cattle
dung (11).
Temporal activity: Flightless; diel periodicity uncertain; sampled from
late winter to the end of spring in the winter rainy season (July to early
Nov.).
Bioregions South Africa: Accurately mapped only to Peninsula Sandstone
Fynbos (FFs 9) (Sandstone Fynbos, Fynbos Biome).
Assessment rationale: EOO = 535 km2 (possibly underestimated); AOO
possibly smaller as it is, currently, known only from FFs 9 covering
± 230 km2 on the Cape Peninsula where it was localised to dense re-
generating cover; although much is highly transformed (75%), most of
the remainder is under protection; flightlessness and very small range
in a highly exploited area would qualify for an IUCN threat category
approaching Endangered (EN), however, currently assessed as Data
2 mm
Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by a survey of Sandstone Fynbos in the vicinity of Cape
Town, particularly lower-altitude parts of Kogelberg Sandstone Fynbos
(FFs 11 – covering 888 km2); further quantitative data on ecological
associations are also required, particularly soil data and more detailed
information on responses to vegetation cover; protected in Cape of
Good Hope Nature Reserve, Cape Peninsula.

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Macroderes foveatus
Frolov & Scholtz, 2004

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘3 km E of Velddrif ’ [Western Cape, South Africa].

Taxonomy: Accepted species, but morphological differences at N localities
(black squares), which fall within the range of the related M. cornutus
Frolov & Scholtz, 2004; may equally be a further undescribed species.
Distribution: Restricted range in a shrubland vegetation unit on dry,
deep, SW coastal sands, South Africa.
Locality data (mean ± SD, range): Altitude: 20 ± 25, 2–38 m; annual
rainfall: 255 ± 16, 244–266 mm; annual temperature (max. + min. /2):
16.6 ± 0.1, 16.5–16.7°C (N=2 – red squares only).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; sampled to human dung bait.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug.).
Bioregions South Africa: Hopefield Sand Fynbos (FFd 3) (Sand Fynbos,
Fynbos Biome) (red squares only).
Assessment rationale: EOO = 105 km2 (red squares only); type series
recorded from two localities in a single vegetation unit that has been
40% transformed for arable farming and pasturage; on the basis of
flightlessness and occurrence at few locations within an extremely
small, threatened range, it would qualify for at least Vunerable (VU),
even Endangered (EN), but currently assessed as Data Deficient (DD)
due to absence of quantitative support.
Conservation measures: To assess conservation status taxonomic ques-
2 mm tions need to be resolved and a quantitative survey is required to de-
termine EOO, AOO, ecological associations and whether clearance of
natural shrubland influences population density or AOO; not currently
known from any protected area.

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Macroderes greeni
(Kirby, 1818)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Onthophagus greeni: ‘Promontorium Bonae Spei’ [Cape

of Good Hope, South Africa].
Taxonomy: Accepted species.
Distribution: Dry coastal sands to N and SE of Cape Town, SW Western
Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 47 ± 41, 0–130 m; annual
rainfall: 324 ± 50, 269–422 mm; annual temperature (max. + min. /2):
16.1 ± 0.5, 15.5–16.6°C (N=7).
Habitat: No adequate quantitative assessment; sampled on deep sand in
open shrubland on farms Modderrivier (6), Pampoenvlei (2); also in
shrubland at Geelbek (13), but not in adjacent sparse pasture cleared of
natural shrubland (0).
Food types: No quantitative assessment; sampled to cattle dung.
Temporal activity: Flightless; diel periodicity uncertain; active in the win-
ter rainy season; on SW coast: sampled primarily in autumn tailing off
through winter to early spring (May to Sept.).
Bioregions South Africa: Langebaan Dune Strandveld (FS 5), Overberg
Dune Strandveld (FS 7) (Western Strandveld); Atlantis Sand Fynbos
(FFd 4) (Sand Fynbos, Fynbos Biome).
Assessment rationale: EOO = 1 535 km2; presumably a deep sand
specialist in natural shrubland although further quantitative support
is required; occupies a small known range, part of which is highly
transformed through cultivation and urban development (W coast:
35–40%, FS 5, FFd 4); it would qualify for an IUCN threat category
2 mm
of at least Vulnerable (VU) on the basis of flightlessness and small range
in few locations, although currently assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by a quantitative survey to determine full EOO, AOO, eco-
logical associations and potential effects of habitat transformation; pro-
tected in Koeberg Nature Reserve and West Coast National Park where
the sparse grassland on Geelbek has now been permitted to regenerate
as natural shrubland.

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Macroderes minutus
Frolov & Scholtz, 2004

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘RSA, Western Cape Prov., 15 km SW of Lutzville’ [South

Africa] – [SW green squares on map].
Taxonomy: Currently an accepted species, but revision will probably re-
veal a complex of two or three species.
Distribution: Different members of species complex depicted in different
colours; mountains (Cederberg) [red squares] and coastal hardeveld or
sandveld [black or green squares], SW Western Cape, South Africa.
Locality data (mean ± SD, range): Black and green squares (lowlands):
altitude: 257 ± 136, 77–404 m; annual rainfall: 155 ± 9, 145–169 mm;
annual temperature (max. + min. /2): 18.4 ± 0.6, 17.6–19.2°C (N=6).
Red squares (Cederberg): altitude: 1 144 ± 702, 563–1 924 m; annual
rainfall: 256 ± 63, 198–322 mm; annual temperature (max. + min. /2):
14.1 ± 3.8, 9.9–17.3°C (N=3).
Habitat: No quantitative assessment; some from deep sands; otherwise no
precise soil or vegetation data.
Food types: No quantitative assessment; records from carrion, plus human
and reedbuck dung.
Temporal activity: All members of complex flightless; diel periodicity un-
certain; active during spring in the winter rainy season (July to Sept.).
Bioregions South Africa: Vegetation units: black squares: Namaqua-
land Hardeveld (SKn 1) (Succulent Karoo Biome); green squares:
Knersvlakte Quartz Vygieveld (SKk 3), Vanrhynsdorp Gannabosveld
(SKk 5) (Succulent Karoo Biome), also Namaqualand Riviere (Azi 1)
(Inland Azonal); red squares: Swartruggens Quartzite Fynbos (FFq 2);
Cederberg Sandstone Fynbos (FFs 4) (Fynbos Biome).
Assessment rationale: EOO for species complex = 5 045 km2, with
2–20% habitat transformation across the combined range; thus, EOO
2 mm could be < 2 000 km2 for each of three species and AOO even smaller;
coupled with flightless habits, each would qualify for an IUCN threat
category approaching Endangered (EN); however, considering the poor
state of taxonomic, distributional and ecological knowledge, currently
assessed as Data Deficient (DD).
Conservation measures: Conservation status may only be assessed once
taxonomic revision has resolved issues of species identity and surveys
have been conducted to determine full EOO, AOO, and ecological
associations; Cederberg localities protected in Cederberg Wilderness
Area; protection of taxa represented by green and black squares un-
certain.

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Macroderes mutilans
Kolbe, 1908

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Klein Namaqualand : Steinkopf ’ [Steinkopf, Namaqua­

land, South Africa].
Taxonomy: Accepted species.
Distribution: Arid upper escarpment, Namaqualand, South Africa.
Locality data (mean ± SD, range): Altitude: 871 ± 98, 739–961 m; an-
nual rainfall: 129 ± 27, 95–158 mm; annual temperature (max. + min.
/2): 17.0 ± 0.8, 15.9–17.6°C (N=5).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; no data available.
Temporal activity: Flightless; diel periodicity uncertain; active during late
summer/autumn and spring in the winter rainy season; (Mar., Aug.,
Sept.).
Bioregions South Africa: Namaqualand Klipkoppe Shrubland (SKn 1),
Namaqualand Blomveld (SKn 3) (Namaqualand Hardeveld Bioregion,
Succulent Karoo Biome).
Assessment rationale: EOO = 1 501 km2 (presumably underestimated);
centred on a large upland area of arid scrub and shrubland on finer-
grained soils; although SKn 1 is little transformed, SKn 3 is 6% trans-
formed by cultivation and highly overgrazed; although the small known
range and low number of localities would qualify for an IUCN threat
category, currently assessed as Data Deficient (DD) pending survey of
SKn 3 and the steep rocky hillsides of SKn1, which cover an area of
± 9 900 km2, but also include the range of M. amplior Frolov & Scholtz
2004 on the lower escarpment to the W.
Conservation measures: Assessment of conservation status requires a 2 mm
quantitative survey to determine the full EOO, AOO, ecological as-
sociations, and particularly if overgrazing in SKn 3 is detrimental to
population density or AOO; not currently recorded from any protected
area.

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Macroderes namakwanus
Frolov & Scholtz, 2004

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Hoekbaai’ [S Namaqualand, Western Cape, South Africa].

Taxonomy: Accepted species.
Distribution: Arid, W coastal deep sands on Western Cape / Northern
Cape border, South Africa; identity of single female paratype from out-
lier locality in Richtersveld (black square) requires validation.
Locality data (mean ± SD, range): Altitude: 148 ± 205, 0–498 m; annual
rainfall: 126 ± 31, 70–144 mm; annual temperature (max. + min. /2):
17.9 ± 0.4, 17.3–18.3°C (N=5).
Habitat: No quantitative assessment; two paratype localities on white
sands found close to the coast, one from yellow sands found more in-
land.
Food types: No quantitative assessment; sampled to human dung bait.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug., Sept.).
Bioregions South Africa: Central Namaqualand Strandveld (SKs 7) (Na-
maqualand Sandveld Bioregion, Succulent Karoo Biome).
Assessment rationale: EOO = 53 km2 (range for red squares only); vali-
dated range extremely small lying between coastal ranges of M. cornutus
Frolov & Scholtz, 2004, to the N and close relative, M. endroedyi Frolov
& Scholtz, 2004, to the S; area around type locality little transformed,
but on the basis of extremely small EOO and records from few loca-
tions, would qualify for an IUCN threat category of at least Vulnerable
(VU); however, in view of severely limited ecological and distributional
data, currently assessed as Data Deficient (DD).
2 mm
Conservation measures: Assessment of conservation status requires that
questions on the identity of the Richtersveld paratype be resolved; a
quantitative survey of the vegetation units around the type locality is
required to establish the EOO, AOO and ecological associations; all
validated localities are in unprotected areas.

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Macroderes nitidus
Harold, 1877b

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cap bon. spei’ [Cape of Good Hope, South Africa].
Taxonomy: Accepted species, but additional material from ‘Cedarberg
Range, east track’ cited in the revision of Frolov and Scholtz (2004) is
considered to belong to an undescribed species.
Distribution: Until recently known only from material bearing type labels
comprising 12 individuals, six from an unstated spot locality in the
Cape of Good Hope and six from ‘Orlog Rivier’ [Oorlogskloof River]
that flows W from Calvinia; all collected by ‘Meyer’, a medical doc-
tor who was based in that town; recently rediscovered after 156 years
± 40 km W of Calvinia.
Locality data (mean ± SD, range): Altitude: 791 ± 86, 730–851 m; an-
nual rainfall: 169 ± 3, 167–171 mm; annual temperature (max. + min.
/2): 17.0 ± 0.3, 16.8–17.3°C (N=2).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; no data available.
Temporal activity: Flightless; diel periodicity uncertain; active during
spring in the winter rainy season (Aug.).
Bioregions South Africa: Vegetation unit: S Hantam Karoo (SKt 2)
(Trans-Escarpment Succulent Karoo Bioregion, Succulent Karoo
Biome).
Assessment rationale: EOO = 80 km2 (underestimated); difficult to assess
conservation status as there is only a single precise spot locality data
and no ecological data; therefore, assessed as Data Deficient (DD);
however, threats may be limited as SKt 2 covers a relatively large area
(7 464 km2) and is little transformed. 2 mm
Conservation measures: To assess conservation status, a quantitative
survey is required to determine the EOO, AOO and ecological associa-
tions; this survey should be focused on the area to the west of Calvinia
and along the Oorlogskloof River in and around the SKt 2 vegetation
unit; not currently known from any protected area.

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Macroderes politulus
Preudhomme de Borre, 1880

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Le patrie présumable est donc la Cafrerie’ [presumably

Caffraria, inexact locality in South Africa].
Taxonomy: Accepted species.
Distribution: Dry climate in Boland region of Western Cape, South Af-
rica.
Locality data (mean ± SD, range): Altitude: 320 ± 245, 62–871 m; an-
nual rainfall: 356 ± 76, 265–466 mm; annual temperature (max. +
min. /2): 16.5 ± 0.7, 15.5–17.8°C (N=8).
Habitat: No quantitative assessment; one individual sampled over an en-
tire year on sandy loam in grassland adjacent to a patch of degraded
shrubland at farm Oranjefontein near Darling, Western Cape.
Food types: No quantitative assessment; one individual sampled to cattle
dung.
Temporal activity: Flightless; diel periodicity uncertain; active during
autumn, early winter and spring in the winter rainy season; (May to
Oct.).
Bioregions South Africa: Various vegetation units centred around the
Boland in the Fynbos Biome: Swartland Granite Renosterveld (Frg 2),
Swartland Shale Renosterveld (Frs 9), Breede Alluvium Renosterveld
(FRa 1), Lourensford Alluvium Fynbos (FFa 4), Winterhoek Sandstone
Fynbos (FFs 5), Piketberg Sandstone Fynbos (FFs 6).
Assessment rationale: EOO = 7 940 km2; small range centred in a region
dominated by finer-grained soils and shrubland, much of which is
highly transformed by agriculture or urban development (Frs 9: 90%;
Frg 2: 80%; FFa 4: 90%; FRa 1: 57%); currently assessed as Data Defi-
cient (DD) due to absence of quantitative data; although would deserve
an IUCN threat category of at least Vulnerable (VU) on the basis of
flightlessness and small, presumably fragmented range at few known
locations; presence in mountain and lowland refuges is supported by
2 mm an older record from the little-transformed FFs 5 (5%) and also several
recent records (2006) from FFs 6 (17% transformed) and a patch of
Renosterveld adjoining the sports field at Stellenbosch University.
Conservation measures: To assess conservation status, a quantitative
survey is required to determine the full EOO, AOO and ecological
associations; as removal of natural shrubland is likely detrimental to
population density and AOO, a survey is required to determine the
number of refuge locations occupied at the present time; not currently
known from any protected area.

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Macroderes undulatus
Preudhomme de Borre, 1880

= Macroderes westwoodi Preudhomme de Borre, 1880

Global: NT (See IUCN Red List – NT)
Endemic: RSA

J A S O N D J F M A M J

Type localities: M. undulatus: ‘Cap de Bonne-Espérance’; M. westwoodi:

‘Cap de Bonne-Espérance’ [both Cape of Good Hope, South Africa].
Taxonomy: Accepted species.
Distribution: Mainly dry Renosterveld localities on low-lying S coastal
hills, Western Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 277 ± 179, 161–541 m;
annual rainfall: 388 ± 21, 366–412 mm; annual temperature (max. +
min. /2): 16.7 ± 0.6, 15.8–17.4°C (N=4).
Habitat: No quantitative assessment; limited DBRU collection records
from a fallow crop field adjoining a patch of natural shrubland after
July rainfall and from sandy clay loam in pasture resulting from clear-
ance of natural shrubland.
Food types: No quantitative assessment; limited DBRU collection records
from cattle dung (2).
Temporal activity: Flightless; diel periodicity uncertain; active during
autumn and winter in the winter rainy season (May, July).
Bioregions South Africa: Various vegetation units in Fynbos Biome along
S coastal hills: Western Rûens Shale Renosterveld (FRs 11), Central
Rûens Shale Renosterveld (FRs 12), Eastern Rûens Shale Renosterveld
(FRs 13); also Swellendam Silcrete Fynbos (FFc 1).
Assessment rationale: EOO = 2 300 km2 (presumably underestimated);
AOO perhaps restricted to finer-grained soils that once supported
Renosterveld shrubland; because of the high level of transformation
by cultivation (80–87%: FRs 11, FRs 12, FRs 13) and low numbers of
records, assessed as Near Threatened (NT) to align with the IUCN Red
List, although essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
2 mm
improved by a quantitative survey to establish the full EOO, AOO,
ecological associations, and how population density or AOO might
be influenced by the extreme habitat transformation, particularly in
Rûens Renosterveld (± 5 870 km2); this is required to justify the NT
assessment; not currently known from any protected area although a
few reserves occur within its range.

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Genus Metacatharsius Montreuil, 1998

Type species and designation: Scarabaeus ferrugineus Olivier, 1789, by original designation.
Synonyms: None.
Last review: All species reviewed by Balthasar (1965b) with a few other species described subse-
quently (Balthasar 1968, 1970; Frey 1975a; Walter 1976).

Metacatharsius Paulian, 1939, was originally described as a subgenus to accommodate 16 species previously described un-
der Catharsius. However, under international rules of nomenclature postdating 1930, the name was considered invalid as
no type species was designated. A type species was only proposed by Montreuil (1998) together with a matrix of characters
to separate the taxon from others at generic level. According to Branco (2011) this action satisfied international nomen-
clatural rules, so that Montreuil became the author of this taxon. However, this change from Metacatharsius Paulian, 1939,
may be open to dispute.

There is no phylogeny and no division into species groups even though the genus comprises 64 species. Of these, 63 are
restricted to the Afrotropical region and one is shared with the Palaearctic region in a Saharo–Sindian pattern of distribu-
tion. Although there are 16 currently valid names for South Africa, Botswana and Namibia, only 11 species accounts are
presented. Arrangements of more than one species per account reflect suspected unpublished synonymies, indicating that
taxonomic revision is required. No species account was prepared for one species, Metacatharsius transvaalensis Balthasar,
1968, as its identity could not be determined with confidence in comparison with a few other unnamed possibly unde-
scribed species.

Metacatharsius species are much smaller-bodied than Catharsius species and, as such, would require less dung for breeding
purposes. Tunnels are constructed from under droppings, but nesting behaviour has not been investigated. All species are
macropterous showing flight activity in darkness. They may show a bias to carrion plus omnivore dung or a bias only to
dung.

As association with sandy soils dominates the distributions of Metacatharsius, all but one southern African species show
occurrence on deep Kalahari sands. The exception is centred on the arid southwest late summer rainfall region. Three other
arid-centred species show distributions restricted to the southwest Kalahari (1), also radiating to the NW along the edge
of the Namib Desert (1), or, with an additional, apparently disjunct occurrence, on the southwest coastal sands (1). Most
other southern African species are centred in moister regions with distributions extending across most of the Kalahari to
E outliers and the NE coastal sands of KwaZulu-Natal (5) with two known only from the NE Kalahari. Some of these
patterns are dependent on the unpublished taxonomic interpretation used in the species accounts.

Because most distributions occur wholly or partly in relatively untransformed regions, taxa cited in eight of the species
accounts are assessed as Least Concern (LC), whereas three are assessed as Data Deficient (DD) due to limited information
or unresolved taxonomic problems. Additional taxonomic scrutiny is also required on three species pages where taxa are
assessed as LC.

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Metacatharsius anderseni
(Waterhouse, 1891)

= Metacatharsius troglodytes Boheman, 1857, sensu Pereira & d’Andretta,

1955
= Catharsius (Metacatharsius) exiguiformis Ferreira, 1964b (pars, 1 paratype)
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Catharsius anderseni: ‘Lake Nyassa’ [=Lake Malawi,

but would be an error; see notes on the Lake Ngami type locality of
Scarabaeus (Scarabaeolus) anderseni Waterhouse, 1890]; M. troglodytes
sensu Pereira & d’Andretta: Not stated; C. (M.) exiguiformis pars: ‘Twee
Rivieren’ [Northern Cape, South Africa].
Taxonomy: Accepted species, but the synonymy of M. troglodytes sensu
Pereira & d’Andretta, 1955, requires re-examination.
Distribution: Deep sands of the SW Kalahari: N South Africa, SW Bot­
swana, E Namibia; recorded distribution in Mozambique is an error.
Locality data (mean ± SD, range): Altitude: 1 077 ± 113, 870–1 461 m;
annual rainfall: 266 ± 50, 179–380 mm; annual temperature (max. +
min. /2): 19.6 ± 0.6, 18.2–21.2°C (N=52).
Habitat: No adequate quantitative data; in Northern Cape: restricted to
SW Kalahari (0.4/trap at 40 sampling sites), Upper Karoo (0), Bush-
manland Karoo (0); few DBRU collection records: deep sand (2), grass-
land (1), open woodland (1).
Food types: On deep sands in SW Botswana (cited as Metacatharsius sp.
A): only attracted to dung with distinct bias in association, sheep (144),
cattle (8), elephant (32), pig (14), carrion (0); in Northern Cape: sam-
pled to composite cattle/sheep dung baits.
Temporal activity: Flight activity in darkness during the late summer
rainy season (Jan. to Mar.).
Ecoregions Namibia, Botswana: Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Kalahari Duneveld (SVkd), E Eastern Kalahari
Bushveld (SVk) (Savanna Biome). 2 mm

Assessment rationale: EOO = 255 955 km2; widespread on deep sands in

a little-transformed area of grassland and open shrubland that is used
primarily for the grazing of domestic livestock, particularly sheep, to
whose dung this species is strongly attracted; assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations; protect-
ed in Kgalagadi Transfrontier Park (South Africa, Botswana), Central
Kalahari Game Reserve (Botswana).

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Metacatharsius dentinum
(Ferreira, 1964b)

Metacatharsius zuluanus
(Balthasar, 1965b)

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: As Catharsius (Metacatharsius) dentinum: ‘Sudoeste Afri-

cano: Matamata, Kalahari Gemsbok Park’ [Mata Mata, South Africa];
As Catharsius (Metacatharsius) zuluanus: ‘Südafrika, Zululand (Mapu-
ta)’ [Manguzi, South Africa].
Taxonomy: Both currently accepted species, but M. zuluanus is possibly a
junior synonym of M. dentinum; type series of both species need to be
examined as possible synonyms of M. pumilionis (Wallengren, 1881)
described from Christiana, NW Province, South Africa.
Distribution: Dry to moist savanna on deep sands of Kalahari, E Kalahari
outliers, pre-Namib, E coastal plain: South Africa, Botwana, Namibia,
S Mozambique, S Zambia; also reported from Angola.
Locality data (mean ± SD, range): M. dentinum: altitude: 1 034 ± 170,
388–1 809 m; annual rainfall: 294 ± 147, 56–710 mm; annual tem-
perature (max. + min. /2): 19.5 ± 1.7, 15.5–23.4°C (N=114). M. zu-
luanus: altitude: 132 ± 185, 0–475 m; annual rainfall: 708 ± 166,
453–1 003 mm; annual temperature (max. + min. /2): 22.5 ± 0.6,
21.7–23.7°C (N=19).
Habitat: No quantitative assessment; limited DBRU collection records
entirely on deep sand (5) in shrubland (5).
Food types: In Botswana: strong bias to carrion (chicken livers: 864), also
attracted to dung, but much less abundant (pig: 151; elephant: 96; cat-
2 mm
tle: 88; sheep: 93); limited DBRU collection records on dead owl (2),
rotten goat meat (1), dead millipede (1), cattle dung (1).
Temporal activity: Flight activity in darkness primarily during the sum-
M. dentinum
mer rainy season (Sept. to May).
Ecoregions Namibia, Botswana, Mozambique: Primarily Namibian
Savanna Woodlands (AT1316), Kalahari Xeric Savanna (AT1309),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea
Woodlands (AT0726), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: N Bushmanland (NKb), N Upper Karoo (NKu)
(Nama Karoo Biome); Kalahari Duneveld (SVkd), Eastern Kalahari
Bushveld (SVk), sand outliers in Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome); Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 111 720 km2; all populations apparently
deep sand specialists although quantitative support is required; popu-
lations currently named M. dentinum are widespread, largely in areas
subject to low habitat transformation; they also show a strong bias to
carrion association; despite apparent specialist attributes, both current-
ly valid taxa deserve Least Concern (LC) status; however, they are as-
sessed as Data Deficient (DD) owing to taxonomic uncertainty as they
may comprise one or two species.
Conservation measures: Taxonomic issues need to be resolved before
conservation status may be adequately assessed; quantitative data on
ecological associations are also required, particularly the food associa-
2 mm tions of populations currently named M. zuluanus; protected in Kga­
lagadi Transfrontier Park (South Africa/Botswana), iSimangaliso Wet-
land Park (World Heritage Site) (South Africa).
M. zuluanus

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Metacatharsius exiguiformis
(Ferreira, 1964b)

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Catharsius (Metacatharsius) exiguiformis: ‘Sudoeste Af-

ricano: Twee Rivieren’ [Twee Rivieren, Northern Cape, South Africa].
Taxonomy: Accepted species, but SW Kalahari populations a little more
shiny, usually with a more indented, less sinuous anterior clypeal mar-
gin.
Distribution: Centred on the SW Kalahari (red squares) and pre-Namib
deep sands (black squares), arid late summer rainfall region: South Af-
rica, Botswana, Namibia.
Locality data (mean ± SD, range): SW Kalahari: altitude: 1 064 ±
125, 802–1 461 m; annual rainfall: 263 ± 67, 142–453 mm; annu-
al temperature (max. + min. /2): 19.7 ± 0.9, 17.9–21.4°C (N=146).
Namib: altitude: 888 ± 392, 283–1 809 m; annual rainfall: 146 ±
114, 35–381 mm; annual temperature (max. + min. /2): 16.9 ± 1.0,
15.8–18.3°C (N=22).
Habitat: No quantitative assessment; DBRU collection records almost
exclusively on deep sand (44), sandy loam (1) in grassland (11), scrub/
shrubland (24), open woodland (8).
Food types: In Botswana: equally abundant on dung of pig (1 367), ele-
phant (1 341) and sheep (1 436); much less abundant on cattle dung
(390), uncommon on carrion (chicken livers: 29); however, DBRU
collection records primarily from dung of cattle (34), also donkey (2),
horse (1), wildebeest (1).
Temporal activity: Flight activity in darkness primarily during the late
summer rainy season (Dec. to May). 2 mm

Ecoregions Namibia, Botswana: Kalahari: SE Kalahari Xeric Savanna

(AT1309); Namib: primarily straddling edges of Namibian Savanna
Woodlands (AT1316), Namib Desert (AT1315), Kaokoveld Desert
(AT1310).
Bioregions South Africa: Kalahari Duneveld (SVkd), Eastern Kalahari
Bushveld (SVk) (Savanna Biome); N extremities of Bushmanland
(NKb) and Upper Karoo (Nama Karoo Biome).
Assessment rationale: EOO = 471 130 km2; AOO would be smaller
due to specialist association with deep sands, which are, nevertheless,
extensive across its range; common in farmland of SW Kalahari and
pre-Namib plains, which are used primarily for the grazing of domes-
tic livestock, especially sheep; little habitat transformation; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations; protected in Kga-
lagadi Transfrontier Park (South Africa/Botswana), Namib-Naukluft
National Park (Namibia).

2 mm

SW Kalahari

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Metacatharsius exiguus
(Boheman, 1860)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Copris exigua: ‘prope lacum N’Gami’ [near Lake Ngami,
N Botswana].
Taxonomy: Accepted species, but slight morphological variation between
geographical centres; Kalahari and E outliers slightly indented clypeal
margin; Zimbabwe and Angola less indented margin; Maputaland less
indented more sinuous margin.
Distribution: Disjunct pattern in three dry to moist savanna centres of
southern Africa related to soil type specialisation: South Africa, Botswa-
na, Namibia, Angola, Zimbabwe; also reported from Mozambique.
Locality data (mean ± SD, range): Altitude: 827 ± 517, 0–1 503 m;
annual rainfall: 557 ± 204, 278–971 mm; annual temperature (max. +
min. /2): 20.7 ± 1.6, 17.6–23.1°C (N=40).
Habitat: No quantitative assessment; DBRU collection records primarily
on deep sand (11), sandy loam (2) in grassland/pasture (6), shrubland
(7), open woodland (3).
Food types: No quantitative assessment; DBRU collection records pri-
marily from dung of cattle (15), also elephant (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Oct. to May).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Kalaha-
ri Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian Baikiaea Woodlands (AT0726), Southern
Miombo Woodlands (AT0719), Maputaland Coastal Forest Mosaic
(AT0119).
Bioregions South Africa: Deep sand patches in Savanna Biome: East-
ern Kalahari Bushveld (SVk), Central Bushveld (SVcb), straddling SE
border of Lowveld (SVl) into Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 680 625 km2; AOO disjunct and smaller
according to occurrence of deep sands, which are widespread in the SW
Kalahari, Maputaland and parts of Zimbabwe; may also be restricted by
occurrence of more open vegetation, but quantitative support required;
habitat transformation low over SW part of range; assessed as Least
5 mm Concern (LC) on basis of large EOO.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Tswalu Kalahari Reserve, Tembe Elephant Park (South Africa).

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Metacatharsius ferreirae
(Balthasar, 1965b)

= Catharsius (Metacatharsius) dubius Ferreira, 1962a

= Catharsius (Metacatharsius) kochi Ferreira, 1964b
Global: DD

J A S O N D J F M A M J

Type localities: As Catharsius (Metacatharsius) ferreirae: nom nov. for

C. (M). dubius [pre-occupied name] ‘Camissombo, Calonda’ [NE
Angola]; C. (M.) kochi: ‘Mangetti, entre Tsumkwe e Omassico’ [NE
Namibia].
Taxonomy: Accepted species; M. kochi is a synonym of M. dubius; as
Metacatharsius was formerly considered a subgenus of Catharsius, C.
(M.) dubius was a pre-occupied name (Catharsius dubius Péringuey,
1901); thus, it was provided with the nom nov. of C. (M.) ferreirae;
types of Metacatharsius rugosipennis Frey, 1975, from ‘Betchuanaland:
Tsano’ [?Tsane, Botswana] need to be compared as a likely junior syn-
onym.
Distribution: Dry to moist savanna of N mega-Kalahari deep sands:
South Africa, Zimbabwe, Botswana, Namibia, Angola.
Locality data (mean ± SD, range): Altitude: 951 ± 246, 499–1 196 m;
annual rainfall: 459 ± 106, 341–608 mm; annual temperature (max. +
min. /2): 22.0 ± 1.5, 20.0–24.3°C (N=8).
Habitat: No quantitative assessment; one DBRU collection record on
deep sand in shrubland.
Food types: No quantitative assessment; one DBRU collection record on
carrion (rotten goat meat).
Temporal activity: Flight activity in darkness during the summer rainy
season (Jan., Feb.).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna
(AT1309), W Zambezian and Mopane Woodlands (AT0725), Kalaha-
ri Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea Wood-
lands (AT0726). 2 mm

Bioregions South Africa: Mopane (SVmp) (Savanna Biome).

Assessment rationale: EOO = 923 845 km2; poorly known, but possibly
a N mega-Kalahari deep sand and carrion specialist although quantita-
tive data are required in support; currently assessed as Data Deficient
(DD) although it is widely distributed across an area in which habitat
transformation is comparatively low, overall.
Conservation measures: A survey is required to determine the full EOO
and AOO; quantitative data on ecological associations are also required
before it is possible to adequately assess conservation status; protected
in Hwange National Park (Zimbabwe).

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Metacatharsius freyi
(Ferreira, 1964b)

No synonyms*
Global: DD
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: As Catharsius (Metacatharsius) freyi: ‘Bechuanalândia:

Ngami’ [Lake Ngami, N Botswana].
Taxonomy: Accepted species, but validation required for species treated
here as M. freyi, which was described from a single holotype male;
*this holotype needs to be compared for synonymy with the type(s) of
M. pusio (Kolbe, 1908) described as Catharsius pusio from ‘Kalahari:
Sekgoma-Khakea......Khakea-Kang’ [Sekumo to Kang, central Kalaha-
ri, Botswana].
Distribution: Dry to moist savanna on deep Kalahari sands and Kalahari
outliers: N South Africa, Botswana; M. pusio cited from Botswana,
Namibia.
Locality data (mean ± SD, range): Altitude: 994 ± 136, 817–1 222 m;
annual rainfall: 512 ± 127, 354–684 mm; annual temperature (max. +
min. /2): 20.8 ± 1.7, 17.9–22.7°C (N=10).
Habitat: No quantitative assessment; distribution coincides with continu-
ous deep sands of the Botswana Kalahari and sand patches in E outliers.
Food types: No quantitative assessment; no data available.
Temporal activity: Like other Metacatharsius species, flight activity pre-
sumably in darkness during the summer rainy season (Oct. to Apr.).
Ecoregions Botswana: Primarily Kalahari Acacia-Baikiaea Woodlands
(AT0709).
Bioregions South Africa: Sand patches in Central Bushveld (SVcb) (Sa-
vanna Biome).
Assessment rationale: EOO = 257 510 km2 (presumably underestimat-
2 mm
ed); widespread in N Botswana and N South Africa; probably a deep
sand specialist and possibly primarily in wooded areas, but quantitative
support required; possibility of threats due to clearance of woodland
need to be investigated; currently assessed as Data Deficient (DD) due
to poor state of knowledge both in terms of species identity and eco-
logical associations.
Conservation measures: Taxonomic issues need to be resolved and quan-
titative data on ecological associations are required before an assessment
may be made of conservation status; protected in Nylsvlei Nature Re-
serve (South Africa).

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Metacatharsius latifrons
(Harold, 1868)

No synonyms
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: As Copris latifrons: ‘Cap bon. spei.’ [Cape of Good Hope,
South Africa].
Taxonomy: Accepted species.
Distribution: Arid deep sands: SW Kalahari (Namibia, Botswana, South
Africa) and W coast, South Africa; uncertain if the disjunct pattern is
real or a collection artefact; report from Mozambique is an error.
Locality data (mean ± SD, range): Altitude: 795 ± 517, 0–1 478 m;
annual rainfall: 279 ± 112, 63–661 mm; annual temperature (max. +
min. /2): 18.4 ± 1.3, 14.5–20.8°C (N=91).
Habitat: No quantitative assessment; DBRU collection records indicate
a deep sand specialist: sand (39), sandy loam (3) in open vegetation:
open woodland (4), scrub/shrubland (17), grassland (16); also fallow
crop fields (2); in Northern Cape: entirely restricted to SW Kalahari
(average of 0.08/sample at 23 study sites), none in Upper Karoo or
Bushmanland Karoo.
Food types: No adequate quantitative assessment; in Botswana: sampled
from pig (3), elephant (2) and sheep dung (2), not from cattle dung
or carrion; DBRU collection records: dung of cattle (36), horse (1),
donkey (1).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season in the Kalahari (Oct. to Apr.); in the W coast winter
rainfall region most records in spring (Sept.) but with activity continu-
ing into the dry season (Sept. to Feb.).
Ecoregions Botswana: Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk); Namaqualand Sandveld (SKs); Sand Fynbos
(FFd), Western Strandveld (FS) (Biomes: Savanna – SV, Succulent
Karoo – SK, Fynbos – F).
Assessment rationale: EOO = 322 620 km2; widespread, but mostly
recorded in low population density; however, much of range little
transformed and used primarily for grazing of domestic livestock; trans- 5 mm
formation extensive only in extreme SW (25–70%) and E of range
(13–42%) in South Africa; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected
in West Coast and Namaqua national parks (South Africa), Kgalaga-
di Transfrontier Park (South Africa, Botswana) and Central Kalahari
Game Reserve (Botswana).

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Metacatharsius marani
(Balthasar, 1940)

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Catharsius (Metacatharsius) mařani: ‘Süd-West Afrika,

Okanjande und Karibib’ [Namibia: Okanjande, Karibib].
Taxonomy: Accepted species.
Distribution: Centred on the arid late summer rainfall region: South Af-
rica, SW extremity of Botswana, Namibia.
Locality data (mean ± SD, range): Altitude: 1 083 ± 199, 424–1 782 m;
annual rainfall: 223 ± 62, 22–407 mm; annual temperature (max. +
min. /2): 18.5 ± 1.8, 14.1–22.2°C (N=239).
Habitat: No adequate quantitative assessment; in Northern Cape: strong
bias to arid Nama Karoo (Upper Karoo: 14.1; Bushmanland: 11.9)
compared to dry SW Kalahari (2.8); DBRU collection records suggest
a bias to sandy soils: sandy clay loam (3), sandy loam (5), sand (21)
in open vegetation: open woodland (4), open scrub/shrubland (18),
grassland (9).
Food types: No adequate quantitative assessment; at range edge in SW
Botswana: attracted only to sheep dung (6), not dung of pig, elephant,
cattle or to carrion (0); DBRU collection records from dung of cattle
(18), wildebeest (1), horse (1), donkey (1), zebra (2).
Temporal activity: Flight activity in darkness during the late summer
rainy season (Dec. to April), attracted to light.
Ecoregions Namibia: Centred on SW Kalahari Xeric Savanna (AT1309),
Namibian Savanna Woodlands (AT1316); margins of two other ecore-
gions.
Bioregions South Africa: Centred on Upper Karoo (NKu), Bushmanland
(NKb) (Nama Karoo Biome); marginal in Kalahari Duneveld (SVkd),
E Eastern Kalahari Bushveld (SVk) (Savanna Biome).
Assessment rationale: EOO = 312 390 km2; centred in a region that is lit-
2 mm tle transformed and used primarily for the grazing of domestic livestock,
particularly sheep; widespread and often abundant in farm rangeland;
readily attracted to composite baits of cattle and sheep dung; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Kgalagadi Transfrontier Park (South Africa).

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Metacatharsius opacus
(Waterhouse, 1891)
= Catharsius cephalotes Péringuey, 1892

Metacatharsius pseudoopacus
(Ferreira, 1965)
No synonyms
Global: LC
J A S O N D J F M A M J

Type localities: As Catharsius opacus: ‘Lake Ngami’ [N Botswana];

C. cephalotes: Not stated; holotype as Catharsius (Metacatharsius) pseu-
doopacus: ‘Província de Moçambique: Changalane’ [S Mozambique]. ♂
Taxonomy: Currently both accepted species, but M. pseudoopacus consid-
ered to be a SE coastal synonym of M. opacus.
Distribution: Dry to moist savanna across N southern Africa and up E
seaboard to E and NE Africa: South Africa, Eswatini, Botswana, Na-
mibia, Angola, Zimbabwe, Zambia, Mozambique, Kenya, Somalia;
also reported from Tanzania.
Locality data (mean ± SD, range): Altitude: 771 ± 432, 0–1 730 m; an-
nual rainfall: 560 ± 189, 171–1 254 mm; annual temperature (max. +
min. /2): 21.3 ± 1.7, 15.4–25.9°C (N=189).
Habitat: In uMkhuze Game Reserve: strong bias to deep sand (14.5), few
on sand over clay (0.4), absent from sandy clay loam (0), clay (0); partly
supported by DBRU collection records, bias to sand (35), but also san-
dy loam (13), sandy clay loam (2), primarily in wooded habitats, forest
(6), open woodland (21), scrub/shrubland (13), grassland (4).
Food types: On deep sand in Botswana: relative food type generalist with
slight bias to carrion (chicken livers: 313) and dung of pig (325), also
dung of elephant (128), cattle (219), sheep (209); similar at Phalabor-
wa: dung of pig (65), elephant (35), cattle (22); DBRU collection re-
cords on dung of human/cattle or buffalo mix (7), elephant (7), rhinoc- 2 mm
eros (1), donkey (2), buffalo (1), cattle (43).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); attracted to light; at Phalaborwa: most active
on warm evening soon after heavy rainfall (86), also cool evening after
light rainfall (29), few on hot, dry evening (7).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: N Na-
mibian Savanna Woodlands (AT1316), N Kalahari Xeric Savanna
(AT1309), Angolan Mopane Woodlands (AT0702), Kalahari Acacia-
Baikiaea Woodlands (AT0709), Zambezian Baikiaea Woodlands
(AT0726), Southern African Bushveld (AT0717), Zambezian and Mo-
pane Woodlands (AT0725), Southern Miombo Woodlands (AT0719),
Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld SVcb), Mopane (SVl),
Lowveld (SVl) (Savanna Biome), N Indian Ocean Coastal Belt Biome
(CB); scattered records in five other bioregions.
Assessment rationale: EOO = 3 129 685 km2; AOO probably smaller
due to bias to coarser-grained soils and lower-altitude, woody vege-
tation; clearance of woodland would presumably reduce population
density to very low levels; range in South Africa subject to variable de-
grees of habitat transformation (0–58%, SVl; 2–49%, SVcb; 0–20%,
SVmp); however, very widely distributed and protected in various re-
serves; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations; in South
Africa: protected in uMkhuze Game Reserve, Kruger National Park. ♀ 2 mm

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Metacatharsius pumilionis
(Wallengren, 1881)

Metacatharsius pumilioniformis
(Ferreira, 1964b)

No synonyms
Global: LC (for M. pumilioniformis – see IUCN Red List – LC)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Copris pumilionis: ‘Ad Christianam semel lecta’ [in-

terpreted as collected at Christiana, NW Province, South Africa]; as
Catharsius (Metacatharsius) pumilioniformis: ‘Sudoeste Africano: Mata-
mata’ [Mata Mata, Northern Cape, South Africa].
Taxonomy: Both accepted species, but validation required in comparison
with type(s) for species treated, here, as M. pumilionis, whose type(s)
also need to be compared with those for M. pumilioniformis as W and
E populations appear to be synonymous, although there are subtle dif-
ferences; also see taxonomic notes for M. dentinum Ferreira, 1964b.
Distribution: M. pumilionis (black squares): dry to moist E savanna:
South Africa, Zimbabwe; also cited from Botswana, Namibia; M. pum-
ilioniformis (red squares): arid W savanna: South Africa, Botswana, Na-
mibia; overlap at two squares in N Cape.
Locality data (mean ± SD, range): M. pumilionis: altitude: 949 ± 421,
60–1 395 m; annual rainfall: 523 ± 135, 328–853 mm; annual temperature
(max. + min. /2): 19.7 ± 1.9, 17.3–24.5°C (N=25). M. pumilioniformis: al-
titude: 1 050 ± 132, 486–1 291 m; annual rainfall: 273 ± 83, 89–555 mm;
annual temperature (max. + min. /2): 19.7 ± 1.1, 17.3–22.8°C (N=43).
Habitat: No adequate quantitative assessment; in a Northern Cape sur-
2 mm vey of the SW Kalahari and N Nama Karoo, M. pumilioniformis was
restricted to the SW Kalahari, primarily on deep sands.
Food types: No adequate quantitative assessment; in Botswana: sampled
in low numbers only to dung of cattle (5) and sheep (1), not to dung of
M. pumilionis
pig, elephant or to carrion (chicken livers); occasionally sampled abun-
dantly to sheep/cattle dung mixture in N Cape.
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: M. pumilioniformis: pri-
marily Kalahari Xeric Savanna (AT1309). M. pumilionis: Southern Af-
rican Bushveld (AT0717).
Bioregions South Africa: M. pumilioniformis: primarily Kalahari Dune­
veld (SVkd), Eastern Kalahari Bushveld (SVk). M. pumilionis: primari-
ly Eastern Kalahari Bushveld (SVk), Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome).
Assessment rationale: EOO = 688 900 km2 (combined); AOO possibly
smaller, restricted to deep sands, possibly in more open vegetation, but
quantitative support required; M. pumilioniformis may be locally abun-
dant in the N Cape (33.0 per sample) and may show a bias to ruminant
herbivore dung; difficult to assess threat levels given poor state of taxo-
nomic and ecological knowledge, but both assessed as Least Concern (LC)
on basis of wide EOO of both populations sets on areas of deep sands.
Conservation measures: Taxonomic issues need to be resolved and quan-
2 mm
titative data on ecological associations are required to improve the as-
sessment of conservation status; M. pumilioniformis protected in Kga-
lagadi Transfrontier Park (South Africa / Botswana), M. pumilionis in
Kruger National Park (South Africa).
M. pumilioniformis

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Metacatharsius troglodytes
(Boheman, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Catharsius troglodytes: ‘juxta fluvium Gariep’ [near Or-

ange River, South Africa].
Taxonomy: Accepted species.
Distribution: Widespread in southern African savanna; mostly on moister
deep sands of the Kalahari, its E outliers, and the SE coastal plain:
South Africa, Mozambique, Zimbabwe, Botswana, Namibia, S Angola.
Locality data (mean ± SD, range): Altitude: 934 ± 408, 0–1 874 m;
annual rainfall: 538 ± 148, 238–972 mm; annual temperature (max. +
min. /2): 20.6 ± 1.8, 16.2–23.7°C (N=173).
Habitat: In Botswana: strong bias to moister NE (1656) compared to arid
SW (156); in Gauteng bushveld: extreme bias to deep sand (129) com-
pared to sandy clay loam (0) and to grassland (118) compared to open
woodland (10) or shaded thicket (1); similar bias in uMkhuze Game
Reserve: deep sand (16.3), sand on clay (1.5), sandy clay loam (0.0),
clay (0.0) with 90% in grassland; only partly supported by DBRU col-
lection records: deep sand (49), sandy loam (21), sandy clay loam (3)
in grassland/pasture (14), scrub/shrubland (17), open woodland (37),
shaded thicket (1).
Food types: In Botswana: attracted primarily to dung (pig: 446, elephant:
631, cattle: 206, sheep: 504) with few to carrion (chicken livers: 25);
somewhat similar at Phalaborwa: pig (51), elephant (16), cattle (49);
DBRU collection records from various dung types: cattle (57), wilde-
beest (1), waterbuck (1), elephant (9), donkey (1), human/cattle mix
(1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); in Gauteng: active from early summer (Oct.)
rising to mid-summer peak (Dec.) tailing off to late summer (Mar.); at
Phalaborwa: much greater activity on a warm evening soon after heavy 2 mm
rainfall (66) than on an evening following a hot, dry day (25) or an
evening following a warm, cloudy day and light rainfall (23); attracted
to light.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: NW Kala-
hari Xeric Savanna (AT1309), Angolan Mopane Woodlands (AT0702),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea
Woodlands (AT0726), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719), Maputaland Coastal Forest
Mosaic (AT0119).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl), Mopane (SVmp) (Savanna Biome);
few records in Dry Highveld Grassland (Gh) (Grassland Biome).
Assessment rationale: EOO = 1 228 465 km2 (estimated); AOO would
be smaller owing to soil type specialisation, but deep sands and spe-
cies range are widespread in southern Africa; bias to open vegetation
suggested by quantitative data so probably not influenced adversely
by clearance of woodland; also readily attracted to various dung types
including that of farm livestock; assessed as Least Concern (LC).
Conservation measures: None recommended at the present time; pro-
tected in Kruger National Park, uMkhuze Game Reserve (South Afri-
ca), Chobe National Park (Botswana).

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Genus Xinidium Harold, 1869a

Type species and designation: Xinidium dentilabris Harold, 1869a, by monotypy.
Synonyms: = Parapinotis Harold, 1878; Type species: Parapinotis dewitzi Harold, 1878, by mono-
typy.
Last review: Generic and species revision (Cambefort 1985a) with one species added subsequently
(Cambefort 1993).

Xinidium Harold, 1869a was created for a single species to which three other species have been added subsequently; one
described under the synonymous generic name, Parapinotis Harold, 1878. The genus is endemic to wet highlands of SE
Africa where three species are endemic to South Africa and one to Zimbabwe. Xinidium species are macropterous and
probably all show flight activity during darkness. All species tunnel from under droppings, but their nesting behaviour is
unknown. Pasture species readily colonise cattle dung.

In South Africa, one species is assessed as Least Concern (LC) as it shows a wide occurrence in grasslands, mainly on wet
E central highlands. The other two are associated with upland and highland forest patches. Although assessed as Data
Deficient (DD), they may deserve an IUCN threat status approaching Vulnerable (VU).

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SURICATA 6 (2020) 259

Xinidium dentilabris
Harold, 1869a

= Copris curvifrons Péringuey, 1885

= Parapinotis dewitzi Harold, 1878, sensu Péringuey, 1901 (pars)
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Port. Natal.’ [Durban, South Africa]; C. curvifrons: ‘Lim-

popo River’ [inexact locality; N South Africa / S Zimbabwe]; P. dewitzi
sensu Péringuey, 1901: ‘Natal (Durban, Estcourt), Southern Rhodesia
(Victoria Falls)’ [South Africa; Zimbabwe; latter citation is undoubt-
edly an error]. ♂
Taxonomy: Accepted species; sexual dimorphism (head, disc of protho-
rax); prominence of characters varies with body size; inexact type lo-
cality and original description suggest C. curvifrons is a synonym of
X. dentilabris Harold, 1869 (known from the Soutpansberg close to
Limpopo River), rather than X. dewitzi; Péringuey’s 1901 description
of P. dewitzi is also X. dentilabris (see Cambefort, 1985).
Distribution: Centred along the E escarpment of Mpumalanga, KwaZulu-
Natal and NE Eastern Cape, South Africa in both cool, wet grasslands
and forest; some coastal records in the S reflected by the type locality;
Péringuey’s, Zimbabwe type locality would represent misidentification
(not Xinidium).
Locality data (mean ± SD, range): Altitude: 1 394 ± 452, 58–2 392 m;
annual rainfall: 838 ± 110, 553–1 271 mm; annual temperature (max.
+ min. /2): 15.5 ± 2.1, 9.9–18.7°C (N=103).
Habitat: No adequate quantitative assessment; along an altitudinal grass-
land gradsect from 500 to 2 800 m in S KwaZulu-Natal; sampled
2 mm
primarily at 1 500 m and not at higher altitude (500 m: 0.2; 1 000 m:
2.2; 1 500 m: 19.1; 1 900 m: 0.2); at Carolina: sampled to cattle dung
in greater numbers in improved Kikuyu pastures (4 163) and fallow
crop fields (2 976) than in natural Themeda grassland (1 433); DBRU
collection records from sand (1), sandy loam (7), sandy clay loam (18),
clay (3) in grassland/pasture (23), forest (2); also fallow crop field (1).
Food types: In upland forest at Mariepskop (1 505–1 554 m): clear bias
to 14 pig dung samples (208) compared to cattle dung (38); DBRU
collection records primarily from cattle dung (31), human/cattle dung
mix (1).
Temporal activity: Flight activity during darkness, primarily during the
summer rainy season (Oct. to May).
Bioregions South Africa: Primarily E Mesic Highveld Grassland (Gm),
N Sub-Escarpment Grassland (Gs) (Grassland Biome); S Indian Ocean
Coastal Belt Biome (CB); marginal in three other bioregions.
Assessment rationale: EOO = 107 815 km2; available data suggest a bias
to finer-grained soils with relatively generalist vegetation associations
characterised by tolerance of pasture improvement and disturbance on
farms where they are attracted to cattle dung in abundance despite a
measured bias to omnivore dung; also occupy a sizeable range; there-
fore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on vegetation associations measured at a
standard altitude around 1 500 m; protected in Golden Gate National
2 mm
Park and Mount Sheba Nature Reserve.
♀

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260 SURICATA 6 (2020)

Xinidium dewitzi
(Harold, 1878)

No synonyms (see taxonomic notes, X. dentilabris, Harold 1869a)

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Parapinotis dewitzi: ‘Cap der guten Hoffnung’ [probably

inaccurate; Cape of Good Hope, South Africa].
Taxonomy: Accepted species; strong sexual dimorphism (head, disc of
prothorax); prominence of characters varies with body size.
Distribution: Presumably forest patches within grasslands on the E es-
♂ carpment, KwaZulu-Natal, South Africa; citation from Zimbabwe is
undoubtedly an error (see taxonomic notes under X. dentilabris Harold,
1869a).
Locality data (mean ± SD, range): Altitude: 1 287 ± 269, 999–1 645 m;
annual rainfall: 813 ± 37, 765–847 mm; annual temperature (max. +
min. /2): 15.1 ± 1.5, 13.1–16.7°C (N=4).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam in pasture; one record from mistbelt forest.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Diel flight activity unknown, but probably in darkness
during the summer rainy season (Jan. to Mar.).
Bioregions South Africa: Three inexact localities map onto Drakensberg
Foothill Moist Grassland (Gs 10) or Midlands Mistbelt Grassland (Gs
9) (Sub-Escarpment Grassland Bioregion, Grand Biome); one exact
locality maps onto Southern Misbelt Forest (FOz 3).
Assessment rationale: EOO = 5 990 km2; AOO probably much small-
er, centred on upland forest patches although quantitative support is
lacking; available observational data in conflict, therefore, assessed as
Data Deficient (DD); however, an IUCN threat category of Vulnerable
2 mm (VU) would be justified due to the low number of known localities
within a small EOO coupled with a single exact locality suggesting
primarily forest associations and a very small fragmented AOO.
Conservation measures: Assessment of conservation status requires quan-
titative data to determine the full EOO, AOO and ecological associa-
tions; one exact locality coincides with 40 ha of indigenous forest on
iNhlosane Mt; unknown if any localities are officially conserved.

2 mm
♀

COPRINI
SURICATA 6 (2020) 261

Xinidium howdeni
Cambefort, 1993

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Afrique du Sud, Natal’, ‘75 km WSW Estcourt, Cathedral

Peaks Forest. Stn’ / ‘Rainbow Gorge...’ [KwaZulu-Natal, South Africa].
Taxonomy: Accepted species; strong sexual dimorphism (head, disc of
prothorax); prominence of characters varies with body size.
Distribution: Drakensberg highlands, NE border of Lesotho and KwaZulu-
Natal, South Africa. ♂
Locality data (mean ± SD, range): Altitude: 1 707 ± 186, 1 500–1 860 m;
annual rainfall: 768 ± 59, 726–810 mm; annual temperature (max. +
min. /2): 10.9 ± 2.5, 9.2–12.7°C (N=2).
Habitat: No quantitative assessment; no soil type data; type locality in
podocarp forest at 1 500 m (altitude possibly underestimated) and
in the veld [?grassland] at 1 860 m; data insufficient to determine if
X. howdeni occurs only in forest patches known to also occur at high
altitude around Botha’s Shelter (Ndedema Gorge) (1 760 m) and the
meteorological station on the Little Berg (1 860 m) (paratype locali-
ties).
Food types: No quantitative assessment; recorded on hyrax dung.
Temporal activity: Diel flight activity unknown, but probably in darkness
during the summer rainy season (Dec.) like other Xinidium species.
Bioregions South Africa: One out of three grid references coincides with
Northern Afrotemperate Forest (FOz 2) (Forest Biome); holotype
locality recorded as forest but grid reference coincides with montane
Drakensberg Grassland (Gd) (Grassland Biome).
Assessment rationale: EOO = ± 20 km2 (presumably grossly underesti-
mated); known from only the five individuals of the type series record-
ed at three localities; occurs at higher mean altitude than close grassland
(X. dentilabris Harold, 1869) and forest relatives (X. dewitzi Harold,
1878) in KwaZulu-Natal; possibly a high altitude forest endemic; if so, 2 mm
AOO would be much smaller than EOO and it could qualify for an
IUCN threat category despite some measure of protection; currently
very poorly known and assessed as Data Deficient (DD).
Conservation measures: A quantitative survey of habitats on the slopes
of the Drakensberg along the SE and NE Lesotho borders is required
to determine the EOO, AOO and ecological associations before an
assessment may be made of conservation status; many FOz 2 forest
patches occur in this region and to the NW as far as S Mpumalanga;
two of three known type localities lie within the Cathedral Peaks State
Forest boundaries.

Photographs: MNHN/A. Mantilleri. ♀ 2 mm

COPRINI
262 SURICATA 6 (2020)

TRIBE GYMNOPLEURINI
Lacordaire, 1856

As Gymnopleurides; Type genus: Gymnopleurus Illiger, 1803.

The tribe, Gymnopleurini, is probably monophyletic. It occurs in the Afrotropical, S to E Palaearctic and
Oriental regions. Species are found in Mediterranean to warm temperate to tropical climates where they
occupy a range of habitats from arid to moist grasslands and savanna with a few in forest. All tribal members
show diurnal, ball-rolling activity.

The Gymnopleurini are currently considered to comprise just four valid genera, of which three occur within
and beyond the Afrotropical region: Gymnopleurus Illiger, 1903; Allogymnopleurus Janssens, 1940a; Garreta
Janssens, 1940a. Paragymnopleurus Shipp, 1897b, is the fourth and most derived genus (Monaghan et al.
2007) with an Oriental and E Palaearctic centre of distribution.
SURICATA 6 (2020) 263

Of three genera and 20 species found in South Africa, Botswana and Namibia, none is considered to be
threatened.
(1) Gymnopleurus Illiger, 1803: Widely distributed genus in Afro-Eurasia represented in southern Africa
by 12 species centred in SW Arid (3), upland grassland (2), Kalahari (1) or savanna (6) regions.
(2) Allogymnopleurus Janssens, 1940a: Widely distributed genus in the Afrotropical and Oriental re-
gions represented by just two savanna species in southern Africa.
(3) Garreta Janssens, 1940a: Widely distributed genus in the Afrotropical and Oriental regions. It was
represented in southern Africa by six species centred on savanna (3) savanna/upland grassland (1),
or E forest (2) when this book was completed in 2017. Subsequently, two new southern African
species were described (Davis & Deschodt 2018).

GYMNOPLEURINI
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Genus Allogymnopleurus Janssens, 1940a

Type species and designation: Gymnopleurus chloris Klug, 1855, by original designation.
Synonyms: None.
Last review: Entire genus reviewed as an illustrated catalogue by Pokorný and Zídek (2009) with
three synonymies.

The genus, Allogymnopleurus Janssens, 1940a, was created by subdivision of Gymnopleurus Illiger, 1803. It is currently
represented by 18 valid species showing primarily savanna and upland grassland distributions in the Afrotropical (16) and
Oriental regions (2).

Relationships have not been investigated by phylogenetic analysis, although three groups may be recognised on the basis
of fore tibial structure in males. One group bears an acute spine at the tip of the inner margin (see A. splendidus (Bertolini,
1849)), one bears a blunt or angular terminal swelling at the tip of the inner margin (see A. consocius (Péringuey, 1901)),
and the other bears a crenulate inner margin. This may or may not have phylogenetic significance and may require re-
interpretation.

Allogymnopleurus species are of moderate body size. They roll balls of dung away from droppings, often in pairs with the
female pulling and male pushing. As the female constructs a single brood spheroid with a slight protuberance above the
egg chamber on each breeding occasion, this strategy may be supported by small amounts of dung.

After the recent synonymy of A. thalassinus (Klug, 1855) and A. chloris (Klug, 1855) with A. splendidus (Bertolini, 1849),
only two species are known to occur in Botswana, Namibia and/or South Africa. Both are centred on southern African
savanna. One shows a large range in the E and smaller range in the W and is assessed as Least Concern (LC). The other is
centred in the E with a range extending onto the Highveld and may have undergone range contraction.

GYMNOPLEURINI
SURICATA 6 (2020) 265

Allogymnopleurus consocius
(Péringuey, 1901)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Transvaal (Potchefstroom), Orange Free State (Bloem-

fontein)’ [South Africa: North-West Province, Free State].
Taxonomy: Accepted species.
Distribution: Drier areas of NW South African Highveld and NE
KwaZulu-Natal lowlands; regional colour variation with coastal indi-
viduals iridescent cupreous and Highveld individuals purple to violet. ♂
Locality data (mean ± SD, range): Highveld: Altitude: 1 440 ± 73,
1 356–1 485 m; annual rainfall: 585 ± 44, 535–619 mm; annu-
al temperature (max. + min. /2): 16.7 ± 0.9, 15.7–17.3°C (N=3).
KwaZulu-Natal/Mpumalanga: Altitude: 202 ± 226, 55–722 m; annual
rainfall: 757 ± 76, 661–900 mm; annual temperature (max. + min. /2):
21.5 ± 0.7, 20.3–22.4°C (N=8).
Habitat: In uMkhuze Game Reserve: extreme bias to clay (143.8) rather
than sandy clay loam (1.4), duplex soils (sand over clay – 0.4) or sand
(0.2); limited support from DBRU collection records: sandy loam (2),
sandy clay loam (1), clay (1) in woodland (4).
Food types: No quantitative assessment; DBRU collection records from
dung of wildebeest (1) and cattle (1).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: Disjunct on Dry Highveld Grassland (Gh)
(Grassland Biome) and S Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 35 250 km2 (full range); AOO = 3 600 km2
(current known range); described from the Highveld in 1901, but since
1970, only recorded on clay in open woodland in game reserves of 2 mm
coastal KwaZulu-Natal, particularly uMkhuze Game Reserve, where it
is often found crowded together on wildebeest droppings, hence the
species name consocius; qualifies for an IUCN threat category because
of possible 90% range contraction to < 5 000 km2, but until that is
confirmed, assessed as Data Deficient (DD).
Conservation measures: Investigations are required to determine if ap-
parent extreme contraction in range is a collection artefact or a true
phenomenon justifying an IUCN threat category; a quantitative survey
is required to confirm or negate its disappearance from former known
localities on the South African Highveld, particularly vegetation types
found in areas of clay soils; owing to observed association with blue
wildebeest dung, research needs to be conducted on the possible role
of wildebeest range contraction; protected in uMkhuze Game Reserve.

GYMNOPLEURINI
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Allogymnopleurus splendidus
(Bertolini, 1849)

= Gymnopleurus chloris Klug, 1855

= Gymnopleurus thalassinus Klug, 1855
= Gymnopleurus coracinus Fahraeus, 1857
= Gymnopleurus subcupratus Fahraeus, 1857
Global: LC (see IUCN Red List – LC)*

J A S O N D J F M A M J

Type localities: As Gymnopleurus splendidus: ‘província Inhambanensi’ [In-

hambane Province, S Mozambique]; G. chloris: ‘Sena’ [Central Mozam-
bique]; G. thalassinus: ‘Tette’ [Tete, Central Mozambique]; G. coracinus:
♂ ‘in terra Natalensi’ [KwaZulu-Natal, South Africa]; G. subcupratus: ‘juxta
fluvium Limpopo’ [near Limpopo River, southern Africa].
Taxonomy: On the basis of precedence, the formerly accepted species
(A. thalassinus, A. chloris) were synonymised with A. splendidus by
Pokorný and Zídek (2009), having been recognised as comprising a
single taxon showing considerable, mostly iridescent, colour variation;
synonym A. thalassinus (red squares): black (N Namibia), cupreous and
green varieties; synonym A. chloris (black squares): comprising shiny
green varieties; varieties overlapping at NE edge of range (blue squares).
Distribution: Sandy savanna in arid upland to moist upland and lowland
areas of southern Africa: N South Africa, N Central Namibia, E Botswa-
na, Mozambique, Zimbabwe, Zambia; citations from Democratic Re-
public of the Congo (DRC), Tanzania require validation; Ethiopia, Ken-
ya citations would represent confusion with A. zavatarii Müller, 1947.
Locality data (mean ± SD, range): Altitude: 956 ± 382, 0–1 958 m; an-
nual rainfall: 477 ± 248, 133–1 659 mm; annual temperature (max. +
min. /2): 19.5 ± 2.0, 13.4–25.9°C (N=413).
Habitat: On Gauteng bushveld: very strong bias to coarse-grained soils
(deep sand: 198; sandy clay loam: 1) in open vegetation (grassland:
5 mm
172; open woodland: 27; shaded thickets: 0); limited support from
DBRU collection records on sand (38), sandy loam (28), sandy clay
loam (11) in grassland/pasture (20), scrub/shrubland (26), open wood-
land (28).
Food types: At Phalaborwa: strong bias to omnivore compared to herbi-
vore dung (pig: 1 046, elephant: 178, cattle: 358); DBRU collection
records on dung of cattle (71), wildebeest (4), elephant (5), rhinoceros
(3), zebra (1), warthog (1), horse (6), donkey (3), impala (1), human/
cattle mix (1).
Temporal activity: Diurnal flight activity, primarily in the summer rainy season (Oct. to May); on Gauteng bushveld: peak activ-
ity early to mid-summer (Oct. to Jan.), tailing off in late summer (Feb. to Mar.); at Phalaborwa: greater abundance on a hot,
dry day (944) than on a warm sunny day following heavy rainfall (380) or a warm cloudy day following light rainfall (129).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: NW Kalahari Xeric Savanna (AT1309), Southern African Bush-
veld (AT0717), Zambezian and Mopane Woodlands (AT0725), Southern Miombo Woodlands (AT0719), Maputaland
Coastal Forest Mosaic (AT0119), margins of other ecoregions.
Bioregions South Africa: Eastern Kalahari Bushveld (SVkb), Central Bushveld (SVcb), Mopane (SVl), Lowveld (SVl), N
Sub-Escarpment Savanna (SVs) (Savanna Biome); NE Bushmanland (NKb), N Upper Karoo (NKu) (Nama Karoo Biome);
N Dry Highveld Grassland (Gh) (Grassland Biome); N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 523 410 km2; SW range coincides with arid, little-transformed areas used primarily for grazing
of domestic livestock (0–2% NK, W SVkb); NE range in regions with greater transformation (0–58% SV); at Phalaborwa:
abundance greater in reserves (332) and farm rangeland (1 167) than in a mined area (83); assessed as Least Concern (LC).
*IUCN Red List entry only for A. chloris.
Conservation measures: None required since A. splendidus is widespread and often abundant in farm rangeland; protected in
various reserves including Kruger National Park and uMkhuze Game Reserve (South Africa).

GYMNOPLEURINI
SURICATA 6 (2020) 267

Genus Garreta Janssens, 1940a

Type species and designation: Ateuchus azureus Fabricius, 1801, by original designation.
Synonyms: None.
Last review: Entire genus created and reviewed by Janssens (1940a); three new species subsequently
added by Balthasar (1961b), Frey (1967, now synonymised) and Mittal (2011); two
species complexes radically revised by Moretto and Génier (2015) with one synonymy
in a third species group; revision of African species (Pokorný & Zídek 2018); two new
species added by Davis and Deschodt (2018).

The genus, Garreta Janssens, 1940a, was created by subdivision of Gymnopleurus Illiger, 1803. When this book was com-
pleted at the end of 2017, the genus was represented by 23 valid species showing savanna or forest distributions in the
Afrotropical (13) and Oriental regions (10). Subsequently, revision of the African component (Pokorný & Zídek 2018)
and addition of species (Davis & Deschodt 2018) raises the African fauna to 15 species. There is no phylogeny nor have
any groups been formally defined for the genus.

Garreta species are of moderate body size and are all macropterous showing diurnal flight activity. They roll balls of dung
away from droppings, often in pairs with the female pushing and male pulling. As the female constructs a single brood
spheroid with a slight protuberance above the egg chamber on each breeding occasion, this strategy may be supported by
small amounts of dung.

Following the partial revision of Moretto and Génier (2015), six valid species are listed, here, for South Africa, Botswana
and Namibia. Changes to the validity of some of these species following subsequent revision (Pokorný & Zídek 2018)
or addition of new species (Davis & Deschodt 2018) are noted in the relevant species accounts. Although G. fastiditus
(Harold,1867b) remains listed as valid by Pokorný & Zídek (2018), it is omitted from the species accounts for southern
Africa as its type locality is considered an error. It may be a synonym of the Oriental G. mundus (Wiedemann, 1819).

Described southern African species are centred in moist savanna (4) or forest (2). The savanna species are centred in the
west (1), east (2) or east extending onto the Highveld (1). Both forest species are centred in the east. Two species from
savanna (1) or forest (1) show distributions extending into the tropics. All but one species show widespread occurrence
and are assessed as Least Concern (LC – 5) or Data Deficient (DD – 1).

GYMNOPLEURINI
268 SURICATA 6 (2020)

Garreta australugens
Davis & Deschodt, 2018

Global: DD

J A S O N D J F M A M J

Type locality: Wildlife College, Site 3, S24°32’27” E31°20’23” [South

African Wildlife College, Kruger National Park, South Africa].
Taxonomy: Accepted species from southern Africa; recently separated
from Garreta lugens (Fairmaire, 1891) described from East Africa ‘la
côte orientale d’Afrique vers le 1er degré de latitude nord…entre les
pays Somalis et le pays de Massai’ [probably NE lowlands of central
Kenya].
Distribution: Moderately dry, hot, eastern lowlands in southern Africa:
Mozambique, E Zimbabwe, NE South Africa.
Locality data (mean ± SD, range): Altitude: 405 ± 301, 96–1 020 m;
annual rainfall: 659 ± 318, 383–1 177 mm; annual temperature (max.
+ min. /2): 22.6 ± 2.0, 19.0–24.8°C (N=7).
Habitat: No quantitative assessment; one DBRU collection record from
sandy clay loam; near Orpen, Kruger National Park: three individuals
sampled on sandy clay loam in very open woodland.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov., Dec.).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Centred on Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 88 890 km2; widespread with population
density apparently low even in the reserves where most of the reference
material has been sporadically recorded; therefore, poorly known and
assessed as Data Deficient (DD).
5 mm Conservation measures: Before an assessment may be made of conserva-
tion status, a survey of E lowlands is required to generate quantitative
ecological data; currently protected in several national parks: Kruger
(South Africa), Gorongosa (Mozambique).

GYMNOPLEURINI
SURICATA 6 (2020) 269

Garreta caffer
(Fahraeus, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Gymnopleurus caffer: ‘in terra Natalensi’ [KwaZulu-

Natal, South Africa].
Taxonomy: Accepted species but exoskeleton sculpture differs only very
slightly from that of the tropical south central African G. laetus subsp.
olivaceus (Quedenfeldt, 1884) and Lake Malawi G. nyassicus (Kolbe,
1897); see notes on viewpoint of Pokorný & Zídek (2018) in section
9 of Preamble; support from a molecular phylogeny would be useful;
G. caffer shows iridescent colour variation: mostly green, but also cu-
preous.
Distribution: Vegetation offering shade on the coastal deep sands of SE
Africa: NE KwaZulu-Natal, South Africa; SE Mozambique (but see
Taxonomy).
Locality data (mean ± SD, range): Altitude: 12 ± 23, 0–88 m; annual
rainfall: 849 ± 114, 659–1 010 mm; annual temperature (max. + min.
/2): 22.0 ± 0.5, 20.5–22.5°C (N=26).
Habitat: On deep sand at Richards Bay: recorded exclusively under shad-
ed vegetation following mining, either regenerating or natural: grass/
shrubland (0), younger woodland (17), older woodland (13), natural
dune forest (32); on deep sand in Maputo Special Reserve: sampled
primarily in cool coastal-fringing dune forest (24.6/trap) rather than
warmer inland sand forest (large patch: 0.5, medium: 1.0, small: 0.1) or
grassland (0); limited DBRU collection records from sand (1) in forest
(1), grassland (1).
Food types: No quantitative assessment; no DBRU collection records;
5 mm
sampled to: mixed pig/cattle dung bait.
Temporal activity: Diurnal flight activity, primarily in the summer rainy
season (Oct. to Feb.); also Aug. in warm coastal range.
Ecoregions Mozambique: Centred on forest in Maputaland Coastal For-
est Mosaic (AT0119).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 8 394 km2; AOO would be smaller due
to habitat specialisation; largely centred on scattered patches of forest,
particularly along the cooler coastline; all known records from deep
sands; relatively small known range in a densely populated area, but
assessed as Least Concern (LC) owing to extensive protection.
Conservation measures: Conservation status would be dependent on
continuing protection of moist forest patches on the SE coast; as-
sessment of status would be improved by quantitative data on AOO
and associations with soil and dung type; protected in iSimangaliso
Wetland Park (World Heritage Site) (South Africa), Maputo Special
Reserve (Mozambique).

GYMNOPLEURINI
270 SURICATA 6 (2020)

Garreta laetus
(Hope, 1842)

subsp. laetus (Hope, 1842)

= Gymnopleurus cupreovirens Kolbe, 1895
= Gymnopleurus azureus var. cupreus Kolbe, 1914
= Gymnopleurus (Paragymnopleurus) azureus var. ebenus Janssens,
1938
= Gymnopleurus (Paragymnopleurus) azureus var. rubrocupreus Jans-
sens, 1938
subsp. olivaceus (Quedenfeldt, 1884)
= Gymnopleurus insidiosus Péringuey, 1901 (but see Taxonomy)
Global: LC (see IUCN Red List – LC as G. azureus (Fabricius, 1801))*

J A S O N D J F M A M J

Type localities: subsp. laetus: as Gymnopleurus laetus: ‘vicinity of Cape Pal-

mas’ [Liberia]; G. cupreovirens: ‘südöstliche vom Victoria-Njansa’ [SE
of Lake Victoria-Nyanza, Tanzania]; var. viridimicans: ‘Undussuma,
südwestlich vom Albert-Nyansa’ [SW of Lake Albert, E central Africa];
var. cupreus: ‘Am Westufer der Russisi, Nord-Tanganjika.... Kiwu-See
(?)’ [? Rusisi River, Lake Kivu, NW Tanzania]; var. ebenus: ‘N.W.Tan-
ganika’ [NW Tanzania]; var. rubrocupreus: Not stated; subsp. olivaceus:
as Gymnopleurus olivaceus: ‘Malange’ [N Angola]; G. insidiosus: ‘South-
ern Rhodesia (Salisbury)’ [Harare, Zimbabwe].
Taxonomy: Accepted species formerly listed under the name G. azureus
(Fabricius, 1801) with exoskeleton sculpture that differs only very
slightly from that of G. caffer (Fahraeus, 1857) on the SE African coast
and G. nyassicus (Kolbe, 1897) from the environs of Lake Malawi; cited
synonyms are those considered valid in a recent revision (Moretto &
Génier 2015); however, the subspecies olivaceus is synonymised with
G. laetus in the most recent review of Pokorný & Zídek (2018); molec-
ular phylogenetic support would be useful; G. laetus shows iridescent
colour variation: green, cupreous.
Distribution: subsp. laetus (range in green): W to E tropical Africa; subsp.
olivaceus (range in black): tropical Africa S of the equatorial Congo
5 mm rain forest reaching the extreme NE of South Africa: also Gabon, S
Democratic Republic of the Congo (DRC), Angola, S Tanzania, N
Mozambique, Zambia, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 754 ± 594, 0–1 478 m; annual rainfall: 913 ± 275, 386–1 233 mm; annual
temperature (max. + min. /2): 21.2 ± 3.1, 16.7–25.5°C (N=18).
Habitat: No quantitative assessment; DBRU collection records from sand (1) in forest (5).
Food types: No quantitative assessment; DBRU collection records from mixed human/cattle or buffalo dung bait (3), elephant
dung (1).
Temporal activity: Diurnal flight activity, primarily in the summer rainy season (Sept. to May).
Ecoregions Zimbabwe, Mozambique: Centred on shaded vegetation in Southern Miombo Woodlands (AT0719), Southern
Zanzibar-Inhambane Coastal Forest Mosaic (AT1028), Eastern Zimbabwean Forest Grassland Mosaic (AT1006); scattered
records in shaded vegetation of Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Marginal occurrence in shaded vegetation of Lowveld (SVl) (Savanna Biome).
Assessment rationale: subsp. olivaceus EOO = 2 848 875 km2; widespread in southern tropical Africa, but as a forest specialist,
AOO would be much smaller and subject to threat in some areas; however, owing to large range, currently assessed as Least
Concern (LC); *IUCN Red List entry was assessed under the name Garreta azureus (Fabricius, 1801) before the revision of
Moretto and Génier (2015) and incorporates both G. laetus and G. caffer (Fahraeus, 1857).
Conservation measures: Assessment of conservation status would be improved by quantitative data on ecological associations
and a survey of forest patches to determine the AOO; protected in Gorongosa National Park (Mozambique), Lake Mutirikwe
Game Reserve (Zimbabwe), dense woodland in extreme NE of Kruger National Park (South Africa).

GYMNOPLEURINI
SURICATA 6 (2020) 271

Garreta nitens
(Olivier, 1789)

= Ateuchus azureus Fabricius, 1801 (but see Taxonomy)

= Gymnopleurus hilaris Hope, 1842
= Gymnopleurus janthinus Castelnau, 1840
= Gymnopleurus splendens Castelnau var. coeruleovirens Kolbe, 1897
= Gymnopleurus splendens Castelnau var. gracilipes Kolbe,1897
= Gymnopleurus splendens Castelnau var. kilimana Kolbe, 1897
Global: LC (see IUCN Red List – LC)*

J A S O N D J F M A M J

Type localities: As Scarabaeus nitens: ‘Sénégal’; A. azureus: ‘Guinea’ [in-

terpreted as Ghana]; G. hilaris: ‘Sierra Leone’; G. janthinus: ‘Barbarie,
Tripoli’ [Libya; probably inaccurate]; G. splendens var. coeruleovirens:
‘Nyassa See’ [Lake Malawi, Tanzania]; var. gracilipes: ‘südöstliche vom
Victoria-Nyansa’ [SE of Lake Victoria-Nyanza, Tanzania]; var. kilima-
na: ‘Kilimandscharo-Gebiet’ [Mt Kilimanjaro region, Tanzania].
Taxonomy: Accepted species, but exoskeleton sculpture differs only very
slightly from the SE African G. wahlbergi (Fahraeus, 1857), which has
been considered a synonym until recently (Moretto & Génier 2015),
and has now been synonymised once more with G. nitens (Pokorný &
Zídek 2018); cited synonyms are those considered valid in the 2015
revision; however, Ateuchus azureus has since been revalidated (Pokorný
& Zídek 2018) as Garreta azureus (Fabricius, 1801), a species apart
from G. nitens; support from a molecular phylogeny would be useful;
G. nitens shows iridescent colour variation from cupreous to green to
blue.
Distribution: Varieties in three disjunct centres; (1) Dry to moist savanna
of N Namibia, Angola: population may be isolated to SW of mega-
Kalahari sands, disjunction within N Namibia probably real at moist-
er localities on finer-grained soils separated by deep sands to N and
NW of Etosha Pan; (2) Moist savanna of S Central to E to W Africa:
24 countries including Democratric Republic of the Congo (DRC), 5 mm

Zambia, Zimbabwe, Malawi, N Mozambique, Tanzania, Kenya, South

Sudan, Nigeria, Benin, Togo, Ghana, Senegal; (3) NE Africa: Eritrea,
Ethiopia.
Locality data (mean ± SD, range): Altitude: 1 282 ± 307, 683–2 196 m;
annual rainfall: 545 ± 341, 356–2 097 mm; annual temperature (max.
+ min. /2): 20.4 ± 2.1, 13.8–23.2°C (N=26).
Habitat: No quantitative assessment; DBRU collection records biased to
finer-grained soils: sand (1), sandy loam (2), sandy clay loam (2) in
wooded habitat: shrubland (1), open woodland (4).
Food types: No quantitative assessment; DBRU collection records on herbivore dung: cattle (5), elephant (1), zebra (1).
Temporal activity: Diurnal flight activity in the summer rainy season (Nov. to Apr.).
Ecoregions Namibia, Zimbabwe: Namibia: NE Namibian Savanna Woodlands (AT1316), S Angolan Mopane Woodlands
(AT0702), margins of two other ecoregions; Zimbabwe: N Southern Miombo Woodlands (AT0719) (not shown on map).
Assessment rationale: EOO = 4 761 275 km2; further study would be useful to determine if exoskeleton structure varies along
one or more clines within the range of G. nitens as well as across the transition to G. wahlbergi; AOO would be smaller than
EOO due to bias to finer-grained soils and woody vegetation; loss of woodland would be a threat across its range; however,
assessed as Least Concern (LC) because of very large EOO and coincidence with several reserves; *IUCN Red List entry for
G. nitens includes data for G. wahlbergi (Fahraeus, 1857) as the assessment predates the recent 2015 revision.
Conservation measures: Assessment of conservation status would be improved by quantitative data on ecological associations;
protected in various national parks, including Etosha (Namibia).

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Garreta unicolor
(Fahraeus, 1857)

= Gymnopleurus delagorguei Waterhouse, 1890

Global: LC

J A S O N D J F M A M J

Type localities: As Gymnopleurus unicolor: ‘Caffraria’ [SE South Africa];

G. delagorguei: ‘Port Natal’ [Durban, South Africa].
Taxonomy: Accepted species, but taxonomy requires revision; often mis-
identified as Garreta fastiditus (Harold, 1867), which was described
as originating from SE Africa, but type material is synonymous with
an Oriental species, G. mundus (Wiedemann, 1819) (P. Moretto pers.
comm.) (see generic notes).
Distribution: Centred on warmer, moist uplands and coastal hills border-
ing and encroaching onto the W, N and E South African Highveld (e.g.
Magaliesberg, Waterberg, Blouberg, Soutpansberg, NE escarpment);
marginal records suggest a distribution continuing N along moister
parts of the SE Mozambique coastline; Zimbabwe citation would be
G. matabelensis Janssens, 1938.
Locality data (mean ± SD, range): Altitude: 1 111 ± 626, 0–2 922 m;
annual rainfall: 797 ± 114, 576–1 048 mm; annual temperature (max.
+ min. /2): 17.3 ± 2.8, 6.6–21.8°C (N=98).
Habitat: No quantitative soil assessment; on pig dung at Ithala Game
Reserve, more abundant in highland grassland (191) than in mid-
altitude grassland (114) or forest (4), lowland shrubland (74) or wood-
land (43); on Wolkberg at 1 400 m: extreme bias to grassland (139),
forest (0); DBRU collection records biased to moderately fine-grained
soils: clay (2), sandy clay loam (8), sandy loam (12), sand (5) in grass-
land/pasture (20), shrubland (3), open woodland (5), forest (1).
Food types: At Ithala Game Reserve: strong attraction to both omnivore
5 mm and ruminant herbivore dung: pig (554), cattle (266), horse (61);
DBRU collection records from various dung types: human/cattle mix
(4), rhinoceros (2), horse (1), wildebeest (1), cattle (25).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: In N: centred on moister vegetation units at the
edge of the Grassland and Savanna Biomes, i.e. upland edges of Mesic
Highveld Grassland (Gm), Central Bushveld (SVcb) and Lowveld
(SVl) Bioregions, including mountains in the Central Bushveld; in SE:
centred on Sub-Escarpment grassland (Gs) (Grassland Biome), Indian
Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 209 670 km2; widespread and readily
attracted to dung of domestic livestock although the range mostly
coincides with vulnerable and endangered natural vegetation; transfor-
mation is extensive in some units of the highland grasslands where it
is most abundant (Gm 33–67%, Gs 10–25%); however, recorded in
various reserves so assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil associations and the impact of
transformation through pasture improvement or creation of crop fields
and tree plantations; protection offered in various game and nature
reserves, including Hluhluwe–iMfolozi, Ithala, uMkhuze (sandy clay
loam in open woodland on Lebombo Mts), Abe Bailey, Suikerbosrand
and Vernon Crookes.

GYMNOPLEURINI
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Garreta wahlbergi
(Fahraeus, 1857) (but see Taxonomy)

= Garreta zumpti Frey, 1967

Global: LC
J A S O N D J F M A M J

Type localities: As Gymnopleurus Wahlbergi: ‘in terra Natalensi’, lectotype:

‘Caffra / ria’ [KwaZulu-Natal, South Africa]; G. zumpti: ‘Transvaal,
Rüstenburg und Umfolozi Zululand’ [South Africa: Rustenburg,
North-West Province; iMfolozi Game Reserve, KwaZulu-Natal].
Taxonomy: Accepted species (in 2017), but exoskeleton sculpture differs
only very slightly from the tropical W, E and south central African
G. nitens (Olivier, 1789) with which it has been synonymised until
recently (Moretto & Génier 2015); now, synonymised once more by
Pokorný and Zídek (2018), who regard the sculpture as intraspecific
variation within G. nitens; support from a molecular phylogeny would
be useful; G. wahlbergi shows iridescent colour variation from cupreous
to green to blue.
Distribution: Widespread on finer-grained soils in woodland savanna
of SE Africa: South Africa, Eswatini, NE Botswana, Zimbabwe, S
Zambia, S Mozambique; Kalahari deep sands probably act as a barrier
between ranges of G. nitens in N Namibia and G. wahlbergi to the E.
Locality data (mean ± SD, range): Altitude: 857 ± 441, 3–1 995 m; an-
nual rainfall: 643 ± 142, 300–1 177 mm; annual temperature (max. +
min. /2): 20.3 ± 2.3, 13.4–25.9°C (N=209).
Habitat: In Gauteng bushveld: strong bias to finer-grained soils: sandy
clay loam (1 125), deep sand (12) in woody vegetation: shaded thickets
(424), open woodland (634), grassland (75); similar bias in uMkhuze
Game Reserve: sand (0.1), duplex soil (sand over clay) (2.4), sandy clay
loam (7.6), clay (3.9); supported by DBRU collection records: clay (1),
sandy clay loam (13), sandy loam (9), sand (1) in forest/thicket (11),
open shrub/woodland (12), grassland/pasture (6).
5 mm
Food types: At Phalaborwa: strong bias to dung of omnivores compared to
herbivores: pig (332), cattle (50), elephant (25); DBRU collection records
on various dung types: human/cattle mix (11), baboon (1), cattle (21), buf-
falo (2), wildebeest (1), elephant (10), rhinoceros (2), hippopotamus (1).
Temporal activity: Diurnal flight activity during the summer rainy season; in Gauteng bushveld: peak activity in early to
mid-summer (Oct. to Jan.) tailing off into late summer (Feb. to Apr.); also recorded in autumn (May) at warmer coastline.
Ecoregions Botswana, Zimbabwe, Mozambique: Centred on Southern African Bushveld (AT0717), Zambezian and Mopane
Woodlands (AT0725), S Southern Miombo Woodlands (AT0719); margins of adjoining ecoregions.
Bioregions South Africa: Centred on Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome); margins of
four adjoining bioregions.
Assessment rationale: EOO = 737 635 km2; readily attracted to cattle dung despite quantitative bias to omnivore dung; AOO
biased by habitat specialisation to finer-grained soils and woody vegetation, which accounts for patchy distribution and
absence from Kalahari deep sands; although clearance of woodland would be detrimental, assessed as Least Concern (LC)
owing to widespread distribution across both farms and reserves; in South Africa, transformation, through cultivation varies
from 0–58% (SVl), 2–49% (SVcb), 0–20% (SVmp).
Conservation measures: Conservation dependent on continuing conservation of woodland savanna on finer-grained soils;
protected in Kruger National Park, uMkhuze and Hluhluwe–iMfolozi game reserves (South Africa), Hwange National Park
(Zimbabwe).

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Genus Gymnopleurus Illiger, 1803

Type species and designation: Scarabaeus geoffroyi Fuesslin, 1775, by subsequent designation (Reiche 1841).
Synonyms: = Symnopleurus Gebler, 1841, considered an error: misspelling of Gymnopleurus.
= Spinigymnopleurus Shipp, 1897b: Type species: Gymnopleurus tristis Castelnau, 1840,
by original designation.
Last review: Entire genus reviewed by Janssens (1940a); new Afrotropical species added, subse-
quently: Ferreira (1954b), Balthasar (1963c), Endrödi (1976), Davis (1986); revision
of one species group (Davis & Génier 2007).

Gymnopleurus Illiger, 1803 shows a distribution centred on the tropics and warm temperate regions of Afro-Eurasia. After
removal of species into other genera in 1897 and 1940, it currently comprises 56 valid species found in the Afrotropical
(35), Oriental (6), Palaearctic (12) regions, or shared between the Oriental and Palaearctic regions (3). These species in-
clude 10 now classified in the subgenus Metagymnopleurus Kabakov, 2006, which are centred on a single region or shared
between regions as for the genus as a whole (4, 2, 1 or 3, respectively). There is no phylogeny to support relationships
within the genus.

Gymnopleurus species are of moderate body size. They roll balls of dung away from droppings, often in pairs with the fe-
male pulling and male pushing. As a single pear-shaped brood is constructed in a chamber at the end of a shallow tunnel,
they are able to breed using small amounts of dung. A number of species is found in arid regions where large droppings
are often absent. Available data suggest a bias to omnivore dung in savanna species.

Gymnopleurus species are macropterous showing diurnal flight activity. A total of 12 valid species is known from South
Africa, Botswana and Namibia although the validity of one species from central Namibia requires further investigation.
Distributions of the remaining 11 species are mainly centred on hotter regions with none recorded from the winter and
bimodal rainfall regions. Several are centred on the arid SW (3) or the Kalahari and pre-Namib sands (1). The remainder
are mostly found in savanna, either centred in the east (4), widespread (1), widespread extending onto the Highveld (1), or
restricted to the Highveld (1). Distributions of four savanna species extend northwards into the tropics, two characterised
by disjuctions between N and S occurrences (although see taxonomic notes for G. pumilus Reiche, 1850).

Gymnopleurus species mostly occupy relatively large ranges. They are often abundant, particularly in drier, little-
transformed regions. They have all been assessed as Least Concern (LC), except for one, particularly, poorly known species
that is assessed as Data Deficient (DD).

GYMNOPLEURINI
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Gymnopleurus aenescens
Wiedemann, 1821

= Scarabaeus bufo (Macleay, 1821)

= Gymnopleurus cupreus Fahraeus, 1857
= Gymnopleurus aenescens Wiedemann var. capensis Ferreira, 1954c
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: G. aenescens: ‘Prom. bon. sp.’ [Cape of Good Hope, South
Africa]; S. bufo: ‘Cap Bonae Spei’ [Cape of Good Hope]; G. cupreus:
‘prope fluvium Limpopo’ [close to Limpopo River, southern Africa];
var. capensis: ‘Província do Cabo: Vryburg’ [South Africa: Vryburg,
North-West Province].
Taxonomy: Accepted species but individuals from the Kaokoveld (arid
NW Namibia) should be compared with G. vanderkelleni van Lans-
berge, 1886 that was probably described from Angola and is currently
synonymised in error with G. virens Erichson, 1843 (see, also, notes in
Taxonomy section of G. virens).
Distribution: Sandy soils in open vegetation of southern African dry sa-
vannas: N South Africa, Namibia, Botswana, SW Angola, W Zimbab­
we, S. Mozambique; outlier record from Gamkaberg (black square)
requires validation.
Locality data (mean ± SD, range): Altitude: 978 ± 293, 9–1 782 m; an-
nual rainfall: 354 ± 151, 22–738 mm; annual temperature (max. +
min. /2): 19.8 ± 1.9, 13.3–23.4°C (N=157) (red squares only).
Habitat: On Gauteng bushveld: extreme association with coarse-grained
soils: deep sand (19), sandy clay loam (0) and strong bias to open veg-
etation: grassland (14), open woodland (5), shaded thickets (0); part-
ly supported by DBRU collection records: sand (12), sandy loam (5),
sandy clay loam (1) in grassland (5) and scrub/shrub/woodland (10). 2 mm

Food types: At Phalaborwa: strong bias to dung of omnivores compared

to herbivores: pig (179), elephant (27), cattle (48); DBRU collection
records on ruminant: cattle (10) and monogastric herbivore dung: ele-
phant (1), rhinoceros (1), zebra (1), donkey (1).
G. aenescens
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.); at Phalaborwa: much greater activity in warm sunshine
after heavy rainfall (195) than on a hot dry day (49) or cool cloudy day
after light rainfall (8).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Namibian
Savanna Woodlands (AT1316), Kalahari Xeric Savanna (AT1309),
Angolan Mopane Woodlands (AT0702), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Southern African Bushveld (AT0717), Zambe-
zian and Mopane Woodlands (AT0725), Maputaland Coastal Forest
Mosaic (AT0119); marginal in four other ecoregions.
Bioregions South Africa: Centred on Mopane (SVmp), Central Bushveld
(SVcb), Eastern Kalahari Bushveld (SVk) (Savanna Biome); marginal in
NE Bushmanland (NKb), N Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 1 407 890 km2 (red squares only); wide-
spread in drier sandy savanna, much of which is relatively untrans-
formed and used for grazing of livestock to whose dung G. aenescens is
frequently attracted despite a bias to omnivore dung; in South Africa:
transformation limited in SW of range (0–2%, W SVk), greater in cen-
tre and NE (13–42%, E SVk; 3–48%, SVcb; 0–20%, SVmp).
Conservation measures: None recommended; widespread in farm range- 2 mm

land and game reserves; protected in Kruger National Park (South Af-
rica), Chobe National Park (Botswana).
G. vanderkelleni

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Gymnopleurus andreaei
Ferreira, 1954b

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Prieska (Província do Cabo)’ [South Africa: Prieska,

Northern Cape].
Taxonomy: Accepted species; mostly uniformly deep purple; some green-
ish individuals at S edge of range.
Distribution: Arid late summer rainfall region in SW southern Africa;
centred on climate types III1 and III2 of Walter and Lieth (1964); N
central South Africa, W Namibia, SW Angola; less abundant at S edge
of range than warmer N.
Locality data (mean ± SD, range): Altitude: 1 003 ± 220, 255–1 818 m;
annual rainfall: 188 ± 54, 35–354 mm; annual temperature (max. +
min. /2): 18.6 ± 1.4, 14.0–21.1°C (N=143).
Habitat: No quantitative assessment; in Northern Cape: centred on hot,
arid Bushmanland (4.6/trap) rather than SW Kalahari deep sands (0.7)
or cooler N Upper Karoo (0.5) although only shows a slight bias to finer-
grained soils: sand (3.0/trap, n = 46 sites), sandy loam (6.7, n = 19);
not supported by DBRU collection records: biased to sand (17), rather
than sandy loam (2), in open vegetation: grassland/pasture (9), scrub/
shrubland (8), open woodland (2).
Food types: No quantitative assessment; DBRU collection records: dung
of baboon (1), donkey (3), horse (3), cattle (8), goat and sheep (1).
Temporal activity: Diurnal flight activity during the late summer rainy
season (Feb. to Apr.).
Ecoregions Namibia: Centred on Namibian Savanna Woodlands
(AT1316); marginal occurrence in two adjoining ecoregions.
Bioregions South Africa: Centred on arid NE Bushmanland (NKb)
(Nama Karoo Biome); marginal occurrence in two adjoining biore-
2 mm gions.
Assessment rationale: EOO = 263 705 km2; widespread; entirely restrict-
ed to an arid area where transformation is limited and land usage is
dominated by grazing of domestic livestock, primarily sheep; as collec-
tion records suggest fairly generalist soil and food associations, G. an-
dreaei would face no threats; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and dung type associations; pro-
tected in Iona National Park (Angola).

GYMNOPLEURINI
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Gymnopleurus asperrimus
Felsche, 1909

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Deutsch-Südwest-Afrika’ [Namibia].

Taxonomy: Accepted species.
Distribution: Straddling borders of Namibia, Botswana and South Africa
in southern part of the arid late summer rainfall region: climate types
III1 and II(III)a of Walter and Lieth (1964).
Locality data (mean ± SD, range): Altitude: 1 020 ± 137, 684–1 508 m;
annual rainfall: 206 ± 43, 114–353 mm; annual temperature (max. +
min. /2): 18.9 ± 1.2, 14.7–21.3°C (N=187).
Habitat: No adequate quantitative assessment; in grassland, scrub and
shrubland of the Northern Cape: primarily in SW Kalahari (6.8/trap),
Bushmanland (13.4), and warm Karoo S of the River Orange (5.6) on
sand (5.6/trap) and sandy loam / sandy clay loam (5.2); only partly
supported by DBRU collection records on sand (21), sandy loam (2)
in open woodland (1), scrub/shrubland (12), grassland/pasture (10).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (12), donkey (3), horse (2).
Temporal activity: Diurnal flight activity during the late summer rainy
season in the arid, late summer rainfall region (Jan. to Apr.).
Ecoregions Namibia: SW Kalahari Xeric Savanna (AT1309), Nama
Karoo (AT1314).
Bioregions South Africa: Kalahari Duneveld (SVkd) (Savanna Biome); E
Bushmanland (NKb), N Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 223 500 km2; shows an appreciable range
in an arid area used primarily for the grazing of livestock where trans-
formation is minimal (SVkd, NKb, NKu); also readily attracted to the
dung of domestic livestock; assessed as Least Concern (LC).
2 mm
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tected in the South African part of Kgalagadi Transfrontier Park.

GYMNOPLEURINI
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Gymnopleurus humanus
Macleay, 1821

= Gymnopleurus sericatus Erichson 1843

= Gymnopleurus modestus van Lansberge, 1886
= Gymnopleurus modestus Péringuey, 1888
= Gymnopleurus peringueyi Shipp, 1895b
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: G. humanus: ‘Cap. Bon. Spei.’ [Cape of Good Hope,

South Africa]; G. sericatus: ‘Angola’; G. modestus van Lansberge: ‘Ben-
guela’ [Angola]; G. peringueyi nom nov. for G. modestus Péringuey:
‘Beaufort West, Cape Colony’ [Western Cape, South Africa].
Taxonomy: Accepted polymorphic species; exoskeleton either blue, green
or cupreous; as in other iridescent species, colour produced by selective
reflection of different wavelengths of light by layers in the exoskeleton.
Distribution: Centred on the arid late summer rainfall region (climate
types III1, III2, II(III)a, II(III)b, II4a) and arid parts of the bimodal
rainfall region (III5) of Walter and Lieth (1964): SW central South
Africa, Namibia, SW Angola; colour polymorphism correlates strong-
ly with the cool, dry season, latitudinal temperature gradient: blue in
cooler S; locally co-occurring blue, green or cupreous in N central Cape
uplands; cupreous in warmer N; it has been suggested that differences
in exoskeleton colour may be driven by the temperatures experienced
during development (see Davis et al. 2008).
Locality data (mean ± SD, range): Altitude: 1 106 ± 282, 113–2 264 m;
annual rainfall: 231 ± 86, 22–556 mm; annual temperature (max. +
min. /2): 17.9 ± 1.9, 11.2–22.4°C (N=380).
Habitat: No adequate quantitative assessment; in Northern Cape: bias to
hot arid Bushmanland (56.0/trap) compared to warm arid N Upper
Karoo (24.5) or arid SW Kalahari deep sands (9.7); in Bushmanland:
slight bias to finer-grained soils, sandy loam (68.4/trap), loamy sand
2 mm (69.6), deep sand (37.6); opposite soil trend in DBRU collection re-
cords: sand (31), sandy loam (17), sandy clay loam (7) in grassland/
pasture (13), scrub/shrubland (24), open woodland (12).
Food types: No quantitative assessment; DBRU collection records on
dung of baboon (1), zebra (2), horse (5), donkey (2), cattle (36), sheep
(2); sampled in abundance to pig dung.
Temporal activity: Diurnal flight activity in bright sunshine during the
late summer rainy season (Dec. to May).
Ecoregions Namibia: Centred on Nama Karoo (AT1314), Namibian
Savanna Woodlands (AT1316); also W margins of Kalahari Xeric Sa-
vanna (AT1309) and margins of four other ecoregions.
Bioregions South Africa: Bushmanland (NKb), Upper Karoo (NKu),
Lower Karoo (NKl) (Nama Karoo Biome); W Eastern Kalahari Bush-
veld (SVkb) (Savanna Biome).
Assessment rationale: EOO = 587 750 km2; widespread across a little-
transformed arid region used primarily for the grazing of domestic live-
stock to whose dung G. humanus is readily attracted; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by additional quantitative ecological data, but in terms of abun-
dance, a dominant species across much of its wide, little-transformed
range; protected in several national parks, Namib-Naukluft and drier
W of Etosha (Namibia), also Iona (Angola).

GYMNOPLEURINI
SURICATA 6 (2020) 279

Gymnopleurus humeralis
Klug, 1855

= Gymnopleurus humanus Macleay, 1821, var. azureus Ferreira, 1954c

Global: LC

J A S O N D J F M A M J

Type localities: ‘Tette’ [Tete, Central Mozambique]; G. humanus var.

azureus: ‘Rodésia do Sul: Hillside’ [possibly Hillside in Harare, Zim-
babwe].
Taxonomy: Accepted polymorphic species; exoskeleton either iridescent
blue, green or cupreous.
Distribution: Southern African wooded savanna: E Botswana, N South
Africa, Eswatini, Zimbabwe, Mozambique, Malawi.
Locality data (mean ± SD, range): Altitude: 735 ± 368, 144–1 850 m;
annual rainfall: 601 ± 158, 297–1 154 mm; annual temperature (max.
+ min. /2): 20.9 ± 2.0, 13.4–25.9°C (N=115).
Habitat: No quantitative assessment; DBRU collection records suggest a
bias to fairly sandy soils: sand (8), sandy loam (12) in wooded savanna:
grassland/pasture (2), open shrub/woodland (16); not found on the
deep sands of the Kalahari; at Phalaborwa: more abundant in reserves
(163) than farmland (5).
Food types: At Phalaborwa: strong bias to dung of omnivores compared
to herbivores: pig (162), elephant (1), cattle (21); DBRU collection
records on dung of omnivores: human (1), baboon (1); monogastric
herbivores: elephant (5), rhinoceros (2), horse (1), donkey (1); and
ruminant herbivores: cattle (9).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Apr.); at Phalaborwa: much more abundant on a hot, dry day
(151) than on a warm sunny day after heavy rainfall (6) or a warm
cloudy day after light rainfall (27).
Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725), Southern African Bushveld (AT0717), Southern Miombo
Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), N
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 534 260 km2; although widespread, possi-
ble bias to woody savanna suggests clearance of woodland would have 2 mm
a negative influence on population density; unclear if the Phalaborwa
results were influenced by vegetation density or dung type availability;
currently widespread in reserves and farm rangeland, so assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on soil and vegetation associations; protect-
ed in various reserves including Kruger National Park (South Africa),
Lake Mutirikwe Game Reserve (Zimbabwe).

GYMNOPLEURINI
280 SURICATA 6 (2020)

Gymnopleurus ignitus
Klug, 1855

= Gymnopleurus smaragdinus Fahraeus, 1857

= Gymnopleurus ignitus Klug var. laeviuscula Kolbe, 1895
= Gymnopleurus ignitus Klug var. nigrocupreus Janssens, 1938a
Global: LC

J A S O N D J F M A M J

Type localities: G. ignitus: ‘Inhambane und Sena.....Tette’ [Inhambane,

Sena, Tete: Mozambique]; G. smaragdinus: ‘prope fluvium Limpopo’
[close to Limpopo River, southern Africa]; var. laeviuscula: ‘Tanga,
Pangani, Mombassa, Sansibar’ [Tanga, Pangani, Zanzibar: Tanzania;
Mombasa, Kenya]; G. nigrocupreus: ‘Kikwit (Congo belge: Kwango)’
[Bandundu Province, W Democratic Republic of the Congo (DRC)].
Taxonomy: Accepted polymorphic species; exoskeleton either iridescent
green or cupreous; difficult to separate from more coarsely punctate
examples of G. pumilus Reiche, 1850.
Distribution: Probably sandy soils in tropical savanna from S central to
E Africa: W Democratic Republic of the Congo (DRC), N Namib-
ia, N Botswana, Zimbabwe, Malawi, Mozambique, coastal Tanzania,
coastal Kenya; Angola record requires validation, but is consistent with
Democratic Republic of the Congo (DRC) record; may also occur in
Limpopo Province, N South Africa; occurrence in central Namibia
tentative (black square).
Locality data (mean ± SD, range): Altitude: 706 ± 400, 0–1 427 m; an-
nual rainfall: 652 ± 225, 252–1 177 mm; annual temperature (max. +
min. /2): 22.8 ± 1.9, 17.8–25.9°C (N=24).
Habitat: No quantitative assessment; limited DBRU collection records
biased to sandy soils: deep sand (3), sandy loam (2) entirely in open
woodland (5).
Food types: No quantitative assessment; limited DBRU collection records
on various dung types: human (1), elephant (1), rhinoceros (1), wart-
hog (1), donkey (1).
Temporal activity: Diurnal flight activity in the summer rainy season
2 mm
(Nov. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian Baikiaea Wood-
lands (AT0726), Zambezian and Mopane Woodlands (AT0725), Kala-
hari Acacia-Baikiaea Woodlands (AT0709), Southern African Bushveld
(AT0717).
Assessment rationale: EOO = 659 605 km2; as the AOO may be limit-
ed by occurrence of sandy soils and woodland, clearance of woodland
might be detrimental to population density; however, quantitative
support is required; occupies a large EOO; therefore, assessed as Least
Concern (LC) although somewhat Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, particularly sup-
port for specialist soil and vegetation associations, which would assist in
determining the AOO; protected in several national parks: Chobe (N
Botswana), Gorongosa (central Mozambique).

GYMNOPLEURINI
SURICATA 6 (2020) 281

Gymnopleurus imitator
Balthasar, 1963c

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Süd-Westafrika, Otjiwarongo-District’ [Namibia].

Taxonomy: Accepted species, currently known only by the holotype; de-
scribed as closely related to G. sericatus Erichson, 1843, and G. humanus
Macleay, 1821, which are now considered synonyms (see G. humanus
species account); further study required to determine taxon status but
probably a synonym of the widespread G. humanus.
Distribution: Only known from Otjiwarongo District, Namibia (grey
shaded area).
Locality data: No precise locality data.
Habitat: Not known.
Food types: Not known.
Temporal activity: Not known, presumably diurnal flight activity like
other Gymnopleurus species; no seasonal data.
Ecoregions Namibia: Locality inexact, probably at S edge Angolan
Mopane Woodlands (AT0702) / NE edge Kalahari Xeric Savanna
(AT1309).
Assessment rationale: EOO unknown; a study is required to determine if
this taxon is a valid species or a synonym of G. humanus with morpho-
logical characteristics representing only intraspecific variation; assessed
as Data Deficient (DD).
Conservation measures: No assessment of conservation status is possible
until taxonomic questions have been resolved; should its valid species
status be upheld, quantitative data on EOO, AOO and ecological asso-
ciations will be required to support an assessment.

GYMNOPLEURINI
282 SURICATA 6 (2020)

Gymnopleurus leei
(Fabricius, 1792)

= Gymnopleurus caelatus Wiedemann & Germar, 1821

= Gymnopleurus bicolor Castelnau, 1840
= Gymnopleurus macleayi Castelnau, 1840
= Gymnopleurus liechtensteini Fahraeus, 1857
Global: LC
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type localities: As Scarabaeus leei: ‘Cap. Bon. Spei’; G. caelatus: ‘Prom.

Bon. Spei.’; G. bicolor: ‘Cap de Bonne-Espérance’; G. macleayi: ‘Cap’ [all
Cape of Good Hope, South Africa]; G. liechtensteini: ‘Natal’ [KwaZulu-
Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on higher or cooler, often drier parts of the sum-
mer rainfall Highveld; also, NE Upper Nama Karoo; with apparent
disjunct occurrence in the bimodal rainfall region along the S coast of
the Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 1 335 ± 408, 21–2 264 m;
annual rainfall: 514 ± 174, 227–845 mm; annual temperature (max. +
min. /2): 15.6 ± 1.5, 10.7–18.6°C (N=74).
Habitat: No quantitative assessment; DBRU collection records on sand
(8), sandy loam (8), sandy clay loam (5) in grassland/pasture (10),
scrub (1), open woodland (10) and old crop fields (3).
Food types: No quantitative assessment; DBRU collection records on
dung: human/cattle mix (1), horse (1), cattle (30).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: Centred on drier vegetation units in the cool
5 mm
Mesic Highveld Grassland (Gm); also moist Sub-Escarpment Grass-
land (Gs), Drakenberg Grassland (Gd) (Grassland Biome); moister NE
parts of Upper Karoo (Nama Karoo Biome) and E vegetation units of
Sandstone Fynbos (FFs) (Fynbos Biome).
Assessment rationale: EOO = 256 000 km2; other than the NE Nama
Karoo, the distribution coincides with vegetation units that are vulner-
able or endangered; transformation 33–67% (Gm); records from old
crop fields suggest some resilience to habitat transformation; however,
at Viljoenskroon only sampled as a rarity in natural grassland (2), not
in a crop field or improved pastures; assessed as Least Concern (LC)
due to large range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; at least 80%
of the range coincides with highly modified regions; therefore, quan-
titative data on effects of pasture improvement and other forms of
transformation would be useful; only very small parts of the range are
protected, e.g. Suikerbosrand and Abe Bailey nature reserves.

GYMNOPLEURINI
SURICATA 6 (2020) 283

Gymnopleurus pumilus
Reiche, 1850 (but see Taxonomy)

= Gymnopleurus gibbosus Roth, 1851

= Gymnopleurus virens Erichson subsp. sternalis Müller, 1942
Global: LC (see IUCN Red List – LC)*

J A S O N D J F M A M J

Type localities: G. pumilus: ‘Abyssinie’ [Ethiopia]; G. gibbosus: ‘abyssinien’

[Ethiopia]; subsp. sternalis: ‘Aethiopia meridionali’ [southern Ethiopia].
Taxonomy: This species is often cited as G. virens Erichson, 1843, but
does not match the type specimen; it more closely resembles species
described from Ethiopia that are currently synonymised in error with
G. virens (the oldest of these is G. pumilus); taxonomic revision is re-
quired to determine if southern African and Ethiopian populations rep-
resent two different species or a single widely distributed species with
a disjunction across several thousands of kilometres; the exoskeletons
of these populations are usually either iridescent yellow-green, green or
cupreous in southern Africa and blue or violet in NE Africa; the protho-
racic discs show an often ill-defined pattern of v- or u-shaped puncta-
tion and shiny apunctate patches; the punctation is usually moderately
strong, but does vary in strength; it may be close to the very strongly
patterned dense punctation of G. ignitis Klug, 1855, or to the weakly
patterned fine punctation of the true G. virens (see species account).
Distribution: Disjunct distribution in hot dry savanna possibly partly re-
lated to soil and climatic specialisation rather than entirely to collection
artefacts; NE Africa: Ethiopia; hiatus to southern Africa: N South Af-
rica, Mozambique, Zimbabwe, N Botswana, N Namibia, SW Angola;
record from Democratic Republic of the Congo (DRC) requires vali-
dation; record from Guinea presumably an error.
Locality data (mean ± SD, range): Altitude: 851 ± 435, 48–1 815 m;
annual rainfall: 550 ± 113, 277–820 mm; annual temperature (max. +
min. /2): 20.8 ± 1.7, 16.3–24.4°C (N=141). 2 mm
Habitat: No adequate quantitative assessment; in uMkhuze Game Reserve:
bias to finer-grained soils: sand (0.3 ), duplex soils (sand over clay) (1.4),
sandy clay loam (1.4), clay (4.8); partly supported by DBRU collection
records: sand (9), sandy loam (16), sandy clay loam (3), with a bias
to open woodland (20) as opposed to shrubland (4) or grassland (4).
Food types: At Phalaborwa: strong bias to dung of omnivores: pig (190), rather than herbivores: elephant (26), cattle (47);
DBRU collection records from a range of dung types: cattle (17), buffalo (1), wildebeest (1), waterbuck (1), impala (1),
elephant (4), rhinoceros (2), donkey (1).
Temporal activity: Diurnal flight activity in the summer rainy season (Oct. to Mar.); at Phalaborwa: much greater abundance
on a warm day soon after heavy rainfall (190) than on a warm cloudy day following light rainfall (26) or a hot dry day (47).
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Woodlands (AT0702), NE Kalahari Xeric Woodlands
(AT1309), NE edge Namibian Savanna Woodlands (AT1316), Zambezian and Mopane Woodlands (AT0725), Kalahari
Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 451 400 km2 (underestimated gross estimate); AOO would be smaller due to soil and vegetation
bias; data suggest two main southern African centres in N Namibia / SW Angola and N South Africa to NW and SE of the
largely unfavourable deep Kalahari sands; dry woodland savanna range maybe influenced by clearance of trees; in South Af-
rica, often extensive habitat transformation within its range (SVl: 0–58%, SVcb: 2–49%, SVmp: 0–20%); however, assessed
as Least Concern (LC) owing to wide range and protection in reserves; *see IUCN Red List entry under G. virens Erichson,
1843, which includes data for this species.
Conservation measures: Taxonomic revision is required to correct errors in nomenclature; assessment of conservation status
would also be improved by quantitative data on habitat associations; protected in Kruger National Park and uMkhuze Game
Reserve (South Africa).

GYMNOPLEURINI
284 SURICATA 6 (2020)

Gymnopleurus reichei
Waterhouse, 1890

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Abyssinia’ [Ethiopia].

Taxonomy: Accepted species, but further investigation of the taxonomic
status of the southern population would be useful.
Distribution: Dry highlands of NE Africa (N Kenya, Ethiopia) and dry
lowlands of NE South Africa; South African population possibly an
outlier rather than S limits of a continuous distribution; regional co-
lour variation with known NE individuals violet and S populations
cupreous.
Locality data (mean ± SD, range): Altitude: 256 ± 22, 237–281 m; an-
nual rainfall: 521 ± 27, 497–559 mm; annual temperature (max. +
min. /2): 22.9 ± 0.3, 22.5–23.1°C (N=4).
Habitat: Not recorded for South Africa.
Food types: No quantitative assessment; DBRU collection records from
dung of baboon (1), elephant (1), zebra (1), waterbuck (1).
Temporal activity: Diurnal flight activity in the summer rainy season
(Mar.).
Bioregions South Africa: Centred on Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 253 700 km2 (assumes unsupported con-
tinuous distribution from S to E Africa); widespread, but with probable
disjunct distribution; known AOO in NE Africa and NE South Africa
much smaller; known distribution in South Africa very restricted, but
entirely confined within a national park; therefore, assessed as Least
Concern (LC) although somewhat Data Deficient (DD).
Conservation measures: Assuming that a review of the taxonomy val-
idates the identity of the South African population as G. reichei, a
quantitative survey is required to determine its full EOO, AOO and
2 mm ecological associations in southern Africa; protected in Kruger National
Park.

GYMNOPLEURINI
SURICATA 6 (2020) 285

Gymnopleurus thelwalli
Waterhouse, 1890

No synonyms
Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type locality: ‘Lake Nyassa’ [environs of Lake Malawi].

Taxonomy: Accepted species; morphologically distant from other south-
ern African taxa.
Distribution: Validated records from Malawi, South Africa; also reported
from Zimbabwe, Tanzania; known only as a rarity from scattered sa-
vanna localities.
Locality data (mean ± SD, range): Altitude: 617 ± 416, 301–1 382 m;
annual rainfall: 661 ± 51, 553–700 mm; annual temperature (max. +
min. /2): 21.5 ± 2.4, 17.8–25.9°C (N=7).
Habitat: Not known.
Food types: Not known.
Temporal activity: Diurnal flight activity in the summer rainy season
(Dec., Feb.).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome).
Assessment rationale: EOO = 555 105 km2 (probably underestimated);
widespread savanna range, but known by only a few museum spec-
imens, possibly because of specialisation; AOO may or may not be
much smaller than EOO; very poorly known both biogeographically
and ecologically; assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, quantitative data
are required to determine if the AOO would be influenced by ecological
specialisation; an ideal focal area would be Skukuza where it has been
recorded with greatest frequency in South Africa; this would permit
predictions of AOO across its widespread recorded EOO; protected in 2 mm
the S Kruger National Park (South Africa).

GYMNOPLEURINI
286 SURICATA 6 (2020)

Gymnopleurus virens
Erichson, 1843

(see Taxonomy section – all synonyms are errors)

= Gymnopleurus pumilus Reiche, 1850
= Gymnopleurus gibbosus Roth, 1851
= Gymnopleurus vanderkelleni van Lansberge, 1886
= Gymnopleurus virens Erichson subsp. sternalis Müller, 1942
Global: LC (see IUCN Red List – LC)*

J A S O N D J F M A M J

Type localities: G. virens: ‘Angola’; G. pumilus: ‘Abyssinie’ [Ethiopia];

G. gibbosus: ‘abyssinien’ [Ethiopia]; as ‘G. van der Kelleni’: ‘West Africa’
[probably Angola]; subsp. sternalis: ‘Aethiopia meridionali’ [southern
Ethiopia].
Taxonomy: Accepted species, but revision is required to correct errors;
this mostly iridescent green species, trending to blue in moister and/
or cooler situations, has often been confused (e.g. Ferreira 1972) with
an E to NE African relative, G. sericeifrons Fairmaire, 1887, described
from ‘Makdischu’ [Mogadishu, Somalia]; however, it most strongly re-
sembles the finer-punctate type of G. virens Erichson, 1843, described
from Angola; all of the cited synonyms are considered to be errors; the
Ethiopian types most strongly resemble a more coarsely punctate south-
ern African taxon, which is the species usually cited as G. virens (but see
account for that species under the name, G. pumilus); G. vanderkelleni
is a misidentified synonym of G. aenescens Wiedemann, 1821, or a
close relative of that species.
Distribution: Possibly disjunct pattern in moist southern African upland
and coastal plain savanna; also marginal in warmer grassland: validated
for N South Africa, Mozambique coast, E Botswana, E Zimbabwe, SW
Angola (Bié Plateau).
Locality data (mean ± SD, range): Average altitude: 1 089 ± 497,
5–2 274 m; average annual rainfall: 739 ± 141, 383–1 142 mm; av-
erage annual temperature (max. + min. /2): 18.5 ± 2.6, 10.6–25.2°C
2 mm (N=114).
Habitat: On Gauteng bushveld: extreme association with finer-grained
soils: sandy clay loam (1 233), deep sand (1), and appreciable bias to
open woodland (838) as opposed to grassland (239) or shaded thickets
(156); supported by DBRU collection records from sand (3), sandy
loam (10), sandy clay loam (3) in grassland/pasture (4), shrubland (4),
open woodland (10), forest (1).
Food types: No quantitative assessment; DBRU collection records presumably biased by more commonly sampled dung types:
human (2), human/cattle mix (2), elephant (3), cattle (17).
Temporal activity: Diurnal flight activity, primarily in the summer rainy season (Oct. to Mar.); on Gauteng bushveld: greatest
activity early in summer rainy season continuing to Mar., declining to low levels in Apr., May, ceasing in June and commenc-
ing again in Sept.
Ecoregions Zimbabwe, Mozambique: Southern Miombo Woodlands (AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Moist S Central Bushveld (SVcb), W and S Lowveld (SVl) (Savanna Biome) along Waterberg and
edge of Highveld; warmer N Mesic Highveld Grassland (Gm) and N Sub-Escarpment Grassland (Gs) (Grassland Biome); N
Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 267 770 km2 (probably underestimated); AOO would be smaller, reflecting soil and vegetation
bias; probably influenced by woodland clearance, but also found in warmer moist grasslands; widespread in farmland and
protected in various nature reserves, assessed as Least Concern (LC); *IUCN Red List entry also includes data for G. pumilus
Reiche, 1850.
Conservation measures: Assessment of conservation status would be improved by quantitative data on food type associations;
protected in Rustenburg and Leeuwfontein nature reserves (South Africa).

GYMNOPLEURINI
288 SURICATA 6 (2020)

TRIBE ONITICELLINI
Kolbe, 1905

Subtribe Drepanocerina van Lansberge, 1875a: As Drepanocérides; Type genus:

Drepanocerus Kirby, 1828.
Subtribe Oniticellina Kolbe, 1905: As Oniticellini; Type genus: Oniticellus De-
jean, 1821.
Subtribe Helictopleurina Janssens, 1946: As Helictopleurides; Type genus: He-
lictopleurus d’Orbigny, 1915.
Subtribe Liatongina Philips, 2016: Type genus: Liatongus Reitter, 1892.
Subtribe Attavicina Philips, 2016: Type genus: Attavicinus Philips & Bell, 2008.

As regards citation of subtribal names in older format, Drepanocerina has precedence over that of Oniti-
cellina. However, at tribal level the name Oniticellini should be maintained as it has been in prevailing
long-term usage (Smith 2006). According to Philips (2016), subtribal divisions (van Lansberge 1875a;
Kolbe 1905; Janssens 1946) are partially valid owing to monophyly in the Drepanocerina and Madagascar
endemic, Helictopleurina. They are partially invalid as regards the Oniticellina, which are paraphyletic.
Therefore, Philips (2016) proposes two new subtribes, Attavicina and Liatongina, of which only Liatongina
are represented in southern Africa by two species.

Phylogenetic analysis of molecular data for selected taxa suggests monophyly at tribal level (Monaghan et
al. 2007) although different analyses of morphological data, together with those for Onthophagini, support
either monophyly or polyphyly (Philips 2016). The tribe currently comprises 25 or 26 genera depending on
the authority followed. A total of 18 are regional endemics: Afrotropical (10), Oriental (3), Palaearctic (1),
Neotropical (1), Jamaica (1) or Madagascar (2). The remainder are shared between regions: (A) Afrotropi-
cal, Oriental (5: Eodrepanus Barbero, Palestrini & Roggero, 2009; Ixodina Roth, 1851; Oniticellus Dejean,
1821; Tibiodrepanus Krikken, 2009; Tiniocellus Péringuey, 1901); (B) Afrotropical, Oriental Palaearctic (1:
Drepanocerus Kirby, 1828); (C) Afrotropical, Oriental, Palaearctic, Cuba (1: Euoniticellus Janssens, 1953);
(D) Afrotropical, Oriental, Palaearctic, Nearctic (1: but polyphyletic; Liatongus Reitter, 1892).

A total of 14 genera and 26 named species are represented in South Africa, Botswana and Namibia (summa-
ry below). They are mostly assessed as Least Concern (LC) owing to mostly very wide distributions (EOO
> 500 000 km2 in 20 out of 26 species), often with high abundance. A few are assessed as Data Deficient
(DD). A few species (6) are associated with monogastric herbivore dung, but also show widespread ranges.

Subtribe Drepanocerina:
(1) Afrodrepanus Krikken, 2009: A single species endemic to SE South African forest and shaded vegetation.
(2) Cyptochirus Lesne, 1900: A single species with a disjunct distribution in upland grassland from SE
to E Africa; patterned variety in moist savanna requires further examination.
(3) Drepanocerus Kirby, 1828: Two species in grassland to savanna; distributed from S to E Africa.
SURICATA 6 (2020) 289

(4) Eodrepanus Barbero, Palestrini & Roggero, 2009: Three species in dry to moist savanna and grass-
lands from SE to E Africa.
(5) Epidrepanus Roggero, Barbero & Palestrini, 2015: Probably a single species in South Africa, but
possibly two species with combined distributions from SE to E to W Africa.
(6) Ixodina Roth, 1851: A single poorly known species distributed from SE to E Africa.
(7) Latodrepanus Krikken, 2009: A single species in sandy savannas from S to E to W Africa.
(8) Paraixodina Roggero, Palestrini & Barbero, 2015: Two species of monogastic herbivore dung dis-
tributed from SE to E African savannas.
(9) Tibiodrepanus Krikken, 2009: A single species of moist SE African grasslands distributed from S to
E to W Africa.

Subtribe Onticellina:
(10) Euoniticellus Janssens, 1953: Five species; two from monogastric herbivore dung distributed from S
to E or S to E to W Africa; three other widespread species centred in cool South African grasslands,
moist grasslands from SE to E Africa, or various non-forest regions from S to E to W Africa.
(11) Oniticellus Dejean, 1821: Four widespread species; one on monogastric herbivore dung distributed
from S to E Africa; two species centred on dry or moist savannas from S to E to W Africa; one spe-
cies centred mostly on cool upland in SE and E central Africa.
(12) Tiniocellus Péringuey, 1901: Two species; one endemic to SE savannas; one widespread in savannas
from S to E Africa.
(13) Tragiscus Klug, 1855: A single species on monogastric herbivore dung in dry to moist savannas;
widespread in savannas from S to E Africa.

Subtribe Liatongina:
(14) Liatongus Reitter, 1892: A single widespread species, primarily centred in moist grasslands from SE
to E Africa; one unidentified forest species in SE South Africa (no species account).

ONITICELLINI
290 SURICATA 6 (2020)

Genus Afrodrepanus Krikken, 2009

Type species and designation: Drepanocerus impressicollis Boheman, 1857, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Krikken (2009), morphological phylogeny Roggero et
al. (2015).

Afrodrepanus Krikken, 2009, comprises only two species endemic to the Afrotropical region. One species shows a tropical
distribution in E and W Africa. The second species has a validated distribution in shaded savanna (thicket) and forest
vegetation of NE South Africa.

ONITICELLINI
SURICATA 6 (2020) 291

Afrodrepanus impressicollis
(Fahraeus, 1857)

= Drepanocerus natalensis Harold, 1859

Global: LC

J A S O N D J F M A M J

Type localities: As Drepanocerus impressicollis: ‘Caffraria meridionali’ [S

southern Africa]; D. natalensis: ‘Port Natal’ [Durban, KwaZulu-Natal,
South Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina.
Distribution: Mostly lower-altitude, moist, shaded vegetation; NE coast
and NE escarpment, South Africa; SE coast Mozambique; citations
from Zimbabwe, Tanzania and Kenya require validation in comparison
with the tropical A. marshalli (Boucomont, 1921).
Locality data (mean ± SD, range): Average altitude: 595 ± 610,
0–1 746 m; average annual rainfall: 875 ± 123, 656–1 094 mm; av-
erage annual temperature (max. + min. /2): 19.5 ± 2.6, 14.0–22.4°C
(N=32).
Habitat: On sandy loam in Ithala Game Reserve: abundant in dense
low-altitude woodland (Ngubu 714 m: 192; Dakaneni 511 m: 621;
Combretum woodland 690 m: 228), becoming uncommon at higher
altitude (Ntshondwe Forest 975 m: 15) and absent from mid-altitude
grassland at 945 m (0); in Wolkberg: present, but uncommon in forest
at 901 m (2.65 per pig dung sample); absent from forest and grassland
at 1 355–1 397 m (0); on deep sand at Richards Bay: bias to natural
dune forest (14.1 per sample) compared to post-mining vegetation
succession: grassland < 1–6 yr (0.0), dense/closed canopy woodland
7–21 yr (0.5), open canopy woodland 21–33 yr (0.0); in uMkhuze
Game Reserve: possible bias to finer-grained soils: sand (0.0), sand on
clay (0.4), sandy clay loam (0.4), clay (0.5).
Food types: In Ithala Game Reserve: very strong bias to omnivore dung
(pig: 890) compared to monogastric (horse: 47) and ruminant herbi-
vore dung (cattle: 119).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Sept. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Indian Ocean Coastal Belt Biome (CB); dense
1 mm
bushveld and thicket at SE and W edges, Lowveld (SVl) (Savanna
Biome); often in patches of Northern Mistbelt Forest (FOz 4) and
Northern Coastal Forest (FOz 7).
Assessment rationale: EOO = 27 860 km2; AOO would be much smaller
according to distribution of dense vegetation offering shade across a
range that is subject to extensive habitat transformation (0–58%, SVl);
however, owing to size of range and protection offered in various re-
serves, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by stronger quantitative data on soil associations and a survey
of dense vegetation patches potentially occupied by A. impressicollis;
protected in Ithala, uMkhuze and Hluhluwe–iMfolozi game reserves
(South Africa), Maputo Elephant Reserve (Mozambique).

ONITICELLINI
292 SURICATA 6 (2020)

Genus Cyptochirus Lesne, 1900

Type species and designation: Onitis ambigua Kirby, 1828, by monotypy.
Synonyms: = Drepanochirus Péringuey, 1901: Type species: Onitis ambigua Kirby, 1828, by mono-
typy.
Last review: Entire genus reviewed by Simonis and Zunino (1980); morphological phylogeny by
Roggero et al. (2015).

Cyptochirus Lesne, 1900, was revalidated from synonymy with Drepanocerus Kirby, 1828, by Simonis and Zunino (1980).
It currently comprises four species endemic to moist savanna and upland grassland of the Afrotropical region where they
construct a brood chamber immediately under droppings (Cambefort 1981).

Morphological phylogeny (Roggero et al. 2015) supports the validity of the four very closely related species, but cited
biogeographical overlap between species in upland centres in southern Africa, E Africa, E central Africa, NE Africa and W
Africa may require further investigation; particularly the claimed occurrence of a second species in South Africa that was
described from Ethiopia.

As ranges in body size overlap between all four Cyptochirus species (Simonis & Zunino 1980), further investigation should
compare specimens of similar body size, e.g. 10 mm long, to ensure claimed specific differences are not related to morpho-
logical variation with body size. A molecular phylogeny would also be useful. Here, only one species is considered to occur
in upland grassland in South Africa although a light and darker brown patterned ‘variety’ has been recorded primarily in
monogastric herbivore dung in lowland savanna of both South Africa and Zimbabwe.

ONITICELLINI
SURICATA 6 (2020) 293

Cyptochirus ambiguus
(Kirby, 1828)

= Oniticellus impressus Castelnau, 1840

Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Onitis ambigua: ‘Promontorium Bonae Spei’; O. impres-

sus: ‘Cap de Bonne-Espérance’ [both Cape of Good Hope, South Africa].
Taxonomy: Accepted species in the subfamily Drepanocerina; limited sex-
ual dimorphism (legs); some individuals from NE KwaZulu-Natal and
moist Zambezi Valley edge show a dark brown pattern on light brown
elytra; taxonomic status of patterned individuals requires assessment;
claimed occurrence of C. trogiformis (Roth, 1851), in South Africa also ♂
requires validation considering it was described from Ethiopia.
Distribution: Moist, warmer parts of Highveld and SE coast of South
Africa (red squares; blue square = outlier): South Africa, Eswatini; plus
claimed disjunct pattern of occurrence in E/NE African highlands:
Tanzania Kenya; further citations from Democratic Republic of the
Congo (DRC), Uganda, Ethiopia suggest sympatry with other Cyp-
tochirus species, but require validation; patterned individuals: South
Africa, Zimbabwe lowland savanna (black squares).
Locality data (mean ± SD, range): Unpatterned: Altitude: 1 046 ± 586,
21–2 286 m; annual rainfall: 775 ± 137, 430–1 047 mm; annual tempera-
ture (max. + min. /2): 16.9 ± 2.4, 10.8–22.6°C (N=112); Patterned: Al-
titude: 328 ± 277, 55–879 m; annual rainfall: 762 ± 107, 649–904 mm;
annual temperature (max. + min. /2): 22.1 ± 2.2, 18.4–25.9°C (N=10).
Habitat: No adequate quantitative assessment; at Foggy Valley unpat-
terned form abundant in natural Themeda grasslands (104), fallow crop
fields (72) and Kikuyu pasture (94); DBRU collection records from
sand (3), sandy loam (18), sandy clay loam (24), clay (6) primarily in
2 mm
grassland/pasture (46), rare in shrubland (1), open woodland (2), forest
(1); patterned form on sand (1), sandy loam (1), sandy clay loam (1) in
open woodland (3) or thicket (1).
Food types: No quantitative assessment; DBRU collection records for un- Patterned variety (black squares on map).
patterned form primarily from dung of cattle (53), also elephant (1);
patterned form primarily from monogastric dung: rhinoceros (4), ele-
phant (1), wildebeest (1).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to May).
Bioregions South Africa: Centred on E Mesic Highveld Grassland (Gm),
Sub-Escarpment Grassland (Gs) (Grassland Biome), transformed habitats
on S coast in Fynbos Biome, marginal along coast in three other biomes; both
unpatterned and patterned forms in moist S Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 323 350 km2; even though S and E African
unpatterned populations may comprise one or two different species; both
are widespread, although not currently known from any highland reserve
in South Africa; frequently recorded from cattle dung in both farmed
and natural moist grasslands; patterned individuals may comprise a fur-
ther species with a much smaller AOO as they are apparently associated
with the dung of large monogastric herbivores that are primarily restrict-
ed to game reserves in moist savanna; assessed as Least Concern (LC).
Conservation measures: A taxonomic revision is required; should pat-
terned individuals deserve species status, a survey to determine their
full EOO, AOO and conservation status would be required; further- 2 mm
more, quantitative ecological data for all populations would improve
assessments; patterned form protected in Tembe Elephant Park and ♀
Hluhluwe–iMfolozi Game Reserve.
Unpatterned variety (red squares on map).

ONITICELLINI
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Genus Drepanocerus Kirby, 1828

Type species and designation: Drepanocerus kirbyi Kirby, 1828, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Krikken (2009); morphological phylogeny (Roggero et al.
2015: three Afrotropical species).

Membership of the genus, Drepanocerus Kirby, 1828, was reduced to only four species by the revision of Krikken (2009).
These comprise three Afrotropical and one Oriental species. Two Afrotropical species occur in the E savanna and upland
grasslands of southern Africa with ranges extending into the E tropics. The third species is centred on E Africa.

ONITICELLINI
SURICATA 6 (2020) 295

Drepanocerus kirbyi
Kirby, 1828

= Oniticellus furcifer Castelnau, 1840

Global: LC

J A S O N D J F M A M J

Type localities: As D. kirbii: ‘Promontorium Bonae Spei’; O. furcifer:

‘Cap de Bonne-Espérance’ [both Cape of Good Hope, South Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina; sexual di-
morphism (head, prothoracic disc), characters varying in prominence
with body size.
Distribution: Savanna and grassland from S and S Central to NE Africa:
South Africa, N Namibia, N Botswana, Zimbabwe, Mozambique, Tan- ♂
zania, Kenya; also reported from Angola, Democratic Republic of the
Congo (DRC), Rwanda, Ethiopia.
Locality data (mean ± SD, range): Altitude: 883 ± 597 0–3 218 m; an-
nual rainfall: 733 ± 175, 306–1 659 mm; annual temperature (max. +
min. /2): 18.4 ± 2.9, 4.8–25.2°C (N=112).
Habitat: Soil generalist with a bias to woody vegetation in Gauteng: deep
sand (1 968), sandy clay loam (2 099); grassland (677), open woodland
(1 890), shaded thicket (1 500); however, DBRU collection records
from sand (10), sandy loam (23), sandy clay loam (10), clay (3) in
grassland/pasture (31), shrubland (6), open woodland (7), forest/thick-
et (2).
Food types: In Gauteng and Phalaborwa: strong bias to dung of rumi-
nant and monogastric herbivores, cattle (307/192), horse/elephant
(169/102), pig (24/17); DBRU collection records primarily from cattle
dung (48), also elephant (2), horse (1).
Temporal activity: Diurnal flight activity; in Gauteng: mostly between
10:00–16:00 with peak at midday (12:00–14:00) or mid-afternoon
(14:00–16:00); active throughout year with peak activity in summer
rainy season (Oct. to May), tailing off in June to low levels in the late
dry season; at Phalaborwa: frequency greater in warm sunny conditions 2 mm
soon after heavy rainfall (226) than in hot, dry (69) or warm cloudy
conditions (14) following light rainfall.
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian and Mopane
Woodlands (AT0725), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); E Mesic Highveld Grassland (Gm), Sub-Escarpment
Grassland (Gs) (Grassland Biome); Indian Ocean Coastal Belt Biome
(CB), Albany Thicket Biome (AT), transformed habitats in E coastal
part of Fynbos Biome.
Assessment rationale: EOO = 3 526 325 km2; widespread in both savan-
na and grassland despite a measured bias to woody habitats; occurs in
both farmland and reserves owing to a strong bias to association with
dung of both indigenous and domestic herbivores; assessed as Least
Concern (LC).
Conservation measures: None recommended; recorded in both farmland
and various game reserves including Hluhluwe–iMfolozi, uMkhuze,
Ithala (South Africa).

2 mm
♀

ONITICELLINI
296 SURICATA 6 (2020)

Drepanocerus patrizii
(Boucomont, 1923a)

= Drepanocerus arthuri Boucomont, 1936

Global: LC

J A S O N D J F M A M J

Type localities: As Cyptochirus patrizii: ‘N. Kenia’ [N Kenya]; D. arthu-

ri: ‘Afrique orientale anglaise: Embaki’ [Tanzania: Lake Embaki in
Ngorongoro Conservation Area].
Taxonomy: Accepted species in the subfamily, Drepanocerina; sexual
dimorphism (head), characters varying in prominence with body size.
Distribution: Disjunct occurrence in dry savanna and grasslands of S and
♂ E Africa separated by moister climate: South Africa, Kenya, Tanzania;
also reported from Namibia, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 129 ± 409, 93–1 759 m;
annual rainfall: 536 ± 189, 254–975 mm; annual temperature (max. +
min. /2): 18.0 ± 2.5, 13.9–23.4°C (N=41).
Habitat: In Gauteng bushveld: relatively generalist with slight bias to finer-
grained soils and more open vegetation: deep sand (333), sandy clay
loam (594); grassland (320), open woodland (482), shaded thickets
(111); largely supported by DBRU collection records: sand (1), sandy
loam (16), sandy clay loam (3), clay (2) in grassland/pasture (10), shru-
bland (6), open woodland (5).
Food types: In Gauteng and Phalaborwa: equal bias to herbivore dung
types: cattle (14/3), horse/elephant (14/3); absent from omnivore
dung: pig (0/0); DBRU collection records from dung of cattle (19),
horse (1), donkey (1).
Temporal activity: Diurnal flight activity; in Gauteng: mostly between
10:00 and 16:00 with a mid-summer peak at midday (12:00–14:00)
or early/late summer peak in mid-afternoon (14:00–16:00); recorded
throughout the year except late dry season (Aug., Sept.), but most
2 mm
abundant early to mid-rainy season (Oct. to Feb.), tailing off into late
summer and the dry season.
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVl),
Lowveld (SVl), Eastern Kalahari Bushveld (SVk) (Savanna Biome);
Dry Highveld Grassland (Gh) (Grassland Biome); E Upper Karoo
(NKu) (Nama Karoo Biome); Albany Thicket Biome (AT); margins of
two other bioregions.
Assessment rationale: EOO = 503 040 km2; widespread under drier
climate in both farmland and game reserves owing to fairly generalist
habitat associations and association with both ruminant and monogas-
tic herbivore dung; thus assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by a distributional survey to confirm the disjunct EOO; pro-
tected in Kruger National Park, Ithala Game Reserve (South Africa),
Amboseli and Marsabit game reserves (Kenya).

5 mm

ONITICELLINI
SURICATA 6 (2020) 297

Genus Eodrepanus Barbero,

Palestrini & Roggero, 2009a
Type species and designation: Drepanocerus parallelus Raffray, 1877, by original designation.
Synonyms: None.
Last review: Entire genus created and reviewed by Barbero et al. (2009a); morphological phylogeny
(Roggero et al. 2015).

Eodrepanus Barbero, Palestrini & Roggero, 2009a was created to accommodate nine species: eight of them extant, pri-
marily from the savannas and grasslands of the Afrotropical (5) and Oriental regions (3), one of them extinct, known as
a fossil from the Palaearctic region (England). All three of the species recorded in southern Africa are only known from N
Botswana and (primarily) E South Africa where two are found in E savanna and one in forest, savanna and grassland. All
three species also range into the E tropics.

ONITICELLINI
298 SURICATA 6 (2020)

Eodrepanus bechynei
(Janssens, 1953)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Drepanocerus bechynei: ‘Abyssinie: lac Daka (Type)’ [Ho-

lotype: Lake Daka, Ethiopia].
Taxonomy: Accepted species in the subfamily, Drepanocerina.
Distribution: Apparently disjunct, high rainfall, lowland and upland
tropical pattern along E seaboard from (1) NE coastal hills of South
Africa to uplands of NE Africa (South Africa, Mozambique, E Dem-
♂ ocratic Republic of the Congo (DRC), Rwanda, Burundi, Tanzania,
Zambia, Kenya, Uganda, Ethiopia) and (2) lowland West Africa (Gam-
bia, Guinea (Conakry), Côte d’Ivoire, Ghana, Benin, Burkina Faso,
Niger, Nigeria, Cameroon); also cited from Senegal, Guinea (Bissau).
Locality data (mean ± SD, range): Altitude: 421 ± 556, 144–1 464 m;
annual rainfall: 1 021 ± 315, 826–1 659 mm; annual temperature
(max. + min. /2): 20.2 ± 1.5, 17.2–21.1°C (N=6).
Habitat: No quantitative assessment; limited DBRU collection records
from deep sand (1), sandy clay loam (1), clay (1) in pasture/grassland
(2).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (1), buffalo (1), rhinoceros (1).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Aug., Nov., Mar., Apr.).
Bioregions South Africa: Moist S Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 2 080 435 km2 (gross estimate); poorly
known ecologically; apparently restricted to a very small, disjunct,
range in southern Africa, but widespread in parts of the wet tropics;
known range in South Africa mostly protected; therefore assessed as
Least Concern (LC) although essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Hluhluwe–iMfolozi Game Reserve (South Africa).

2 mm

ONITICELLINI
SURICATA 6 (2020) 299

Eodrepanus fastiditus
(Péringuey, 1901)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Drepanocerus fastiditus; ‘Natal (Frere, Durban, Est-

court)’ [KwaZulu-Natal, South Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina; limited
sexual dimorphism (prothoracic disc, elytra).
Distribution: Moist coastal and upland areas along E seaboard from S
to NE Africa: South Africa, Zimbabwe, Mozambique, E Democratic
Republic of the Congo (DRC), Burundi, Kenya, Uganda, Ethiopia, ♀
Eritrea; also cited from Rwanda, Tanzania; record from Guinea (Bissau)
requires validation; southern African disjunctions probably a mix of
collection artefacts and unsuitable habitat or climate.
Locality data (mean ± SD, range): Altitude: 941 ± 511, 1–1 782 m; an-
nual rainfall: 759 ± 209, 240–1 659 mm; annual temperature (max. +
min. /2): 18.5 ± 2.3, 13.8–25.2°C (N=56).
Habitat: In Gauteng bushveld: bias to sandy clay loam (95) compared
to deep sand (24) and bias to open vegetation, grassland (69), open
woodland (46), shaded thicket (4); largely supported by DBRU collec-
tion records: bias to finer-grained soils, deep sand (4), sandy loam (4),
sandy clay loam (3), clay (2) in grassland/pasture (8), open woodland
(4), forest (1), also fallow crop fields (2).
Food types: At Phalaborwa: strong bias to monogastric herbivore dung
(elephant: 13) compared to ruminant herbivore (cattle: 2) and omni-
vore dung (0); DBRU collection records primarily from dung of cattle
(9), horse (2), elephant (1), human/cattle mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Aug. to Apr.); at Phalaborwa: primarily active in hot
sunny weather (13), few in warm sunny weather soon after heavy rain-
fall (2), absent in warm cloudy weather following light rainfall (0).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719). 2 mm
Bioregions South Africa: Biomes: primarily straddling edges of Indian
Ocean Coastal Belt Biome (CB) to Savanna (SV) and Savanna to
Grassland (G); Bioregions: Central Bushveld (SVcb), Lowveld (SVl),
Sub-Escarpment Savanna (SVs), Mesic Highveld Grassland (Gm),
Sub-Escarpment Grassland (Gs); marginal in three other bioregions.
Assessment rationale: EOO = 1 517 320 km2 (gross estimate); extremely
widespread in open vegetation along E seaboard of Africa so soil and
climatic bias would not be a threat; however, bias to monogastric her-
bivore dung would have implications for population density outside of
reserves; even so, assessed as Least Concen (LC).
Conservation measures: None recommended owing to widespread
occurrence; protected in Leeuwfontein and Tswaing nature reserves,
Hluhluwe–iMfolozi Game Reserve (South Africa).

ONITICELLINI
300 SURICATA 6 (2020)

Eodrepanus parallelus
(Raffray, 1877)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Drepanocerus? parallelus: ‘Abyssinie: Province de Tem-

biène’ [Tembien Province, Ethiopia].
Taxonomy: Accepted species in the subfamily, Drepanocerina.
Distribution: Apparent disjunct pattern, primarily in moist savanna along
E seaboard from uplands of NE Africa to NE lowlands of South Africa:
Eritrea, Ethiopia, Uganda, Kenya, Tanzania, E Democratic Republic of
the Congo, Rwanda, Burundi, Malawi, Zambia, Zimbabwe, Mozam-
♀
bique, Botswana, South Africa.
Locality data (mean ± SD, range): Altitude: 754 ± 381, 144–1 108 m;
annual rainfall: 755 ± 122, 589–904 mm; annual temperature (max. +
min. /2): 21.0 ± 1.8, 19.0–24.4°C (N=8).
Habitat: No quantitative assessment; limited DBRU collection records
from sand (2), sandy clay loam (1) in shrubland (1), woodland (1),
fallow crop field (1).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (2), buffalo (1), rhinoceros (1).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Aug., Jan., Apr.).
Ecoregions Botswana, Zimbabwe: Southern Miombo Woodlands
(AT0719), Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Moist S Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 797 475 km2 (gross estimate); poorly
known ecologically; patchy records from moist savanna of southern
Africa, but widespread in parts of the tropics; known range in South
Africa mostly protected; therefore, assessed as Least Concern (LC) al-
though essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by a quantitative data on ecological associations; protected
in Hluhluwe–iMfolozi Game Reserve (South Africa), Chobe National
Park (Botswana), Lake Mutirikwe Game Reserve (Zimbabwe).

2 mm

ONITICELLINI
SURICATA 6 (2020) 301

Genus Epidrepanus Roggero,

Barbero & Palestrini, 2015
Type species and designation: Oniticellus caelatus Gerstaecker, 1871, by original designation.
Synonyms: None.
Last review: Genus created and reviewed with a morphological phylogeny by Roggero et al. (2015).

Epidrepanus Roggero, Barbero & Palestrini, 2015 was created to accommodate three Afrotropical species previously in-
cluded in the genus, Latodrepanus Krikken, 2009. Two species have been recorded in South African uplands. However,
these were described from the same type locality in Tanzania and there seems to be much confusion in their identification.

ONITICELLINI
302 SURICATA 6 (2020)

Epidrepanus caelatus
(Gerstaecker, 1871)

Epidrepanus pulvinarius
(Balthasar, 1963c)

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: As Oniticellus caelatus: ‘Aruscha’; as Drepanocerus pulvi-

narius: ‘Ost-Afrika, Tanganjika, Aruscha’ [both Arusha, N Tanzania].
Taxonomy: Both accepted species in the subfamily, Drepanocerina; how-
♀ ever, each described from the same type locality from only a single un-
sexed (E. caelatus) or a single male specimen (E. pulvinarius); material
from South African museums variously identified as E. caelatus or
E. pulvinarius, but seems confused.
Distribution: In South Africa: Epidrepanus species are centred along the
Waterberg, N edge of Highveld Grassland, and edge of NE escarpment;
they are primarily recorded in moist open woodland on E coastal hills
and in Highveld valleys; also widespread, but rare, across Highveld Grass-
land; on the map Epidrepanus individuals are identified as E. caelatus (red
squares), E. pulvinarius (black squares), or are unidentified to species
level (grey squares); Roggero et al. (2017) cite E. pulvinarius only from
the SE coast and E. caelatus primarily from the Highveld and NE coast;
across Africa E. pulvinarius is cited from Kenya, Tanzania and South
Africa (Roggero et al. 2017); E. caelatus is cited from Democratic Re-
public of the Congo, Tanzania, Kenya, Ethiopia, Guinea (or in all seven
regions defined from southern to E to W Africa) (Roggero et al. 2015).
Locality data (mean ± SD, range): Altitude: 1 199 ± 408, 273–1 824 m;
annual rainfall: 786 ± 117, 576–1 157 mm; annual temperature (max.
+ min. /2): 17.2 ± 1.8, 14.6–20.5°C (N=24).
Habitat: No adequate quantitative assessment; on finer-grained soils at
Suikerbosrand Nature Reserve: only in open woodland (26 at 1 665 m)
not Highveld Grassland (0 at 1 662 m) or montane grassland (0 at
1 910 m); on finer-grained soils at Ithala Game Reserve: primarily
in shaded woodland at Dakaneni (602 at 472 m), dense shrubland
at Ngubu (458 at 706 m) and open Combretum woodland (235 at
1 mm 711 m), rare in Highveld Grassland (1 at 1 325 m).
Food types: In Ithala Game Reserve: abundant on monogastric herbivore,
ruminant herbivore and omnivore dung (horse: 546; cattle: 349; pig: 400).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Oct. to Feb.).
Bioregions South Africa: Primarily straddling edges of Savanna and
Grassland biomes; Central Bushveld (SVcb), Lowveld (SVl) (Savanna
Biome); N and E edges: both Mesic Highveld Grassland (Gm) and
Sub-Escarpment Grassland (Gs) (Grassland Biome).
Assessment rationale: EOO = 160 840 km2 (range in South Africa); tax-
onomy unclear, possibly confused; ecological data related to E. pulvi-
narius at Ithala and Epidrepanus sp. at Suikerbosrand; both data sets sug-
gest population density would be influenced by clearance of woodland;
assessed as Data Deficient (DD) because of taxonomic uncertainty.
Conservation measures: It seems that further taxonomic revision is re-
quired to adequately assess the conservation status of Epidrepanus species
in South Africa; however, ecological data suggest preservation of moist
woodland is a requirement for conservation of the species; protected in
Suikerbosrand Nature Reserve and Ithala Game Reserve (see above).

ONITICELLINI
SURICATA 6 (2020) 303

Genus Euoniticellus Janssens, 1953

Type species and designation: Scarabaeus fulvus Goeze, 1777, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Janssens (1953); new species added by Cambefort and
Mauchamps (1980), Cambefort (1983), Biswas and Chatterjee (1985, cited as Oniti-
cellus), Masumoto (1987) and by Cambefort (1996a) with a phylogeny of all but the
Biswas and Chatterjee species.

Euoniticellus Janssens, 1953, separates groups of larger-bodied tunnelling species and smaller-bodied species of unknown
habits from the genus Oniticellus Dejean, 1821, which now contains only species showing pad-dwelling, endocoprid
breeding habits. The Biswas and Chatterjee species from N India were not included in the phylogeny of Cambefort
(1996a). Two of these N Indian species are stated to be close to E. pallipes (Fabricius, 1781) and E. pallens (Olivier, 1789),
both of which have also been recorded in N India.

Euoniticellus shows a distribution in Afro-Eurasia with one species isolated on the Caribbean island of Cuba. It comprises a
total of 19 or 21 species represented in the Afrotropical (14), Oriental (1 +?2) and Palaearctic regions (2), with one further
species shared between the Afrotropical, W Oriental and S Palaearctic regions.

Phylogeny (Cambefort, 1996a) splits the genus into eight more basally derived, smaller-bodied species and 11 more ter-
minally derived, larger-bodied species, which include E. pallens and E. pallipes. Further study is required to determine the
evolutionary, ecological and taxonomic implications of this topology, if any.

In South Africa, Botswana and Namibia, Euoniticellus is represented by four larger-bodied and one smaller-bodied species.

Smaller-bodied: One E savanna species associated with elephant dung with a range extending into the E tropics.

Larger-bodied: Four species; one with a savanna distribution and widespead into tropical savanna on elephant dung;
three species more frequently recorded on dung of large ruminant herbivores; one widespread thoughout southern Africa
and all savanna S of the Sahara; one centred in the S Cape, moist SE savanna and moist N Highveld Grassland with addi-
tional occurrence range on E tropical highlands; the third centred on Highveld Grassland, Upper Karoo and the S Cape
of South Africa.

The three most terminally derived species showing ruminant dung bias are assessed as Least Concern (LC) as they are
widespread and abundant. Those more basally derived species showing elephant dung associations are assessed as Data
Deficient (DD).

ONITICELLINI
304 SURICATA 6 (2020)

Euoniticellus africanus
(Harold, 1873)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Oniticellus africanus: ‘Cap bon. spei; Bloemfontein’

[South Africa: Cape of Good Hope; Free State].
Taxonomy: Accepted species; limited sexual dimorphism (head).
Distribution: Cool Highveld, Upper Karoo, and SE to S lowlands of
South Africa; also reported from Lesotho.
♂ Locality data (mean ± SD, range): Altitude: 1 218 ± 473, 8–2 700 m;
annual rainfall: 518 ± 201, 118–1 101 mm; annual temperature (max.
+ min. /2): 15.8 ± 1.6, 8.0–20.8°C (N=215).
Habitat: No adequate quantitative assessment; in Western Cape: strong
bias to transformed pasture habitats (Geelbek: 46, Groote Post: 532,
Oranjefontein: 180, Waylands: 454) as opposed to natural shrubland
(Geelbek: 1, Modderrivier: 6, Pampoenvlei (sandy loam): 8, Pampoen-
vlei (sand): 37); DBRU collection records suggest a possible bias to
finer-grained soils: sand (17), sandy loam (33), sandy clay loam (50),
clay (2) in open vegetation: grassland/pasture (57), scrub (8), open
woodland (6), fallow crop fields (4).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (136), buffalo (3), horse (2), zebra (2).
Temporal activity: Diurnal flight activity in spring, summer and autumn
across the winter, bimodal (spring, autumn peaks), late summer and
mid-summer rainfall regions (Aug. to May).
Bioregions South Africa: Dry Highveld Grassland (Gh), Mesic Highveld
Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland Biome);
Eastern Kalahari Bushveld (SVk) (Savanna Biome); Upper Nama
Karoo (NKu) (Nama Karoo Biome); Albany Thicket Biome (AT),
transformed habitats of Fynbos Biome in S of Western and Eastern
2 mm Cape; marginal in four other bioregions
Assessment rationale: EOO = 677 390 km2; few known records from
protected areas; however, widespread in farmland in South Africa where
it is readily attracted to the dung of farm livestock, especially that of
cattle; although transformation is extensive in the moister parts of its
range (Gh, 4–63%; Gm, 6–69%; Gs, 2–50%); not currently consid-
ered threatened; therefore, assessed as Least Concern (LC).
Conservation measures: Quantitative data on ecological associations
would improve the assessment of conservation status, particularly,
effects of pasture improvement; however, records from transformed
habitats in the Fynbos Biome and from fallow crop fields suggest some
tolerance of habitat modification; protected in Addo Elephant National
Park.

♀
2 mm

ONITICELLINI
SURICATA 6 (2020) 305

Euoniticellus intermedius
(Reiche, 1849)
= Oniticellus pallens Olivier, 1789, sensu Castelnau, 1840
= Oniticellus clavatus Roth, 1851
= Oniticellus speciosus Costa, 1853
= Oniticellus nasicornis Reiche, 1849, sensu Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As Oniticellus intermedius: ‘Cap de Bonne-Espérance,

d’Abyssinie et d’Algérie’ [Cape of Good Hope, South Africa; Ethiopia;
Algeria]; O. pallens sensu Castelnau: ‘Sénégal’ [Senegal]; O. clavatus:
Not stated; O. speciosus ‘Calabrie’ [Calabria, S Italy; synonymy requires ♂
validation]; O. nasicornis sensu Péringuey: ‘Cape Colony (Worcester,
Knysna, Queen’s Town, Kimberley, Graham’s Town, Uitenhage, Albert),
Natal (Durban, Maritzburg), Transvaal (Potchefstroom, Klerksdorp),
Lower Rhodesia (Buluwayo, Enkeldoorn)’ [South Africa, Zimbabwe].
Taxonomy: Accepted species; clear sexual dimorphism (head, prothoracic
disc), prominence of characters varies with body size.
Distribution: Distributed throughout savannas of S, E, and W Africa with
a bias to drier climate; cited from at least 25 sub-Saharan countries; also
reported from N Africa (Algeria), Arabia, and as a rarity in S Italy (re-
quires validation); introduced into Australia and USA; dispersed into
central America (Mexico, Nicaragua, Costa Rica).
Locality data (mean ± SD, range): Altitude: 967 ± 459, 0–2 286 m; an-
nual rainfall: 507 ± 248, 34–1 659 mm; annual temperature (max. +
min. /2): 18.7 ± 2.3, 10.6–25.2°C (N=580).
Habitat: In Gauteng bushveld: no particular soil bias, deep sand (3 676),
sandy clay loam (4 464), but much more abundant in grassland (3 429)
or open woodland (4 165) as opposed to shaded thickets (546); in West-
ern Cape: strong bias to transformed pasture habitats (Geelbek: 34, 2 mm
Groote Post: 656, Oranjefontein: 364, Waylands: 936) as opposed to
natural shrubland (Geelbek: 3, Modderrivier: 29, Pampoenvlei (sandy
loam): 21, Pampoenvlei (sand): 106); supported by DBRU collection
records from sand (72), sandy loam (56), sandy clay loam (75), clay
(13) in grassland/pasture (98), scrub/shrubland (31), open woodland
(55), forest/thicket (2); also fallow crop fields (14).
Food types: In Gauteng bushveld: strong bias to dung of pad-dropping
ruminant herbivores: cattle (306), horse (46), pig (3), carrion (0);
similar bias at Phalaborwa: cattle (1 049), elephant (115), pig (23);
DBRU collection records primarily from dung of cattle (255); also,
buffalo (13), elephant (4), horse (4), rhinoceros (3), donkey (1), zebra
(1), human/cattle or buffalo mixed bait (6), baboon (1).
Temporal activity: Diurnal flight activity peaking at midday (12:00–
14:00); in both Gauteng bushveld (summer rainfall) and Western
Cape (winter rainfall): active throughout entire year showing greatest
abundance in summer, whether dry or wet; at Phalaborwa: frequency
much greater in warm sunny conditions soon after heavy rainfall (951)
than in warm cloudy conditions following light rainfall (30) or hot, dry
conditions (157).
Ecoregions Namibia, Botswana, Zimbabwe: Recorded from most ecore-
gions of Namibia and Botswana (see summary tables for ecoregions),
also Southern Miombo Woodlands (AT0719) in Zimbabwe.
Bioregions South Africa: Recorded from all biomes (Savanna (SV),
Grassland (G), Nama Karoo (NK), Succulent Karoo (SK), Indian ♀
2 mm

ONITICELLINI
306 SURICATA 6 (2020)

Euoniticellus intermedius (continued)

Ocean Coastal Belt (CB), Albany Thicket (AT), Fynbos (F)) and most
bioregions, but biased to warmer, drier areas; rare or absent from cooler
wetter parts of Highveld.
Assessment rationale: EOO = 9 060 360 km2 (underestimated sub-
Saharan range); open habitat generalist, tolerant of habitat modifica-
tion; exceptionally widespread throughout African savannas into the
Palaearctic region; widely recorded in reserves with further populations
resulting from introductions into the Americas and Australia; assessed
as Least Concern (LC).
Conservation measures: None recommended; adaptable with a bias to
dung pads of ruminant herbivores that are abundant in both farmed
areas and game reserves such as Kruger National Park (South Africa).

ONITICELLINI
SURICATA 6 (2020) 307

Euoniticellus kawanus
(Janssens, 1939b)

No synonyms
Global: DD (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Forêt de Kawa, Lac Albert’ [not traced; presumably E

Democratic Republic of the Congo (DRC)].
Taxonomy: Accepted species, but this savanna-centred species should be
validated by comparison with the forest-collected type.
Distribution: Widespread, but patchy occurrence in dry savanna due
to dung type specialisation; recorded from S to E to W Africa: South
Africa, Namibia, Botswana, Zimbabwe, Democratic Republic of the ♂
Congo (DRC), Kenya, Côte d’Ivoire.
Locality data (mean ± SD, range): Altitude: 968 ± 299, 274–1 110 m;
annual rainfall: 444 ± 154, 209–608 mm; annual temperature (max. +
min. /2): 20.6 ± 2.9, 15.7–23.6°C (N=8).
Habitat: No quantitative data; limited DBRU collection records from
sand (1) or sandy loam (3), only in open woodland (3).
Food types: No adequate quantitative data; in Botswana: recorded on
elephant (2) and pig dung (1); DBRU collection records from elephant
(4) and zebra (1) dung.
Temporal activity: Diurnal flight activity during the summer rainy season
(Nov. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian and Mopane
Woodlands (AT0725), Angolan Mopane Woodlands (AT0702), Na-
mibian Savanna Woodlands (AT1316); margins of one other ecoregion.
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 2 691 645 km2; widespread, but infre-
quently recorded in low numbers suggesting low population density;
most available records with ecological data (12) recorded from mono-
gastric herbivore dung; thus AOO would be influenced by the range
contraction experienced by these taxa all over Africa with most records
2 mm
from game reserves; ecological data too limited to clearly determine if
AOO is further influenced by any habitat bias; assessed as Data Defi-
cient (DD).
Conservation measures: Quantitative data on ecological associations is
required to assess conservation status; however, limited observations
suggest conservation status would be dependent on the continuing pro-
tection of large monogastric herbivores; protected in various national
parks and game reserves, including Kruger (South Africa), Hwange
(Zimbabwe), Chobe (Botswana), Etosha (Namibia), Meru (Kenya).

ONITICELLINI
308 SURICATA 6 (2020)

Euoniticellus triangulatus
(Harold, 1873)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Oniticellus triangulatus: ‘Africa australis’ [southern Af-

rica].
Taxonomy: Accepted species; limited sexual dimorphism (head).
Distribution: Bias to cool, moist grasslands and transformed pastures
from the S and SE coast plus moist Highveld of South Africa to uplands
of E Africa (E African DBRU records mostly above 1 400 m: n=28/32;
♂ with bias to pasture/grass: n=20/33): South Africa, Namibia, Zimbab­
we, Tanzania, Kenya, Eritrea; also reported from Angola, Democratic
Republic of the Congo (DRC), Mozambique; W African occurrence
requires validation: Guinea, Benin; southern African outliers (black
squares) possibly in shade.
Locality data (mean ± SD, range): Altitude: 887 ± 660, 0–3 218 m; an-
nual rainfall: 673 ± 206, 166–1 101 mm; annual temperature (max. +
min. /2): 16.9 ± 2.6, 4.8–24.4°C (N=178).
Habitat: No adequate quantitative assessment; in Western Cape: strong
bias to cool transformed pasture habitats as opposed to natural shrub­
land; cool Cape Peninsula (Kikuyu grass pasture on Bonne Attente:
4 478 vs shrubland of Cape of Good Hope Nature Reserve: 2); warmer
Darling Hills pasture (Groote Post: 1 229, Oranjefontein: 722, Way-
lands: 1 394) vs shrubland of adjacent coastline (Modderrivier: 29,
Pampoenvlei (sandy loam): 18, Pampoenvlei (sand): 184); DBRU col-
lection records from sand (10), sandy loam (26), sandy clay loam (27),
clay (3) with bias to grassland/pasture (53) rather than shrubland (4) or
open woodland (3); also fallow crop fields (6).
Food types: No quantitative assessment; DBRU collection records pri-
marily from dung of cattle (90); also buffalo (1), horse (3).
2 mm Temporal activity: Diurnal flight activity throughout entire year over
entire South African range; to the N: greatest frequency of DBRU re-
cords in summer rainy season (Oct. to May); similar to the SW in the
Western Cape winter rainfall region: greatest abundance in moist late
spring throughout the dry summer till moist autumn (Sept. to May).
Bioregions South Africa: Warmer parts of Grassland Biome: N and E
Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs);
Indian Ocean Coastal Belt Biome (CB); transformed habitats along
S coast in Albany Thicket (AT) and Fynbos Biomes; also margins of
Savanna and Nama Karoo Biomes.
Assessment rationale: EOO = 2 234 985 km2 (underestimated, also re-
corded in Eritrea); AOO would be somewhat smaller, fragmented, and
restricted to grassy highlands and cool dry (to the S) or wet coastal
regions (to the N); recorded from few protected areas; however, wide-
spread in farmland where it is readily attracted to cattle dung in natural
grassland, habitats transformed to pasture, even fallow crop fields; as-
sessed as Least Concern (LC).
Conservation measures: Quantitative data on ecological associations
would improve the assessment of conservation status; protected in
Bontebok National Park.

2 mm ♀

ONITICELLINI
SURICATA 6 (2020) 309

Euoniticellus zumpti
Janssens, 1953

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Hluhluewe, Zululand’ [presumably Hluhluwe Game Re-

serve, KwaZulu-Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Moist to dry lowland savanna from SE to E Africa: South
Africa, Mozambique, Kenya; also cited from Tanzania.
Locality data (mean ± SD, range): Altitude: 310 ± 314, 19–1 126 m;
annual rainfall: 778 ± 162, 539–1 136 mm; annual temperature (max. ♂
+ min. /2): 21.4 ± 1.9, 16.8–25.2°C (N=15).
Habitat: No quantitative assessment; observed on deep sand and finer-
grained soils; limited DBRU collection records from sand (1) in open
woodland (2).
Food types: At Phalaborwa: sampled to dung of elephant (5), pig (1), but
not cattle (0); DBRU collection records from dung of elephant (1),
rhinoceros (1), zebra (1).
Temporal activity: Diurnal flight activity primarily during the summer
rainy season (Oct. to Mar.).
Bioregions South Africa: Lowveld (SVl) (Savanna Biome); marginal in
two other bioregions.
Assessment rationale: EOO = 273 965 km2 (gross estimate); AOO would
presumably be smaller because of bias to dung of monogastric herbi-
vores; this association is reflected by the disjunct patchy distribution
primarily in game reserves into which the ranges of large indigenous
herbivores have contracted; as quantitative ecological data is limited;
the species is assessed as Data Deficient (DD) although it would prob-
ably qualify for Least Concern (LC) owing to widespread occurrence 2 mm
in various reserves.
Conservation measures: Quantitative data on habitat associations and a
survey to determine full EOO and AOO are required before an ade-
quate assessment may be made of conservation status; protected in var-
ious game reserves and national parks, including Hluhluwe–iMfolozi,
Ithala, Kruger (South Africa), Masai Mara (Kenya).

♀
2 mm

ONITICELLINI
310 SURICATA 6 (2020)

Genus Ixodina Roth, 1851

Type species and designation: Ixodina abyssinica Roth, 1851, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Krikken (2009).

Ixodina Roth, 1851, was revalidated from synonymy with Drepanocerus Kirby, 1828, by Krikken (2009). It now comprises
five species in moist savannas of the Afrotropical region following the recent transfer of three species to the new genus,
Paraixodina Roggero, Barbero & Palestrini, 2015. Roggero et al. (2015) note that there are no significant differences be-
tween the genitalia and epipharynx of four Ixodina species described by Endrödi (1971, 1976). The alpha taxonomy may
require revision.

A single species is found in NE savannas of KwaZulu-Natal, South Africa, where it has been recorded only from game
reserves. However, it ranges into the tropics as S to E, and, E to W African subspecies.

ONITICELLINI
SURICATA 6 (2020) 311

Ixodina abyssinica
Roth, 1851

subsp. abyssinica Roth, 1851

subsp. tangana (Endrödi, 1971)
Global: DD

J A S O N D J F M A M J

Type localities: I. abyssinica: ‘Tigré in N. Abyssinien’ [Tigray, N Ethio-

pia]; as Drepanocerus abyssinicus Roth subsp. tanganus: ‘Mt. Meru E
slope, 5700 feet’ [N Tanzania].
Taxonomy: Accepted species in the subfamily Drepanocerina; southern
African material should be compared with types of subspecies; sexual
dimorphism (head, prothoracic disc) characters varying in prominence
with body size.
Distribution: Widespread in dry to moister savanna regions of Africa;
subsp. abyssinica: W to NE Africa, Guinea (Bissau), Guinea (Conakry),
Ivory Coast, Niger, Cameroon, Democratic Republic of the Congo
(DRC), Ethiopia, Djibouti; subsp. tangana: E to S Africa, Kenya, Tan-
zania, Mozambique, South Africa (see range map).
Locality data (mean ± SD, range): Altitude: 229 ± 145, 55–450 m; an-
nual rainfall: 888 ± 110, 743–1 075 mm; annual temperature (max. +
min. /2): 21.3 ± 0.9, 20.5–22.8°C (N=6).
Habitat: No quantitative assessment; limited DBRU collection records
from sand (2) in open woodland (1) and pasture (1).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (2), buffalo (1), rhinoceros (1).
Temporal activity: Diurnal flight activity, primarily in the summer rainy
season (Aug. to Apr.).
Bioregions South Africa: S extremity of Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 347 255 km2 (underestimated); AOO
would be smaller and limited to moister climate; restricted distribution
pattern possibly a mix of localisation and collection artefact due to
small body size; ecologically poorly known; apparently widespread, so
may deserve Least Concern (LC) status, however, currently assessed as
Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by more comprehensive data as known range may be underes-
timated and ecological associations are poorly known; in South Africa,
all known localities protected in uMkhuze and Hluhluwe–iMfolozi 2 mm
game reserves.

ONITICELLINI
312 SURICATA 6 (2020)

Genus Latodrepanus Krikken, 2009

Type species and designation: Drepanocerus laticollis Fahraeus, 1857, by original designation.
Synonyms: = Drepanellus Barbero, Palestrini & Roggero, 2009b: Type species: Drepanocerus lati-
collis Fahraeus, 1857, by original designation.
Last review: Entire genus reviewed by Krikken (2009), new species added by Barbero et al. (2009b).

The Afrotropical genus, Latodrepanus Krikken, 2009, now comprises three species following the description of two new
species from West (Guinea) and SE Africa (Malawi) (Barbero et al. 2009b) and the transfer of three W to E or E African
species to the new genus Epidrepanus Roggero, Barbero & Palestrini, 2015. A single species occurs in South Africa. It
shows a dry, sandy, savanna distribution from S to S Central to E Africa.

ONITICELLINI
SURICATA 6 (2020) 313

Latodrepanus laticollis
(Fahraeus, 1857)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Drepanocerus laticollis: ‘juxta fluvium Limpopo’ [near

Limpopo River, southern Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina.
Distribution: Moist savanna on sandy soils from S to E Africa: NE South
Africa, NE Botswana, Angola, Zimbabwe, Zambia, Mozambique,
Democratic Republic of the Congo (DRC), Tanzania, Kenya; citations
from W Africa (Senegal, Guinea, Benin) represent confusion with ♂
other Latodrepanus or Epidrepanus species; disjunctions may partly
reflect distribution of suitable climate and sandy soils.
Locality data (mean ± SD, range): Altitude: 832 ± 427, 0–1 462 m; an-
nual rainfall: 701 ± 184, 418–1 275 mm; annual temperature (max. +
min. /2): 20.4 ± 1.7, 17.3–24.7°C (N=80).
Habitat: In Gauteng bushveld: extreme bias to deep sand (2 080) com-
pared to sandy clay loam (54) with bias to open woodland (1 714)
compared to grassland (386) or shaded thickets (34); DBRU collection
records biased to sandy soils: sand (20), sandy loam (12), sandy clay
loam (8) in woody vegetation: grassland/pasture (12), shrubland (8),
open woodland (19), shaded thicket (1).
Food types: At Phalaborwa: extreme bias to herbivore dung (ruminant
cattle: 240; monogastric elephant: 169) compared to omnivore dung
(pig: 9); similar in Gauteng bushveld: cattle (327), horse (206), pig
(14), carrion (0); DBRU collection records from dung of cattle (42),
rhinoceros (2), donkey (1), waterbuck (1), human (1).
Temporal activity: Diurnal flight activity peaking at midday and early
afternoon (12:00–16:00) in Gauteng bushveld; primarily active in the
summer rainy season (Oct. to Apr.); abundant from Oct. to Jan., tailing
off Feb. to Apr., rare May to July; at Phalaborwa: more abundant in
warm sunny weather soon after heavy rainfall (951) than on a hot dry 1 mm

day (157) or a warm cloudy day following light rainfall (30).

Ecoregions Botswana, Zimbabwe, Mozambique: Southern African
Bushveld (AT0717), Southern Miombo Woodlands (AT0719), moist-
er W of Zambezian and Mopane Woodlands (AT0725); Maputaland
Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Primarily centred on Central Bushveld (SVcb),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 4 469 700 km2 (rough estimate); AOO
would be smaller according to soil and vegetation specialisation; clear-
ance of woody vegetation on sandy soils would presumably reduce
population density; however, very widespread and protected in various
reserves; therefore, assessed as Least Concern (LC).
Conservation measures: None recommended; protected in Kruger Na-
tional Park, Leeuwfontein and Tswaing nature reserves (South Africa).

ONITICELLINI
314 SURICATA 6 (2020)

Genus Liatongus Reitter, 1893

Type species and designation: Onthophagus phanaeoides Westwood, 1839, by subsequent designation (Arrow 1931).
Synonyms: None (but see below).
Last review: Entire genus reviewed by Janssens (1953); new species added by Balthasar (1956,
1964a), Scheuern (1988), Král and Rejsek (1999); two species removed to other gen-
era Attavicinus monstosus (Bates 1887), Paroniticellus festivus (Steven 1809); one species
transferred from Oniticellus Dejean, 1821, may be misclassified within Liatongus (of-
ten cited as Scaptodera rhadamistus (Fabricius, 1775)).

Because of common usage, Liatongus Reitter, 1893, is retained here as the generic name, although it has recently been
noted (Branco 2011) that it is a junior synonym of Onthosphaenus Motschulsky, 1860 (Type species: Scarabaeus vertagus
Fabricius, 1798, by subsequent designation (Paulian 1986)).

Liatongus is widespread in moist forest, savanna and upland grassland regions where it is represented by a total of 44
species in the Afrotropical (16), Oriental (23), Palaearctic (3) and Nearctic regions (2), with a further three species shared
between the Oriental and Palaearctic. One further Oriental species, formally transferred by Lumaret and Moretto (1983)
from Oniticellus Dejean, 1821, into the Liatongus subgenus Paraliatongus Balthasar, 1963d, may be misclassified and is
often cited within the genus Scaptodera Hope, 1837 (see above).

Since Janssens’ review (1953), the single Neotropical species has been removed from Liatongus into the new genus Atta-
vicinus Philips & Bell, 2008. A Palaearctic species has been removed into the genus, Paroniticellus Balthasar, 1963d (see
above). However, as presently constituted the genus remains paraphyletic (Philips 2016) based on relationships of two
species, one Afrotropical, one Nearctic.

Only two Liatongus species are found within the study area of Namibia, Botswana and South Africa. One of these is ap-
parently an undescribed forest endemic with a restricted range (no species account prepared). The other is widespread in
moist E savannas and grasslands with a range extending into the S central and E tropics.

ONITICELLINI
SURICATA 6 (2020) 315

Liatongus militaris
(Castelnau, 1840)

= Oniticellus tridens Roth, 1851

= Oniticellus quadrituberculatus van Lansberge, 1886
Global: LC

J A S O N D J F M A M J

Type localities: As Oniticellus militaris: ‘Cap de Bonne-Espérance’ [Cape

of Good Hope, South Africa]; O. tridens: ‘Tigré, N. Abyssinien’ [Tigray
Region, Ethiopia]; O. quadrituberculatus: ‘l’Afrique Occidentale’
[Humpata, Angola].
Taxonomy: Accepted species; limited sexual dimorphism (head, protho-
racic disc). ♂
Distribution: Moist grassland and savanna from S and S Central to NE
Africa: South Africa, Zimbabwe, Mozambique, Angola, Kenya, Tan-
zania; marginal in N Botswana; also cited from Eritrea, Sudan, Egypt.
Locality data (mean ± SD, range): Altitude: 949 ± 532, 0–2 274 m; an-
nual rainfall: 720 ± 181, 309–1 659 mm; annual temperature (max. +
min. /2): 18.4 ± 2.4, 10.6–25.2°C (N=315).
Habitat: In Gauteng bushveld: strong bias to finer-grained soils: deep
sand (66), sandy clay loam (1 261) in open vegetation: grassland
(1 013), open woodland (297), shaded thickets (17); partly supported
by DBRU collection records: sand (35), sandy loam (44), sandy clay
loam (49), clay (14) in grassland/pasture (97), scrub/shrubland (10),
open woodland (19), thickets (3).
Food types: At Phalaborwa: strong bias to dung of elephant (22) rather
than that of cattle (4) or pig (0); DBRU collection records dominated
by common herbivore dung types outside of reserves: cattle (163), buf-
falo (6), elephant (1), horse (1).
Temporal activity: Diurnal flight activity primarily during the summer
rainy season (Oct. to Apr.); at Phalaborwa: frequency much greater in
hot, dry conditions (20) compared to warm cloudy conditions follow-
ing light rainfall (4) or warm sunny weather soon after heavy rainfall
(2).
Ecoregions Botswana, Zimbabwe: Primarily Southern Miombo Wood- 2 mm
lands (AT0719), Southern African Bushveld (AT0717).
Bioregions South Africa: Centred in Savanna (SV), Grassland (G), In-
dian Ocean Coastal Belt (CB) biomes; Bioregions: Central Bushveld
(SVcb), Lowveld (SVl), Mesic Highveld Grassland (Gm), Dry High-
veld Grassland (Gh), Sub-Escarpment Grassland (Gs); also Albany
Thicket Biome (AT), SE Fynbos Biome, primarily in transformed
habitats; marginal in Eastern Kalahari Bushveld (SVk).
Assessment rationale: EOO = 3 632 435 km2 (underestimated); wide-
spread with a bias to open habitats and a high frequency of records
from cattle dung in farmland despite a measured bias to elephant dung;
assessed as Least Concern (LC).
Conservation measures: None recommended; protected in Hluhluwe–
iMfolozi Game Reserve (South Africa) and Gorongosa National Park
(Mozambique).

ONITICELLINI
316 SURICATA 6 (2020)

Genus Oniticellus Dejean, 1821

Type species and designation: Scarabaeus cinctus Fabricius, 1775, by subsequent designation (Arrow 1931).
Synonyms: = Pseudoniticellus Kraatz, 1895: Type species: Oniticellus planatus Castelnau, 1840, by
monotypy.
Last review: Entire genus reviewed by Janssens (1953) with one species, thereafter, formally trans-
ferred to Paraliatongus Balthasar, 1963d (Lumaret & Moretto 1983); new species add-
ed by Balthasar (1964a), Biswas and Chatterjee (1985).

Addition and subtraction of species following the review of Janssens (1953), reduces membership of Oniticellus Dejean,
1821, to seven, perhaps eight, species found in the Afrotropical (5) or Oriental regions (2 +?1). The included species all
show essentially endocoprid breeding habits and have a history of misidentification and assignment to different genera,
including Pseudoniticellus Kraatz, 1895; Liatongus Reitter, 1893; and Pseudoniticellus sensu Paulian, 1945. Two new species
added after Janssens review comprise an Afrotropical forest species (O. pseudoplanatus Balthasar, 1964a) and an Oriental
species (O. gayeni Biswas & Chatterjee, 1985) that requires validation as it is cited from the same droppings as a close rel-
ative. One Oriental species subtracted after Janssens review has been variously cited as a species of Paraliatongus Balthasar,
1963; Liatongus (Paraliatongus), Liatongus or Scaptodera Hope, 1837 (cf. S. rhadamistus (Fabricius, 1775)).

In South Africa, Botswana and Namibia, Oniticellus is represented by four widespread species, all assessed as Least Con-
cern (LC), two with ranges extending into the E tropics and two with ranges extending into the E tropics and onwards into
W Africa. In southern Africa, the species are centred on elephant or equine dung in savanna (1); or various dung types in E
savanna (1); E savanna plus E upland grassland and the S Cape coast (1); or just E upland grassland and the S Cape coast
(1). These four species either construct brood chambers in droppings without tunnelling in the soil (2) or they construct
brood chambers in the soil in contact with the dropping (2). Monitoring of two species for an entire year has shown that
they remain active as adults throughout the rainy and dry seasons without any detected dormancy.

ONITICELLINI
SURICATA 6 (2020) 317

Oniticellus egregius
Klug, 1855

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Tette’ [Tete, Central Mozambique].

Taxonomy: Accepted species, once separated into the genus Pseudoniticel-
lus sensu Paulian, 1945, but last reviewed as Oniticellus (Janssens 1953).
Distribution: Widespread in dry savanna from S to E Africa: South Afri-
ca, Namibia, Botswana, Mozambique, Tanzania, Kenya; also reported
from Zimbabwe, Democratic Republic of the Congo (DRC), Somalia.
Locality data (mean ± SD, range): Altitude: 748 ± 450, 33–1 524 m; ♂
annual rainfall: 597 ± 156, 263–904 mm; annual temperature (max. +
min. /2): 20.7 ± 2.3, 15.6–23.6°C (N=38, exact records).
Habitat: No quantitative assessment; limited DBRU collection records
from sandy loam (3) in grassland/pasture (1) or open woodland (3).
Food types: No adequate quantitative assessment; at Phalaborwa (2) and
in Botswana (4): only sampled to elephant dung; DBRU collection
records from dung of elephant (6), rhinoceros (1) and horse (1).
Temporal activity: Diurnal flight activity recorded in the summer rainy
season (Oct. to Mar.); may show greater seasonality than strictly endo-
coprid Oniticellus species as nests are sited in pits in the soil under the
edge of droppings and contain clay coated broods that are not tended
by the parent female during larval development.
Ecoregions Namibia, Botswana: Namibian Savanna Woodlands
(AT1316), Angolan Mopane Woodlands (AT0702), Kalahari Acacia-
Baikiaea Woodlands (AT0709).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); margins of two other bioregions.
Assessment rationale: EOO = 2 407 180 km2; AOO much smaller
owing to specialisation to coarse-fibred dung of monogastric herbi-
vores whose distribution has largely contracted to within the borders
of game reserves or to areas such as Gauteng (South Africa) where there
are adequate populations of indigenous (zebra) or domestic equines
(horse); however, assessed as Least Concern (LC) owing to widespread
protection in many game reserves which account for 65% of available 2 mm

southern African records (46).

Conservation measures: Conservation status of this species would be
dependent on continuing protection of large monogastric herbivores
in savanna reserves and maintenance of equine populations elsewhere;
assessment of conservation status would be improved by quantitative
data on habitat associations and how these might further influence the
AOO; protected in various national parks and game reserves including
Kruger, uMkhuze, Hluhluwe–iMfolozi (South Africa), Etosha (Namib-
ia), Manyara (Tanzania), Tsavo, Masai Mara, Amboseli, Meru (Kenya).

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Oniticellus formosus
Chevrolat, 1830

= Oniticellus pictus Hausmann sensu Péringuey, 1901 (pars)

= Oniticellus parapictus Balthasar, 1941a
Global: LC

J A S O N D J F M A M J

Type localities: O. formosus: ‘Sénégal’ [Senegal]; O. pictus sensu Péringuey

(pars): ‘Transvaal (Rustenburg)’ [South Africa]; O. parapictus: ‘Senegal’.
Taxonomy: Accepted species assigned to the genus, Liatongus Reitter, 1893,
by d’Orbigny (1916), but reviewed as Oniticellus by Janssens (1953).
Distribution: Mainly in dry savanna from W to E to southern Africa: Sene-
gal, Côte d’Ivoire, Niger, Nigeria, Kenya, Tanzania, Mozambique, Zim-
babwe, Botswana, South Africa; also reported from Mauritania, Angola.
Locality data (mean ± SD, range): Altitude: 752 ± 471, 8–1 706 m; an-
nual rainfall: 608 ± 207, 205–1 168 mm; annual temperature (max. +
min. /2): 20.4 ± 2.2, 14.4–25.2°C (N=77).
Habitat: In Gauteng bushveld: bias to finer-grained soils: sandy clay loam
(96) rather than deep sand (19) where there is greater dung removal,
also bias to more open habitats: grassland (80), open woodland (34)
rather than shaded thickets (1); largely supported by DBRU collection
records: sand (13), sandy loam (12), sandy clay loam (9), clay (7), in
grassland/pasture (22), shrub/woodland (17).
Food types: At Phalaborwa: strong bias to elephant dung (32) as opposed to
that of cattle (2) and pig (0); similar results in Botswana: elephant (16), cattle
(2), sheep (3), pig (1); at Abokouamekro (Côte d’Ivoire): recorded to cat-
tle (41), but not pig dung; DBRU collection records primarily from rumi-
nant herbivore dung: cattle (33), buffalo (6), rather than monogastric her-
bivore dung: elephant (4), rhinoceros (1), donkey (1); also waterbuck (1).
Temporal activity: Diurnal flight activity; recorded during every month
of the year; probably because this species does not tunnel in the soil and
therefore does not remain dormant in tunnels during unsuitably cold
weather; at Phalaborwa: frequency much greater in hot, dry conditions
(29) than in warm cloudy conditions following light rainfall (0), or in
warm sunny conditions soon after heavy rainfall (6); in Gauteng bush-
2 mm
veld: greatest breeding success within cattle pads at the end of the dry
season (Aug., Sept.) when dung removal is minimal and temperatures
are increasing.
Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725), E Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern
Miombo Woodlands (AT0719).
Bioregions South Africa: Centred in Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp) (Savanna Biome); also N Indian Ocean Coastal
Belt Biome (CB) with sporadic occurrence in Eastern Kalahari Bush-
veld (SVk), NE Bushmanland (NKb).
Assessment rationale: EOO = 4 827 535 km2; strong bias to large droppings
of herbivores reflects endocoprid breeding habits (nest constructed within
dropping containing broods that are tended by parent female during lar-
val development); population density is low reflecting exploitation com-
petition with dung-burying beetles and, probably, density dependence of
breeding sites within dung (primarily, one nest per cattle pad); over most
of the range elephant droppings no longer occur; however, extremely
widespread, therefore, assessed as Least Concern (LC).
Conservation measures: None recommended; commonly recorded in
low numbers from cattle dung in farmland despite a measured bias
to elephant dung; protected in Kruger National Park and Hluhluwe–
iMfolozi Game Reserve.

ONITICELLINI
SURICATA 6 (2020) 319

Oniticellus pictus
(Hausmann, 1807)

= Oniticellus pictus Péringuey, 1901 (pars)

= Oniticellus pictus orientalis Janssens, 1953
Global: LC

J A S O N D J F M A M J

Type localities: As Aphodius pictus: Not stated; O. pictus Péringuey: ‘Cape

Colony (Uitenhage, Graham’s Town, Queen’s Town, Griqualand West,
Knysna, Somerset East), Natal (Frere, Durban, Maritzburg)’ [Eastern
Cape, South Africa; KwaZulu-Natal citations may be either O. pictus
or O. formosus Chevrolat]; subsp. orientalis: ‘Est du Congo Belge: Parc
National Albert’ [Virunga National Park, E Democratic Republic of
the Congo (DRC)].
Taxonomy: Accepted species.
Distribution: Disjunct distribution on the East Central African highlands
(Democratic Republic of the Congo (DRC)) and the South African
Highveld to SE coastal lowlands: additional reports from Rwanda,
Tanzania; Mozambique report requires validation; Guinea presumably
an error.
Locality data (mean ± SD, range): Altitude: 834 ± 701, 16–2 194 m;
annual rainfall: 661 ± 188, 412–1 065 mm; annual temperature (max.
+ min. /2): 16.5 ± 2.0, 11.0–20.3°C (N=35).
Habitat: No quantitative assessment; DBRU collection records only from
finer-grained soils: sandy loam (6), sandy clay loam (10), clay (1), pri-
marily in grassland/pasture (12), less frequently in shrub/woodland (4).
Food types: No quantitative assessment; DBRU collection records mainly
from cattle dung (23), rarely horse dung (1).
Temporal activity: Diurnal flight activity; recorded during both the dry
(May, Aug., Sept.) and rainy seasons (Oct. to Mar.); several individuals
observed clustered together in the centre of a cattle pad during a snow
storm over natural grassland (Machadodorp [= Makhado] 16 km E),
reflecting non-soil tunnelling, endocoprid habits (breed within drop-
pings).
Bioregions South Africa: E Mesic Highveld Grassland (Gm), Sub- 2 mm

Escarpment Grassland (Gs) (Grassland Biome); S Indian Ocean Coast-

al Belt Biome (CB); Albany Thicket Biome (AT); transformed S coastal
habitats in Fynbos Biome.
Assessment rationale: EOO = 131 510 km2; widely recorded in low
population density from cattle dung in farmland; owing to widespread
distribution, not thought to be threatened, thus, assessed as Least Con-
cern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
whether or not pasture improvement influences its occurrence; not
known from any nature reserve at the present time.

ONITICELLINI
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Oniticellus planatus
Castelnau, 1840

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Cap de Bonne-Espérance’ [Cape of Good Hope, South

Africa].
Taxonomy: Accepted species assigned to the genus Pseudoniticellus Kraatz,
1895, then to Liatongus Reitter, 1893, by d’Orbigny (1916), but re-
viewed as Oniticellus by Janssens (1953).
Distribution: Mainly in warmer moist savanna and warmer moist grass-
lands from W to E to southern Africa: Côte d’ Ivoire, Kenya, Tanzania,
Mozambique, Angola, Zimbabwe, NE Namibia, N Botswana,
Eswatini, South Africa; also reported from Guinea (Conakry), Central
African Republic, Democratic Republic of the Congo (DRC), Ethiopia,
Somalia, Burundi, Rwanda.
Locality data (mean ± SD, range): Altitude: 790 ± 583, 0–2 194 m; an-
nual rainfall: 731 ± 195, 365–1 659 mm; annual temperature (max. +
min. /2): 18.8 ± 2.7, 11.0–25.2°C (N=186).
Habitat: No adequate quantitative data; DBRU collection records from
sand (14), sandy loam (24), sandy clay loam (23), clay (8) in grassland/
pasture (45), shrub/woodland (21), forest (2).
Food types: No adequate quantitative data; both at Phalaborwa and in Bot­
swana: recorded only to elephant dung (5 at each), not ruminant her-
bivore or omnivore dung; however, DBRU collection records primarily
from ruminant herbivore dung: cattle (80), buffalo (4); also monogas-
tric herbivore dung: elephant (13), rhinoceros (4), horse (2), zebra (1).
Temporal activity: Diurnal flight activity; recorded during every month of
the year; probably because this species is not known to tunnel in the soil
under field conditions or to remain dormant in tunnels during unsuitably
cold weather; in Gauteng bushveld: greatest breeding success at the in-
terface between soil and cattle pads was at the end of the cool, dry season
(Aug., Sept.) when dung burial is minimal and temperatures are increasing.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Southern
Miombo Woodlands (AT0719), Southern African Bushveld (AT0717),
2 mm moister margins of Zambezian and Mopane Woodlands (AT0725),
Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome); sporadic occurrence in Grassland Bi-
ome with greater frequency only in warmer E Mesic Highveld Grass-
land (Gm) and N Sub-Escarpment Grassland (Gs); Indian Ocean
Coastal Belt (CB) and Albany Thicket (AT) biomes; transformed habi-
tats in Eastern Cape part of Fynbos Biome.
Assessment rationale: EOO = 4 837 160 km2; bias to large droppings of
herbivores reflects endocoprid breeding habits (nest constructed at drop-
ping/soil interface containing broods that are tended by female parent
during larval development); population density is low, reflecting ex-
ploitation competition with dung-burying beetles and probable density
dependence of breeding sites within dung (primarily, one nest per cattle
pad); over most of the range elephant droppings no longer occur; how-
ever, extremely widespread, therefore, assessed as Least Concern (LC).
Conservation measures: None recommended; commonly recorded in
low numbers from cattle dung in farmland despite a measured bias to
elephant dung; protected in moister parts of Kruger National Park and
Hluhluwe–iMfolozi Game Reserve.

ONITICELLINI
SURICATA 6 (2020) 321

Genus Paraixodina Roggero,

Barbero & Palestrini, 2015
Type species and designation: Drepanocerus runicus Arrow, 1909, by original designation.
Synonyms: None.
Last review: Genus created and reviewed with a morphological phylogeny by Roggero et al. 2015.

Paraixodina Roggero, Barbero & Palestrini, 2015, currently comprises three very small-bodied species from savannas in
the E Afrotropical (2) and E Oriental regions (1). The two Afrotropical species range from E Savanna of South Africa into
the E tropics. Both are primarily associated with monogastric herbivore dung.

ONITICELLINI
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Paraixodina freyi
(Janssens, 1953)

No synonyms
Global: DD (see IUCN Red List as Ixodina freyi – DD)

J A S O N D J F M A M J

Type locality: As Drepanocerus freyi: ‘Zululand : Hluhluewe’ [presumably,

Hluhluwe Game Reserve, KwaZulu-Natal, South Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina.
Distribution: Limited patchy records along the E seaboard of Africa;
moist lowland savanna in NE South Africa, upland grassland in Kenya
(1 500 m); also cited from Tanzania.
Locality data (mean ± SD, range): Altitude: 246 ± 253, 62–879 m; an-
nual rainfall: 795 ± 135, 566–982 mm; annual temperature (max. +
min. /2): 21.2 ± 1.2, 18.4–22.4°C (N=9).
Habitat: No quantitative assessment; extremely limited DBRU collection
records from sandy clay loam (1), clay (1) in open woodland (2).
Food types: No quantitative assessment; extremely limited DBRU col-
lection records from dung of cattle (1), rhinoceros (1); recorded from
zebra dung in Kenya (1).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Nov. to Feb., May).
Bioregions South Africa: Centred on Lowveld (SVl) (Savanna Biome),
marginal in extreme N tip of Sub-Escarpment Grassland (Gs) (Grass-
land Biome).
Assessment rationale: EOO = 450 215 km2 (gross estimate); poorly
known ecologically; unclear if soil type or vegetation type influences
occurrence; food associations unclear although all but one of the avail-
able records are from game reserves where elephants and rhinoceros
occur; thus, known range in South Africa mostly protected; assessed as
Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
dung type associations; P. freyi is thought to be a monogastric dung
specialist, which would have important conservation implications as
regards a possibly highly restricted AOO within a large EOO; pro-
2 mm tected in Hluhluwe–iMfolozi and uMkhuze game reserves plus Kruger
National Park (South Africa), Masai Mara Game Reserve (Kenya).

ONITICELLINI
SURICATA 6 (2020) 323

Paraixodina saegeri
(Balthasar, 1963e)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: As Drepanocerus saegeri: ‘National Parc de la Garamba’ [Ga-

ramba National Park, NE Democratic Republic of the Congo (DRC)].
Taxonomy: Accepted species in the subfamily, Drepanocerina; morphol-
ogy of southern African and Tanzanian individuals matches original
description, but identity should be validated by comparison with type
material.
Distribution: Limited patchy records in moist lowland savanna along the
E seaboard of Africa; South Africa, Botswana, Tanzania, NE Democrat-
ic Republic of the Congo (DRC); also cited from Kenya; citation from
Côte d’Ivoire requires validation; disjunctions on map reflect samples
recorded within or close to game reserves where monogastric herbivores
occur.
Locality data (mean ± SD, range): Altitude: 480 ± 311, 117–1 000 m;
annual rainfall: 668 ± 142, 529–904 mm; annual temperature (max. +
min. /2): 21.7 ± 0.6, 20.5–22.5°C (N=12).
Habitat: No quantitative assessment; observed on sand, sandy loam and
sandy clay loam in open woodland and forest.
Food types: At Phalaborwa: extreme bias to monogastric herbivore dung
(elephant: 27) compared to dung of ruminant herbivores (cattle: 1)
and omnivores (pig: 2); one DBRU collection record from rhinoceros
dung; also recorded in zebra dung.
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Feb.); at Phalaborwa: primarily active on a hot sunny day
(28), few active in warm cloudy weather following light rain (2) or in
warm weather soon after heavy rainfall (0). 1 mm
Ecoregions Botswana: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 073 235 km2 (gross estimate for E sea-
board); AOO would be much smaller because of range contraction
by large indigenous monogastric herbivores to whose dung P. saegeri
is specialised; limited patchy records probably a collection artefact re-
lated to both dung type specialisation and small body size; probably
not threatened due to large range and protection in various reserves;
however, poorly known and assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations and further survey
data to determine full EOO and AOO; protected in various national
parks and game reserves, including Hluhluwe–iMfolozi and Kruger
(South Africa), Chobe (Botswana), Manyara (Tanzania).

ONITICELLINI
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Genus Tibiodrepanus Krikken, 2009

Type species and designation: Copris setosus Wiedemann, 1823, by original designation.
Synonyms: = Sulcodrepanus Krikken, 2009: Type species: Drepanocerus sulcicollis Castelnau, 1840,
by original designation.
Last review: All but one species of the genus reviewed by Barbero et al. (2011); phylogeny (Roggero
et al. 2015).

Tibiodrepanus Krikken, 2009, currently comprises seven species found in moist forest, savanna or grassland of the Afro-
tropical (2), Oriental (4) or S Palaearctic (Afghanistan: 1) regions. The single species represented in moist SE grasslands of
South Africa is widely represented across a range extending into the E and W tropics.

ONITICELLINI
SURICATA 6 (2020) 325

Tibiodrepanus sulcicollis
(Castelnau, 1840)

= Drepanocerus dispar Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type localities: As Oniticellus sulcicollis: ‘Cap de Bonne-Espérance’;

D. dispar: ‘Caffraria meridionali.....Cap. Bonae Spei.’ [both Cape of
Good Hope, South Africa].
Taxonomy: Accepted species in the subfamily, Drepanocerina; sexual
dimorphism varying in prominence with body size (prothoracic disc).
Distribution: Moist grassland from S to E to W Africa: South Africa,
Mozambique, Democratic Republic of the Congo (DRC), Kenya, ♂
Ethiopia, Cameroon, Guinea (Conakry), Côte d’Ivoire; also cited from
13 other countries in S (1), E (4) and W (8) Africa.
Locality data (mean ± SD, range): Altitude: 893 ± 601, 55–2 140 m;
annual rainfall: 725 ± 171, 389–1 065 mm; annual temperature (max.
+ min. /2): 16.6 ± 2.0, 11.3–22.6°C (N=38).
Habitat: No quantitative assessment; DBRU collection records all from
finer-grained soils; sandy loam (7), sandy clay loam (5), clay (1) in open
vegetation: grassland/pasture (11), scrub (2).
Food types: No adequate quantitative assessment; DBRU collection re-
cords entirely from cattle dung (14) although sampled to dung of horse
(5) rather than cattle (1) in Ithala Game Reserve.
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Oct. to Apr.).
Bioregions South Africa: E Mesic Highveld Grassland (Gm), Sub-
Escarpment Grassland (Gs) (Grassland Biome); Indian Ocean Coastal
Belt Biome (CB); mostly transformed habitats in Albany Thicket (AT)
and SE coastal Fynbos Biomes.
Assessment rationale: EOO = 5 683 200 km2 based on limited data; in
the SE: AOO centred on cool, moist climate; in the E: AOO centred
on tropical uplands; in the W: AOO apparently centred on wet tropical 2 mm
lowlands; AOO smaller than EOO, but also, apparently, large with
high frequency of records from cattle dung in open vegetation of farm-
land; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Ithala Game Reserve (South Africa).

♀ 2 mm

ONITICELLINI
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Genus Tiniocellus Péringuey, 1901

Type species and designation: Oniticellus spinipes Roth, 1851, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Branco (2010).

After the review of Branco (2010), Tiniocellus Péringuey, 1901, currently comprises six species represented in the savannas
of the Afrotropical (5) and W Oriental regions (1). In the study area of Namibia, Botswana and South Africa, the genus is
represented by two species, both centred in E savanna with one showing a range extending into the S central and E tropics.

ONITICELLINI
SURICATA 6 (2020) 327

Tiniocellus eurypygus
Branco, 2010

subsp. eurypygus Branco, 2010

subsp. transdrakensbergensis Branco, 2010
Global: LC
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type localities: subsp. eurypygus: ‘RSA, Limpopo, Potgietersrus’ [Moko-

pane, Limpopo Province]; subsp. transdrakensbergensis: ‘RSA, KwaZulu-
Natal, Itala Reserve’ [Ithala Game Reserve] [both South Africa].
Taxonomy: Accepted species.
Distribution: subsp. eurypygus: bushveld savanna, NW South Africa (red
squares); subsp. transdrakensbergensis: NE lowland savanna, South Afri-
ca plus Eswatini (black squares).
Locality data (mean ± SD, range): subsp. eurypygus: altitude: 1 212 ±
146, 943–1 491 m; annual rainfall: 638 ± 45, 528–709 mm; annual
temperature (max. + min. /2): 18.7 ± 0.8, 17.2–20.4°C (N=32). subsp.
transdrakensbergensis: altitude: 467 ± 383, 55–1 736 m; annual rainfall:
700 ± 150, 436–964 mm; annual temperature (max. + min. /2): 21.1
± 2.2, 14.0–23.7°C (N=35).
Habitat: In Gauteng bushveld, subsp. eurypygus: bias to deep sand (941)
compared to sandy clay loam (282), strong bias to open woodland
(1 043) compared to grassland (92) or shaded thickets (88) (cited as
T. spinipes (Roth)); at Roodeplaat Nature Reserve: grassland (1.1), open
woodland (6.3), shaded thickets (0.1); supported by DBRU collection
records for both subsp. on sand (8), sandy loam (7), sandy clay loam (3)
in pasture (2), open shrub/woodland (15).
Food types: At Phalaborwa, subsp. transdrakensbergensis: bias to dung of
monogastric herbivores (elephant: 117) compared to omnivores (pig:
2 mm
55) or ruminant herbivores (cattle: 30); DBRU collection records for
both subspp. from various dung types: elephant (2), rhinoceros (5),
Subsp. eurypygus
zebra (1), horse (3), baboon (1), cattle (8), buffalo (1), wildebeest (1),
waterbuck (1).
Temporal activity: Diurnal flight activity recorded in most months (July
to May); in Gauteng bushveld, subsp. eurypygus: primarily active in the
rainy season from early to mid-summer (Oct. to Jan.), tailing off in late
summer (Feb. to Apr.); very low activity in the cool dry season (May
to Sept.); at Phalaborwa, subsp. transdrakensbergensis: less active on a
warm, moist cloudy day (23) than on warm, moist sunny (96) or hot,
dry sunny (84) days.
Bioregions South Africa: subsp. eurypygus: Central Bushveld (SVcb);
subsp. transdrakensbergensis: Lowveld (SVl) (both Savanna Biome);
marginal in three other bioregions.
Assessment rationale: EOO = 123 625 km2; widespread in NE South
African savanna, but population density would be influenced by clear-
ance of partially shaded woody vegetation and contraction in ranges of
monogastric herbivores (habitat transformation: 0–58% (SVl), 2–49%
(SVcb)); however, frequently recorded in both reserves and farmland;
therefore, assessed as Least Concern (LC).
Conservation measures: Continuing conservation of monogastric her-
2 mm
bivores in woody vegetation would favour higher population density;
protected in Kruger National Park (subsp. transdrakensbergensis) and
various nature reserves: Leeuwfontein, Tswaing (subsp. eurypygus).
Subsp. transdrakensbergensis

ONITICELLINI
328 SURICATA 6 (2020)

Tiniocellus spinipes
(Roth, 1851)

= Oniticellus variegatus Fahraeus, 1857

= Onticellus humilis Gerstaecker, 1871
= Tiniocellus asmarensis Balthasar, 1968
Global: LC

J A S O N D J F M A M J

Type localities: As Oniticellus spinipes: ‘Abyssinien’ [Tigray, N Ethiopia];

O. variegatus: ‘juxta fluvium Limpopo’ [near Limpopo River, southern
Africa]; O. humilis: ‘Mbaramu’ [Mbalamu, NE Tanzania]; T. asmaren-
sis: ‘Ost-Afrika: Aethiopien, Erythraea, Umgebung von Asmara’ [envi-
rons of Asmara, Eritrea].
Taxonomy: Accepted species.
Distribution: Hot savanna from SE to NE Africa: NE South Africa,
Mozambique, SE Zimbabwe, N Botswana, NE Namibia, S Angola, S
Democratic Republic of the Congo (DRC), Zambia, Malawi, Tanza-
nia, Kenya, Ethiopia, Eritrea, Somalia; one tentative record from the
Sahel in Burkina Faso, W Africa.
Locality data (mean ± SD, range): Altitude: 470 ± 346, 28–1 397 m;
annual rainfall: 636 ± 219, 383–1 177 mm; annual temperature (max.
+ min. /2): 22.6 ± 1.8, 17.1–25.5°C (N=27).
Habitat: No quantitative assessment; limited DBRU collection records
from sandy loam (1), sandy clay loam (2), clay (1) in open woodland
(4), forest (1).
Food types: No quantitative assessment; DBRU collection records from
dung of elephant (8), baboon (1), buffalo (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Sept. to Mar.).
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian and Mopane
2 mm
Woodlands (AT0725); marginal occurrence in three other ecoregions.
Bioregions South Africa: N Lowveld (SVl), Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 2 951 525 km2; almost all South African
records from the Kruger National Park; unclear if dominance of col-
lection records from finer-grained soils in woodland on elephant dung
represent a soil, vegetation and food type bias that would influence the
AOO as regards habitat transformation and contraction in range of
large monogastric herbivores and their coarse-fibred dung types; how-
ever, extremely widespread in hot savanna with most of South African
range under protection; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; protected in var-
ious national parks: Kruger (South Africa), Gorongosa (Mozambique),
South Luangwa (Zambia), Tsavo (Kenya).

ONITICELLINI
SURICATA 6 (2020) 329

Genus Tragiscus Klug, 1855

Type species and designation: Tragiscus dimidiatus Klug, 1855, by monotypy.
Synonyms: = Deronitis Arrow, 1933: Type species: Deronitis epphipiatus Arrow, 1933, by monotypy.
Last review: Janssens (1953).

Tragiscus Klug, 1855, is monotypic and endemic to the Afrotropical region where it shows endocoprid breeding habits
within elephant and rhinoceros droppings. The single species shows a savanna distribution in the N of Namibia, Botswana
and South Africa with a range extending into the E tropics.

ONITICELLINI
330 SURICATA 6 (2020)

Tragiscus dimidiatus
Klug, 1855

= Deronitis ephippiatus Arrow, 1933

Global: LC

J A S O N D J F M A M J

Type localities: ‘Inhambane’ [SE Mozambique coast]; D. ephippiatus:

‘UGANDA: Katunguru, Kasinga Channel’.
Taxonomy: Accepted species.
Distribution: Dry to moist savanna from S to E Africa: South Africa,
Namibia, Botswana, Zimbabwe, Mozambique, Kenya, Uganda.
Locality data (mean ± SD, range): Altitude: 471 ± 354, 0–1 259 m; an-
♂ nual rainfall: 602 ± 178, 328–1 136 mm; annual temperature (max. +
min. /2): 22.4 ± 1.3, 19.6–25.2°C (N=24).
Habitat: No quantitative assessment; DBRU collection records from vari-
ous soil types: sand (2), sandy loam (3), clay (1), all in open woodland
(4).
Food types: No quantitative assessment; DBRU collection records all
from monogastric herbivore dung: elephant (7), rhinoceros (1).
Temporal activity: Diurnal flight activity; recorded during the summer
rainy season (Nov. to Apr.), but as a non-soil-tunnelling species con-
structing nest chambers within older droppings containing broods that
are tended by the parent female during larval development, it probably
remains active throughout the year as in other endocoprid taxa.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 2 192 930 km2; AOO would be much
smaller since all available records are from game reserves except for
two on game farms (square-lipped (white) rhinoceros dung) and one
in a hunting area (elephant dung); AOO reflects bias to dung of large
2 mm
monogastric herbivores whose ranges have now largely contracted to
within the borders of protected areas; currently assessed as Least Con-
cern (LC) since the EOO is large with records from various reserves.
Conservation measures: Conservation status of this species would depend
on continuing protection of elephants and rhinoceros in dry savanna
and would be improved by an investigation of soil and vegetation type
associations as a possible further limitation on AOO; protected in vari-
ous game reserves and national parks, including Kruger, Tembe (South
Africa), Lake Mutirikwe (Zimbabwe) Chobe (Botswana), Etosha (Na-
mibia), Gorongosa (Mozambique), Amboseli (Kenya).

♀
2 mm

ONITICELLINI
332 SURICATA 6 (2020)

TRIBE ONITINI
Castelnau, 1840

As Onitides; Type genus: Onitis Fabricius, 1798

The monophyletic, large-bodied, tunnelling tribe, Onitini, is represented across Afro-Eurasia although
most of the genera are Afrotropical endemics. After the recent synonymy of Janssensellus Cambefort, 1976,
with Acanthonitis Janssens, 1937 (Cambefort et al. 2010), the tribe now comprises 16 valid genera. These
are endemic to the Afrotropical (13) or S Palaearctic regions (1) with the two most speciose genera shared
between the Palaearctic, Oriental and Afrotropical regions.

Although primarily associated with savanna, grassland or winter rainfall shrublands, at least two monotypic
genera are found in rainforest (Allonitis Janssens, 1936; Lophodonitis Janssens, 1938b) and one ranges into
arid regions (Cheironitis van Lansberge, 1875b). There is also a strong trend to association with the dung
of monogastric herbivores in all or some members of at least nine genera, which has implications for their
conservation status given the restricted ranges now occupied by large indigenous monogastrics.

A total of 10 genera have been recorded for the Onitini in Botswana, Namibia and/or South Africa. Eight
are species-poor whereas two are widespread in Afro-Eurasia and are species rich.
SURICATA 6 (2020) 333

In southern Africa:

Two widespread Afro-Eurasian genera

(1) Cheironitis van Lansberge, 1875b: Five validated southern African species showing a bias to SW
Arid regions (3) or dry savanna (2); some with a bias to monogastric herbivore dung.
(2) Onitis Fabricius, 1798: A total of 32 southern African species with distributions centred on savanna
(20) upland grassland (6) savanna/grassland (2) or winter/bimodal rainfall shrublands (4). Close to
half of the species may show a bias to the dung of monogastric herbivores.

Three Afrotropical genera with small savanna ranges

(3) Anonychonitis Janssens, 1950: Monotypic, monogastric dung specialist in one reserve on NE South
African coastal hills.
(4) Kolbeellus Jacobson, 1906: Monotypic genus recently rediscovered in SE South Africa after a hiatus
of at least 100 years.
(5) Tropidonitis Boheman, 1857: Monotypic, monogastric dung specialist on NE South African and SE
Mozambique coastal sands.

Five Afrotropical genera with large savanna ranges and localised occurrence

(6) Gilletellus Janssens, 1937: One species recorded from buck pellets in N Namibia; possible wider
range to South Africa depends on accuracy of identification.
(7) Heteronitis Gillet, 1911: One monogastric dung specialist ranging from S to E Africa.
(8) Megalonitis Janssens, 1937: One southern African monogastric herbivore dung specialist species.
(9) Neonitis Péringuey, 1901: Monotypic, monogastric dung specialist centred on savannas from N
South Africa to E Democratic Republic of the Congo (DRC).
(10) Platyonitis Janssens, 1942: One species on moist deep sands of N Namibia and SW Angola; mono-
gastric herbivore dung specialist.

Owing to various attributes: small ranges, dung type specialisation and/or infrequency of records, several
monotypic genera or taxa of species-poor genera are assessed as threatened (NT: 1; VU: 2; DD/EN: 2).

ONITINI
334 SURICATA 6 (2020)

Genus Anonychonitis Janssens, 1950

Type species and designation: Anonychonitis freyi Janssens, 1950, by monotypy.
Synonyms: None.
Last review: Genus created by Janssens (1950).

Anonychonitis Janssens, 1950, is a monotypic genus recorded solely from rhinoceros dung in a small protected area of NE
KwaZulu-Natal, South Africa. It is apparent that its conservation would be dependent on the continuing conservation of
rhinoceros in Hluhluwe–iMfolozi Game Reserve in the face of an upsurge in poaching over the past few years.

ONITINI
SURICATA 6 (2020) 335

Anonychonitis freyi
Janssens, 1950

No synonyms
Global: NT (see IUCN Red List – NT)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Zululand: Umfulozi’ [Hluhluwe–iMfolozi Game Reserve,

KwaZulu-Natal, South Africa].
Taxonomy: Accepted species; limited sexual dimorphism (head, protho-
racic disc).
Distribution: Past EOO unknown; now restricted to lowland savanna in
Hluhluwe–iMfolozi Game Reserve, KwaZulu-Natal, South Africa. ♂
Locality data (mean ± SD, range): Altitude: 199 ± 65, 110–273 m; an-
nual rainfall: 878 ± 22, 878–932 mm; annual temperature (max. +
min. /2): 20.8 ± 0.3, 20.5–21.3°C (N=5).
Habitat: No quantitative assessment; DBRU collection records from clay
(1) in open woodland (2), grassland clearing (1).
Food types: No quantitative assessment; all DBRU collection records
from square-lipped (white) rhinoceros dung (4), which is dropped in
middens; often found at the base of older droppings; cryptic owing
to its habit of covering itself in dung fibres; constructs a single brood
ovoid at the end of a tunnel in the soil.
Temporal activity: Diel flight periodicity unknown; active in the summer
rainy season (Oct. to Mar.).
Bioregions South Africa: moist SE Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 960 km2 (area, Hluhluwe–iMfolozi Game
Reserve); AOO may be even smaller if soil and vegetation specialisa-
tions exist; known range entirely protected within a game reserve where
it has been recorded only from the middens of square-lipped (white)
rhinoceros, a monogastric herbivore species whose population density
2 mm
is being reduced by poaching throughout southern Africa, including
Hluhluwe–iMfolozi where, recently (2014), there has been an increase
in poaching; owing to very small EOO, AOO and potential poaching
pressures on rhinoceros, A. freyi is currently assessed as Near Threatened
(NT).
Conservation measures: Continued conservation of rhinoceros is essen-
tial for conservation of this species; a survey to precisely determine its
ecological associations, AOO, and population density would be advis-
able, including an investigation to determine if it colonises the dung
of elephants, which have been re-introduced into Hluhluwe–iMfolozi
Game Reserve.

♀
2 mm

ONITINI
336 SURICATA 6 (2020)

Genus Cheironitis van Lansberge, 1875b

Type species and designation: Scarabaeus furcifer Rossi, 1792, by subsequent designation (Arrow 1931).
Synonyms: = Uposlotus Costa, 1853, nomen oblitum: Type species: Scarabaeus furcifer Rossi, 1792,
by subsequent designation (Arrow 1931).
= Chironitis van Lansberge sensu Janssens, 1937, misspelling of Cheironitis.
Last review: Entire genus reviewed by Janssens (1937); one species subsequently made a syn-
onym of a species transferred from Heteronitis Gillet, 1911 (C. audens (Péringuey,
1901)); new Afrotropical species added by Janssens (1943), Balthasar (1963b), Ferreira
(1976a); subtraction of two Afrotropical species into a new genus, Pseudochironitis
Ferreira (1977).

Although often spelt as ‘Chironitis’ following publications by Bedel (1892a) and Reitter (1893), Branco and Ziani (2005)
made a case for van Lansberge’s original spelling of Cheironitis as the valid spelling for this genus, citing rules of nomen-
clature in support. They also suggest that the senior synonym, Uposlotus Costa, 1853, should be considered invalidated as
a nomen oblitum (= forgotten name) having been used as the genus name only once (Ádám 2003) since 1899.

A total of 22 species are currently placed in Cheironitis. These show distributions centred on drier regions of Afro-Eurasia.
They are either endemic to the Palaearctic (10) and Afrotropical regions (10), or shared between the Afrotropical and
Palaearctic (1), or Palaearctic and W Oriental regions (1).

In Africa, the general pattern of distribution is to the drier parts of the SW with a disjunction to the drier parts of the NE
and W. Five species currently occur in southern Africa with another two remaining undescribed. Three described species
show distributions centred on the SW arid late summer rainfall region with one also overlapping into the winter and bi-
modal rainfall regions during summer. One is distributed across Karoo, savanna and dry grassland. The fifth occurs in E
dry savanna where it is associated with monogastric herbivore dung. Although species found in dry or arid areas are not
threatened, possible effects of grassland modification and loss of zebra need to be investigated. Furthermore, adjustments
to generic membership may be necessary for several taxa currently classified as Cheironitis species.

ONITINI
SURICATA 6 (2020) 337

Cheironitis audens
(Péringuey, 1901)

= Chironitis damarensis Felsche, 1907

Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: As Onitis audens: ‘Cape Colony (Prieska)’ [Prieska,

Northern Cape, South Africa]; C. damarensis: ‘Kubub, D. S.-W. Afrika’
[?river or farm in Namibia].
Taxonomy: Accepted species; some sexual dimorphism (legs, prosternum).
Distribution: Arid late summer rainfall region with populations disjunct
in South Africa / S Namibia and N Namibia Plateau; disjunction does ♂
not appear to be a collection artefact.
Locality data (mean ± SD, range): Altitude: 1 100 ± 214, 608–1 855 m;
annual rainfall: 241 ± 74, 118–455 mm; annual temperature (max. +
min. /2): 17.9 ± 2.0, 11.9–23.0°C (N=105).
Habitat: No formal quantitative assessment; soil generalisation shown in
Northern Cape: sand (3.9), sandy loam (3.7), sandy clay loam (2.7);
sampled equally in Upper Karoo and Bushmanland (0.2/trap), rare in
SW Kalahari (0.001); largely supported by DBRU collection records
on sand (8), sandy loam (8), sandy clay loam (11) primarily in scrub/
shrubland (11) and open woodland (6), few in pasture (2).
Food types: No quantitative assessment; DBRU collection records from
ruminant: cattle (17), wildebeest (2) and monogastric herbivore dung:
zebra (4), horse (2), donkey (2).
Temporal activity: Diurnal flight activity in the late summer rainy season
(Dec. to Apr.).
Ecoregions Namibia: In S: Nama Karoo (AT1314); in N: restricted to
Angolan Mopane Woodlands (AT0702).
Bioregions South Africa: Upper Karoo (Nku), Bushmanland (NKb)
(Nama Karoo Biome); marginal in four other bioregions.
Assessment rationale: EOO = 208 390 km2 (assumes continuous range);
2 mm
widespread across arid regions used primarily for the grazing of live-
stock; readily attracted to the dung of both ruminant and monogastric
herbivores on farms where there is little transformation (NKb, NKu);
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations despite
observed, large, little-transformed range and association with dung of
farm livestock; a survey to demonstrate a truly disjunct or continuous
range would also be useful; little protection across main South African
range; however, protected in Etosha National Park (Namibia).

♀ 2 mm

ONITINI
338 SURICATA 6 (2020)

Cheironitis hoplosternus
(Harold, 1868)

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Onitis hoplosternus ‘Afric. austr. inter.’ [interior of south-

ern Africa].
Taxonomy: Accepted species; some sexual dimorphism (legs, prosternum).
Distribution: Dry W uplands, central South Africa encompassing: SE
Kalahari, NE Karoo, W Highveld grasslands; also sporadic marginal
♂ records from dry savanna: South Africa, Botswana, N Namibia Plateau.
Locality data (mean ± SD, range): Altitude: 1 187 ± 260, 62–1 964 m;
annual rainfall: 353 ± 153, 172–815 mm; annual temperature (max. +
min. /2): 17.7 ± 2.0, 12.5–22.9°C (N=138).
Habitat: No adequate quantitative assessment; in Northern Cape: primar-
ily sampled from SW Kalahari (0.7/trap), less in Upper Karoo (0.4)
and arid Bushmanland (0.002); DBRU collection records from a range
of soil and vegetation types: clay (3), sandy clay loam (5), sandy loam
(13), sand (12), on pasture/grassland (15), scrub/shrubland (6) and
open woodland (4); also fallow crop fields (2).
Food types: No quantitative assessment; DBRU collection records pri-
marily from dung of ruminant herbivores: cattle (33); but also mono-
gastric herbivore dung: horse (3), zebra (2), donkey (1); trapped to pig
dung in Botswana (1 individual).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Apr.)
Ecoregions Namibia, Botswana Kalahari Xeric Savanna (AT1309), Ka-
lahari Acacia-Baikiaea Woodlands (AT0709), Angolan Mopane Wood-
lands (AT0726).
Bioregions South Africa: N Upper Karoo (NKu) (Nama Karoo Biome);
2 mm
East Kalahari Bushveld (SVk), S Central Bushveld (SVcb) (Savanna
Biome); also Dry Highveld Grassland (Gh) (Grassland Biome); sporad-
ic occurrence in five other bioregions across four biomes.
Assessment rationale: EOO = 651 685 km2; main SW range little
transformed (0–2%; SVk, NKu); main NE range more transformed
(10–63%; SVk, Gh); isolated records in SW Cape pastures may be a re-
sponse to transformation of natural shrubland; widespread and readily
attracted to the dung of domestic livestock in farm rangeland; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would ben-
efit from quantitative data on ecological associations and population
density as there are currently few records from reserves in South Africa;
protected in Etosha National Park (Namibia).

2 mm ♀

ONITINI
SURICATA 6 (2020) 339

Cheironitis imitator
(Balthasar, 1963b)

No synonyms
Global: DD (see IUCN Red List as C. indicus – DD)

J A S O N D J F M A M J

Type locality: As Chironitis imitator: Holotype: ‘Sudan (Mongalla, Lado)’

[South Sudan: Mongalla, Lado].
Taxonomy: Accepted species, but southern African individuals need to be
compared with type material from NE Africa to validate their identity
as C. imitator rather than an extremely close undescribed relative; from
the descriptions of the prosternal spurs, it seems that Balthasar (1963b)
compared C. imitator with the true C. indicus from arid SW Africa ♂
(see species page for C. indicus van Lansberge, 1875b) rather than with
these southern African individuals treated here as C. imitator; some
sexual dimorphism (legs, prosternum).
Distribution: In southern Africa: shows an apparent disjunct pattern
in two lowland centres; Savuti, Okavango Delta, Botswana and NE
lowlands plus Limpopo Valley, South Africa; the major disjunction in
overall species range between S and NE plus E Africa may or may not
indicate two closely related species.
Locality data (mean ± SD, range): Altitude: 541 ± 251, 278–1 000 m;
annual rainfall: 486 ± 104, 309–655 mm; annual temperature (max. +
min. /2): 22.3 ± 0.6, 21.3–23.1°C (N=16).
Habitat: No quantitative data; DBRU collection records from various soil
types: sand (2), sandy loam (3), sandy clay loam (1) entirely in open
woodland (4).
Food types: No adequate quantitative data; at Phalaborwa: sampled to
pig dung (2); however, DBRU collection records from southern Africa
indicate a bias to coarse-fibred dung of monogastric herbivores: ele-
phant (5), donkey (3) plus cattle dung (1); many E African records for
C. imitator also from elephant and zebra dung. 2 mm
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Mar.).
Ecoregions Botswana: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Dry lowlands of Central Bushveld (SVcb),
Lowveld (SVl) and especially Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 669 950 km2 (gross estimate); AOO would
be smaller if disjunct pattern is real rather than a collection artefact;
disjunction possibly related to dung type association reflecting range
contraction onto game reserves for indigenous monogastric herbivores
and scattered occurrence of domestic monogastric herbivores in farm-
land; this could represent a threat over most of the potential EOO
in southern Africa; owing to limited ecological data, assessed as Data
Deficient (DD).
Conservation measures: Before conservation status may be assessed, taxo-
nomic uncertainties need to be resolved regarding identity as C. imitator
Balthasar, 1963, or a close undescribed relative; a survey to determine
the full EOO and AOO is also required as well as quantitative data on
ecological associations; protected in two national parks, Kruger (South
Africa), Chobe (Botswana) that support large populations of elephants.

♀
2 mm

ONITINI
340 SURICATA 6 (2020)

Cheironitis indicus
van Lansberge, 1875b

No synonyms
Gloal: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: Cited as ‘Inde en deçà du Gange. Arabie’ [India ‘below’

the Ganges; Arabia], but see Taxonomy section.
Taxonomy: Accepted species, but type unknown (Janssens, 1937); Jans-
sens (1937) states that material from Windhoek, Namibia, in Gillet’s
collection corresponds to van Lansberge’s original description, whereas
♂ it does not match Arrow’s (1931) description of material from India
[and Karachi, Pakistan] cited as C. indicus van Lansberge, 1875b;
this Indian material is now classified under the nom nov. of C. arrowi
(Janssens, 1937); also see the species account for C. imitator Balthasar,
1963b; some sexual dimorphism (legs, prosternum).
Distribution: Arid SW summer rainfall region from Bushmanland, South
Africa, N along and below the W escarpment as far N as Windhoek,
Namibia.
Locality data (mean ± SD, range): Altitude: 1 028 ± 203, 58–1 762 m;
annual rainfall: 196 ± 47, 14–255 mm; annual temperature (max. +
min. /2): 18.5 ± 1.4, 13.0–20.3°C (N=61).
Habitat: In Northern Cape: recorded to composite sheep/cattle baits in
low abundance (0.3–0.4/trap per study site) in 70 out of 406 sites;
Bushmanland (27 sites), Upper Karoo extension (27), SW Kalahari
(14), Upper Karoo (2); frequency of records on deep sand, loamy or
rocky sand sites (47) greater than sandy loam (22) or sandy clay loam
(1); limited DBRU collection records entirely on deep sand (4) in scrub
(2) or open woodland (2).
Food types: No quantitative assessment; limited DBRU collection records
on dung of horse (1), donkey (1), cattle (1).
Temporal activity: Diurnal flight activity during the late summer rainy
2 mm
season (Feb., Mar.).
Ecoregions Namibia: S Namibian Savanna Woodlands (AT1316), Namib
Desert (AT1315).
Bioregions South Africa: N Bushmanland (NKb), marginal in W Upper
Karoo (NKu).
Assessment rationale: EOO = 113 785 km2; centred in an area that has
undergone little habitat transformation and is used primarily for graz-
ing of farm livestock; not considered to face current threats as it has
been widely recorded in dung on farms in the Northern Cape; there-
fore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative assessment of ecological associations, particu-
larly as existing data suggest a bias to sandy soils; high spatial frequency,
but low abundance in samples, need to be investigated as they may
result from bias to monogastric herbivore dung like some relatives; may
be protected in parts of the Namib/Naukluft National Park, Namibia.

♀
2 mm

ONITINI
SURICATA 6 (2020) 341

Cheironitis scabrosus
(Fabricius, 1776)

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus scabrosus: ‘Surinami’ [Surinam – undoubt-

edly an error].
Taxonomy: Accepted species some sexual dimorphism (legs, prosternum).
Distribution: Dry to arid areas in the late summer, winter and bimodal
rainfall regions of SW South Africa and SW Namibia.
Locality data (mean ± SD, range): Altitude: 1 017 ± 321, 16–1 855 m; ♂
annual rainfall: 244 ± 81, 29–509 mm; annual temperature (max. +
min. /2): 17.6 ± 1.8, 11.9–20.5°C (N=242).
Habitat: No quantitative soil type assessment; in the SW on deep sands of
West Coast National Park: primarily sampled in pasture (46) following
clearance of natural shrubland, few in natural shrubland (4); in the NE,
primarily sampled from scrub in the Upper Karoo (5.6/trap), less in
arid Bushmanland (1.0) and deep sands of SW Kalahari (0.2); DBRU
collection records from clay (2), sandy clay loam (15), sandy loam (14),
sand (11) in shrubland (4), scrub (19), pasture/grassland (9) and crop
fields (3).
Food types: No quantitative assessment; DBRU collection records dom-
inated by ruminant herbivore dung: cattle (42), sheep (1); also mono-
gastric herbivore dung: horse (4)
Temporal activity: Diurnal flight activity during the summer rainy season
to the NE (Sept.–Apr.); similar seasonality during the dry mid-summer
of the winter rainfall region to the SW (Nov.–Feb.) peaking in Dec. in
West Coast National Park.
Ecoregions Namibia: S Namibian Savanna Woodlands (AT1316), Nama
Karoo (AT1314), Succulent Karoo (AT1322).
Bioregions South Africa: Upper Karoo (NKu), Bushmanland (NKb)
2 mm
(Nama Karoo Biome); also in Succulent Karoo, Fynbos (FF, FR) and
SW Savanna Biomes.
Assessment rationale: EOO = 478 375 km2; widespread; readily attracted
to the dung of domestic livestock in rangeland; little transformation
in scrub habitats in the arid NE of its range (0–2%; NK); to the SW
where transformation is high (5–80%; FF, FR), dry mid-summer ac-
tivity and much greater abundance in pastures may indicate range (or
at least population) expansion in response to clearance of indigenous
shrubland; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; few
records from protected areas, but widespread on domestic dung types
in farm rangeland and adaptable to transformation in the SW; despite
widespread occurrence and abundance, no current records from pro-
tected areas.

♀
2 mm

ONITINI
342 SURICATA 6 (2020)

Genus Gilletellus Janssens, 1937

Type species and designation: Onitis porculus Boheman, 1857, by subsequent designation (Janssens 1937).
Synonyms: None.
Last review: Entire genus reviewed by Janssens (1937).

Gilletellus Janssens, 1937, is represented by just two S Afrotropical species described from inexact localities presumed to be
in South Africa and possibly Mozambique, (‘Zambèze’ Province of Mozambique or elsewhere along the Zambezi River?).
Genus poorly known due to extremely infrequent records with little precise information. Assessed as Data Deficient (DD),
but only a single specimen known to have been recorded in the past 40 years.

ONITINI
SURICATA 6 (2020) 343

Gilletellus porculus
(Boheman, 1857)

No synonyms
Global: DD
Endemic: ?RSA, Namibia

J A S O N D J F M A M J

Type locality: As Onitis porculus: ‘Caffraria interiore’ [inexact, interior of

southern or South Africa].
Taxonomy: Accepted species; limited sexual dimorphism (legs); species
identification requires validation.
Distribution: A single record from the dry W of Etosha National Park
suggests that the inexact type locality may refer to the interior of ♂
southern Africa within the arid late summer rainfall region, but this is
speculative given its extreme rarity in collections.
Locality data (average): Altitude: 1 156 m; annual rainfall: 391 mm;
annual temperature (max. + min. /2): 22.2°C (N=1).
Habitat: No quantitative assessment; a single DBRU collection record
from sandy clay loam in scrub.
Food types: No quantitative assessment; a single DBRU collection record
from springbok pellets.
Temporal activity: Unknown.
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702).
Assessment rationale: EOO impossible to determine; a single specimen
recorded on springbok dung may or may not be indicative of dung type
specialisation in this excessively rare species; must be assessed as Data
Deficient (DD) because of the paucity of data, but its apparent rarity
is a cause for great concern; the single record would currently justify
Endangered (EN) or even Critically Endangered (CR) status.
Conservation measures: A survey from dry to arid savanna is required to
determine the local EOO, ecological associations, AOO and popula- 2 mm

tion density of G. porculus in W Etosha National Park where it is under

protection; such a survey would provide insights on how to assess its
conservation status across a potentially much larger EOO and AOO,
possibly southwards to South Africa.

ONITINI
344 SURICATA 6 (2020)

Genus Heteronitis Gillet, 1911

Type species and designation: Onitis tridens Castelnau, 1840, by subsequent designation (Janssens 1937).
Synonyms: None.
Last review: Entire genus reviewed by Janssens (1937) with subsequent removal of one species to
Cheironitis van Lansberge, 1875b (see C. audens (Péringuey, 1901)).

Heteronitis Gillet, 1911, currently comprises five Afrotropical species found in savannas from S to E to W Africa where
they are, apparently, associated primarily with elephant dung. Most species ranges are centred in E (2) or W Africa (2).
Only a single species shows a range from southern to E Africa; largely restricted to reserves in southern Africa, but assessed
as Least Concern (LC) on basis of wide EOO and protection in many reserves.

ONITINI
SURICATA 6 (2020) 345

Heteronitis castelnaui
(Harold, 1862)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Onitis castelnaui: ‘Zanzibar’ [Tanzania].

Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc,
legs), prominence of characters varies with body size.
Distribution: S to E African savanna; patchy occurrence due to dung type
specialisation; primarily centred on game reserves: N South Africa, N
Namibia, N Botswana, Zimbabwe, Mozambique, Angola, Zambia,
♂
Tanzania, Kenya; also reported from Democratic Republic of the
Congo (DRC), Rwanda.
Locality data (mean ± SD, range): Altitude: 565 ± 394, 55–1 448 m;
annual rainfall: 605 ± 168, 172–1 136 mm; annual temperature (max.
+ min. /2): 22.0 ± 1.7, 15.7–25.9°C (N=74).
Habitat: No quantitative assessment; DBRU collection records indicate a
bias to finer-grained soils: sandy clay loam (5), sandy loam (11), sand
(5), and woody vegetation: dry forest (1), open shrub/woodland (14),
scrub (3), grassland (2).
Food types: No quantitative assessment; DBRU collection records domi-
nated by monogastric dung types: elephant (25), hooked-lipped (black)
(2) and square-lipped (white) (4) rhinoceros, zebra (3), horse (1), don-
key (1); also warthog (1); few on ruminant dung types: cattle (3).
Temporal activity: Diel flight activity in fading light and crepuscular
darkness (late afternoon to dusk or dawn to early morning) (Caveney et
al. 1989) primarily in the summer rainy season (Oct. to May); attracted
to light.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), N Namibian Savanna Woodlands (AT1316), Zambe-
zian Baikiaea Woodlands (A0726), Zambezian and Mopane Woodlands
(AT0725), Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern 5 mm

African Bushveld (AT0717).

Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 3 997 830 km2 (gross estimate); AOO very
much smaller due to dung, and possible soil and vegetation specialisa-
tion; 82% (N=42/51) of DBRU collection records (1971–1986) from
proclaimed reserves; such concentration of recent records indicates a
clear effect of range contraction by large monogastric herbivores; single
records in areas distant from reserves (black squares) suggest very low
density relict populations (Buffelsdoorn in South Africa; West Nich-
olson in Zimbabwe); greater numbers have been recorded from areas
where rhinoceros have been re-introduced, e.g. reserves near Thabazim-
bi, South Africa (blue square); because of wide range and protection in
many reserves, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, including
population density both inside and outside of reserves; conservation
dependent on continuing protection of indigenous monogastric her-
bivores, particularly elephant and rhinoceros; currently protected in
many national parks and game reserves: Hluhluwe–iMfolozi, Kruger
(South Africa), Hwange (Zimbabwe), Etosha (Namibia), Serengeti,
Manyara (Tanzania), Masai Mara, Meru (Kenya). ♀ 5 mm

ONITINI
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Genus Kolbeellus Jacobson, 1906

Type species and designation: Tapeinopterus ateuchoides van Lansberge, 1875b, by subsequent designation (Janssens
1937).
Synonyms: = Tapeinopterus van Lansberge, 1875b: Type species: Tapeinopterus ateuchoides van
Lansberge, 1875b, by monotypy.
Last review: Entire genus reviewed by Janssens (1937).

Kolbeellus Jacobson, 1906, is a nom nov. for Tapeinopterus van Lansberge, 1875b, which is a previously occupied name for
a genus of Carabidae. Kolbeellus is monotypic, exceedingly rare, occupying a small, ill-defined range in the Eastern Cape,
South Africa, where protection of the genus is urgently required.

ONITINI
SURICATA 6 (2020) 347

Kolbeellus ateuchoides
(van Lansberge, 1875b)

Global: DD (see IUCN Red List – DD)

Endemic: RSA

J A S O N D J F M A M J

Type locality: As Tapeinopterus ateuchoides: ‘Afrique austral’ [southern

Africa].
Taxonomy: Accepted species; Kolbeellus ateuchoides (van Lansberge,
1875b) is a nom nov. for Tapeinopterus ateuchoides van Lansberge,
1875b.
Distribution: Dry upland localities, at the SE edge of the Upper Karoo,
Eastern Cape, South Africa. ♂
Locality data (mean ± SD, range): Altitude: 1 890 ± 735, 1 371–2 410 m;
annual rainfall: 429 ± 21, 414–443 mm; annual temperature (max. +
min. /2): 11.9 ± 4.2, 8.9–14.9°C (N=2).
Habitat: No quantitative assessment; unknown.
Food types: No quantitative assessment; unknown.
Temporal activity: Unknown.
Bioregions South Africa: Upper Karoo Hardeveld (NKu 2), Eastern
Upper Karoo (NKu 4) (Upper Karoo Bioregion, Nama Karoo Biome);
both localities very close to outliers of Karoo Escarpment Grassland
(Gh 1) (Dry Highveld Grassland Bioregion, Grassland Biome).
Assessment rationale: EOO = 1 395 km2; known only by the holotype
male recorded in the 19th century and a few specimens recently re-
corded in the Eastern Cape, South Africa (2004); the hiatus between
collections suggest genuine rarity, this may be related to range contrac-
tion of monogastic herbivores, such as elephant and zebra, since many
species-poor onitine genera are specialists on their coarse-fibred dung;
however, this possibility currently remains speculative; owing to pauci-
ty of data, assessed as Data Deficient (DD), but known distribution of
2 mm
< 5 000 km2 at < five known locations qualifies for Endangered status
(EN).
Conservation measures: A quantitative survey is required to determine
the EOO, AOO and ecological associations; a study at known localities
is required to determine if specialist dung type associations might be
the reason for rare occurrence in drier upland islands at SE edge of the
Upper Karoo; not currently known from any protected area.

ONITINI
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Genus Megalonitis Janssens, 1937

Type species and designation: Onitis gigas Bertolini, 1855, by subsequent designation (Janssens 1937).
Synonyms: None.
Last review: Genus created and reviewed by Janssens (1937).

Megalonitis Janssens, 1937, currently comprises two species found in dry savanna of southern Africa where they are prob-
ably associated with elephant dung. Despite the wide range, Megalonitis species are rarely recorded and then only in game
reserves, hence the assessment of Data Deficient (DD). Janssens (1937) did not see the type species of the genus (M. gigas
(Bertolini, 1855)) and notes that differences to M. bohemani (van Lansberge, 1875b) may merely represent morphological
variation associated with body size. If so, M. bohemani would become a junior synonym.

ONITINI
SURICATA 6 (2020) 349

Megalonitis bohemani
(van Lansberge, 1875b)

No synonyms
Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type locality: As Onitis bohemanni: ‘Caffrerie intérieure (N’Gami)’ [inte-

rior of SE Africa = Lake Ngami, Botswana].
Taxonomy: Accepted species; however, comparison of type material is
necessary to determine if M. bohemani might be a junior synonym of
M. gigas (Bertolini, 1855), described from Mozambique; limited sexual
dimorphism (legs, head).
Distribution: Fragmented range in dry southern African savanna: N Na- ♂
mibia, N Botswana, Zimbabwe, NE South Africa.
Locality data (mean ± SD, range): Altitude: 928 ± 241, 429–1 127 m;
annual rainfall: 528 ± 87, 414–659 mm; annual temperature (max. +
min. /2): 22.4 ± 0.6, 21.1–23.1°C (N=7).
Habitat: No quantitative assessment; DBRU collection records from sand
(1), sandy loam (2), sandy clay loam (1) in open woodland (2), grass-
land (1).
Food types: No quantitative assessment; DBRU collection records entire-
ly from monogastric herbivore dung: elephant (5).
Temporal activity: Flight activity during darkness in the summer rainy
season (Nov. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Zambezian Baikiaea Woodlands (AT0726), Zam-
bezian and Mopane Woodlands (AT0725), Kalahari Acacia-Baikiaea
Woodlands (AT0709).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 317 740 km2 (?underestimated); AOO
much smaller; entirely recorded from game reserves, presumably due 5 mm
to specialisation on the dung of elephants whose ranges are now frag-
mented and restricted; widespread, but rare, suggesting low population
density even in game parks; assessed as Data Deficient (DD) because of
limited data although may deserve a threat category despite wide range.
Conservation measures: Assessment of conservation status will only be
possible once a quantitative survey has determined its ecological associ-
ations, AOO and population density; limited available data suggest that
continuing protection of elephants in dry savanna reserves of southern
Africa is essential for the conservation of this species; protected in vari-
ous national parks and game reserves: Etosha (Namibia), Moremi (Bot­
swana), Victoria Falls, Hwange (Zimbabwe), Kruger (South Africa).

♀ 5 mm

ONITINI
350 SURICATA 6 (2020)

Genus Neonitis Péringuey, 1901

Type species and designation: Neonitis porculus (Boheman) sensu Péringuey, 1901, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Cambefort et al. (2010).

Neonitis Péringuey, 1901, was described to accommodate a single misidentified species (Neonitis porculus (Boheman) sensu
Péringuey, 1901, ≠ Gilletellus porculus (Boheman, 1857)). Gillet subsequently recognised the error and renamed the spe-
cies Neonitis rhodesiae Gillet, 1918, although Cambefort et al. (2010), thereafter, synonymised it with Neonitis nigritiae
Gillet, 1918. Two new species of ‘Neonitis’ described by Cambefort (1995) are now transferred by Cambefort et al. (2010)
to the genus, Acanthonitis Janssens, 1937, with its synonym Janssensellus Cambefort, 1975. Neonitis now comprises a sin-
gle, rarely recorded, SE African species. It has been recorded from elephant dung in South Africa whereas Cambefort et al.
(2010) record diurnal activity by the species in Tanzania.

ONITINI
SURICATA 6 (2020) 351

Neonitis nigritiae
Gillet, 1918

= Neonitis rhodesiae Gillet, 1918

= Neonitis porculus (Boheman, 1857) sensu Péringuey, 1901
Global: DD

J A S O N D J F M A M J

Type localities: N. nigritiae: ‘Afrique orientale: Ile de Wau dans le lac

Kivu’ [?Isle of Idjwi, Lake Kivu, Congo DRC]; N. rhodesiae, nom nov.
for N. porculus sensu Péringuey: ‘Southern Rhodesia (Salisbury, Mazoe)’
[Zimbabwe: Harare, Mazoe].
Taxonomy: Accepted, sexually dimorphic species (head, legs); Neonits
porculus (Boheman) sensu Péringuey (1901) was confused with Onitis
porculus Boheman, 1857 (type loc. ‘Caffraria interiore’) (now Gilletel-
lus porculus (Boheman)); Gillet (1918) renamed Neonitis porculus as
N. rhodesiae; southern African N. rhodesiae was recently (2010) syn-
onymised with East African N. nigritiae; female specimen from South
Africa illustrated here requires validation as N. rhodesiae / N. nigritiae.
Distribution: Difficult to assess owing to rarity in collections, but known
from moist savanna from SE to E Africa; type localities in E Demo-
cratic Republic of the Congo (DRC) and N Zimbabwe; also, recently
cited from Tanzania (2010); new (2009) record from NW South Africa
requires validation; citation from Free State, South Africa, also requires
validation.
Locality data (mean ± SD, range): Altitude: 1 355 ± 107, 1 204–1 448 m;
annual rainfall: 748 ± 117, 627–861 mm; annual temperature (max. +
min. /2): 18.7 ± 0.5, 18.2–19.4°C (N=4).
Habitat: No quantitative assessment; southern African savanna presum-
ably differs from tropical habitat on islands in Lake Kivu.
Food types: No quantitative assessment; unknown, but suspected to be a
monogastric dung specialist.
Temporal activity: Diurnal flight activity during the summer rainy season
2 mm
(Dec.).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Close to Waterberg in Central Bushveld (SVcb)
(Savanna Biome).
Assessment rationale: EOO = 123 785 km2 (underestimated); AOO
undoubtedly much smaller; rarity in collections probably related to
extremely low population density with specialisation to the coarse-
fibred dung of large monogastric herbivores whose ranges have now
largely contracted within the boundaries of game reserves; after revi-
sion (2010), Neonitis becomes a monotypic genus; poorly known, so
assessed as Data Deficient (DD); however, rarity and probable small
AOO suggests it deserves some level of threat category despite a large
EOO.
Conservation measures: A quantitative survey focused on game reserves
is required to determine the EOO, AOO, ecological associations and
population density; unknown whether or not it is adequately protected
in game reserves; record from Mabalingwe Game Reserve, South Africa
remains unvalidated; known older records from type localities lie out-
side of protected areas.

ONITINI
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Genus Onitis Fabricius, 1798

Type species and designation: Scarabaeus sphinx Olivier, 1789, by subsequent designation (Latreille 1810).
Synonyms: None.
Last review: All Afrotropical species reviewed by Ferreira (1978); further new Afrotropical spe-
cies described, subsequently (Houston 1983; Cambefort 1984; Davis 1986; Moretto
2010), plus reviews and additions to species groups XI (Josso 2012) and XIII (Moretto
2007).

The long-established genus, Onitis Fabricius, shows a distribution centred on the tropics and warm temperate regions of
Afro-Eurasia. It is currently divided into 19 species groups comprising 170 species. These are, essentially, endemic to the
Palaearctic (10), Oriental (24) and Afrotropical regions (134) with two shared between the Afrotropics and Palaearctic.

Eight groups and 32 species of Onitis are represented within South Africa, Botswana and Namibia. Some species show
specialisations that may influence their conservation status. In particular, associations with the dung of range-restricted
monogastric herbivore influences the AOO of about 14 species comprising some or all species in six of the eight groups.
AOO is also influenced by bias to particular soil types (at least some species of six groups) or to woody vegetation cover
(particularly Group XVII).

Onitis species are relatively large bodied and construct broods at the ends of tunnels leading from under droppings.
These may comprise single brood ovoids at the branched tips of the tunnel, separated by soil (Group XVII) or clustered
together (some Group III) (Edwards & Aschenborn 1987). Or, they may comprise brood sausages with eggs laid at
intervals, either along a single sausage (Group XVIII) or along multiple sausages clustered together at the branched tips
of the tunnel (some Group III, Group VII). This nest architecture requires large amounts of dung. Therefore, they are
found primarily in moister regions that support large herbivores dropping large amounts of dung. Few species occur
where such large herbivores do not occur naturally, such as particularly arid regions or the central Kalahari where sur-
face water is scarce.

Onitis species are macropterous flying during darkness at dusk, dusk and dawn, or dusk, night and dawn, although some
species at the more temperate northernmost and southernmost limits of their latitudinal range are day fliers (see Group
VII). Of the 32 species in southern Africa, 24 are restricted to southern Africa with only eight showing larger, more wide-
spread ranges into the tropics. In southern Africa, most species show northerly ranges centred on savanna (20) either in
the east (12), the west (5) or both (3). Others are centred in savanna and Highveld Grassland (2), Highveld Grassland (6)
or moister regions of the southwest (4).

Many Onitis species are assessed as Least Concern (LC) since they are mostly widespread in farmland and reserves (18) or
are currently well protected in reserves along with elephants and/or zebra (5). However, a number (9) are assessed as Data
Deficient (DD) since they are poorly known ecologically and have been recorded from few localities over large or very
small areas. In some cases, species assessed as DD may be threatened.

In southern Africa:

Group III: Seven species; one savanna species widespread to E Africa, one savanna and Highveld Grassland species
widespread into the SW Palaearctic region; five southern African species, either centred to the south (1),
on Highveld grasslands (3) or in savanna and Highveld Grassland (1); mostly widespread in farmland.
Group VII: Five species in a cool-adapted species group; all restricted to southern Africa; four showing cool season
activity, either southerly biased in winter rainfall (1) or mainly summer rainfall regions (2) or with rela-
tively widespread range (1); one showing summer activity centred on Highveld Grassland; diurnal flight
activity in two southernmost species.
Group IX: Two savanna species with widespread distributions to E or E and W Africa; at least one is a monogastric
dung specialist on finer-grained soils in woodland.

ONITINI
SURICATA 6 (2020) 353

Group XI: Four southern African species centred on arid to dry savanna with a probable bias to association with
dung of monogastric herbivores; bias in soil associations shown by at least two species. One further spe-
cies possibly undescribed at present.
Group XII: One southern African, dry savanna species with probable specialist associations.
Group XVII: Four savanna species; three centred in southern Africa, one distributed to E Africa; at least three associ-
ated primarily with woody habitat.
Group XVIII: Five species, four centred in arid to mostly dry savanna, one on Highveld Grassland; all associated with
dung of monogastric herbivores although two equally attracted to pads of ruminant herbivores; four spe-
cies restricted to southern Africa, including a sand specialist, one finer-grained soil specialist distributed
to E Africa.
Group XIX: Four savanna species with fragmented distributions centred on game reserves owing to association with
dung of monogastric herbivores; possibly all associated with woody vegetation; two southern African
sand specialists, two on finer-grained soils with distributions extending to E Africa.

ONITINI
354 SURICATA 6 (2020)

Onitis aeruginosus
Klug, 1855

= Onitis granulisetosus Ferreira, 1976a

Global: LC (see IUCN Red List: O. aeruginosus – DD, O. granuliseto-
sus – LC)

J A S O N D J F M A M J

Type localities: O. aeruginosus: ‘Sena’ [Sena, Zambezi Valley, Mozam-

bique]; O. granulisetosus: ‘Rhodesia : no 1310, Wankie N.P. (Nr.
Sinamatella)’ [Hwange National Park, Zimbabwe].
Taxonomy: Accepted Group XI species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
♂ Distribution: Dry to moist lowland savanna on deep sands at NE edge
of Kalahari plus Zambezi Valley, also inland sand pockets on Mozam-
bique Coastal Plain: NE Namibia, N Botswana, N Zimbabwe, S Mo-
zambique, NE South Africa.
Locality data (mean ± SD, range): Altitude: 701 ± 331, 114–1 106 m;
annual rainfall: 586 ± 93, 418–681 mm; annual temperature (max. +
min. /2): 23.0 ± 1.6, 21.1–26.7°C (N=10).
Habitat: No quantitative assessment; severely limited DBRU collection
records on sand (1) or sandy loam (1) in open shrub/woodland (1).
Food types: On deep sand in Botswana: sampled only to dung of elephant
(15) and pig (10), not dung of cattle (0), sheep (0) or carrion (chicken
livers: 0); severely limited DBRU collection records on dung of ele-
phant (1), cattle (1).
Temporal activity: Flight activity in darkness during summer rainy season
(Dec. to Mar.).
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian and Mopane
Woodlands (AT0725), Zambezian Baikiaea Woodlands (AT0726),
Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
2 mm Assessment rationale: EOO = 134 325 km2; AOO probably much small-
er due to putative soil and dung type specialisation; thought to be a
deep sand specialist in open woodland, but support currently severely
limited; limited data suggest specialisation to dung of omnivores and
large monogastric herbivores; assessed as Least Concern (LC) since
most records are from protected areas.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations and data to de-
termine the importance to population density of large monogastric her-
bivores given the greater frequency of O. aeruginosus records in reserves;
protected in Hwange, Chobe and Kruger national parks (Zimbabwe,
Botswana, South Africa).

♀
2 mm

ONITINI
SURICATA 6 (2020) 355

Onitis alexis
Klug, 1835

= Onitis aygulus Latreille, 1804

= Onitis inuus Klug, 1862
= Onitis sphinx Harold, 1871b
= Onitis africanus Gillet, 1909
= Onitis africanus var. tuberculatus Gillet, 1909
subsp. septentrionalis Balthasar, 1942
Global: LC

J A S O N D J F M A M J

Type localities: O. alexis: nom nov. for O. aygulus Latreille (pre-occupied ♂

name): ‘Depuis la Barbarie jusqu’au cap du Bonne-Espérance’ [from
North Africa almost to the Cape of Good Hope]; O. inuus: ‘von Inham-
bane als früher von Tette und Sena’ [Mozambique: Inhambane, Tete and
Sena]; O. sphinx: ‘Keren und Insaba’ [Eritrea: Keren and Ansaba]; O. af-
ricanus plus var.: ‘Koulikoro’ [Mali]; subsp. septentrionalis: Not stated.
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Arid to moist African savannas S of the Sahara: South
Africa, Namibia, Botswana, Eswatini, Zimbabwe, Angola, Zambia,
Mozambique, Tanzania, Kenya, Eritrea, Nigeria, Côte d’Ivoire, Mali;
also reported from Cameroon, Niger, Ghana, Senegal, Mauritania;
circum-Mediterranean populations of N Africa, Middle East and
S Europe (Morocco to Syria, Greece to S Spain) separated as subsp.
septentrionalis (range not shown).
Locality data (mean ± SD, range): Altitude: 981 ± 452, 0–2 199 m; an-
nual rainfall: 600 ± 206, 71–1 659 mm; annual temperature (max. +
min. /2): 19.3 ± 2.4, 11.2–25.2°C (N=574).
Habitat: In Gauteng bushveld: soil generalist with a bias to more open
vegetation, deep sand (56), sandy clay loam (84), grassland (79), open
2 mm
woodland (35), shaded thickets (26); largely supported by DBRU col-
lection records: clay (13), sandy clay loam (101), sandy loam (112),
sand (100) in crop fields (11), pasture/grassland (154), open shrub/
woodland (128), forest/thickets (7).
Food types: No quantitative assessment; DBRU collection records on
cattle (370) and horse (6) dung in farmland, also in reserves on dung
of buffalo (13), elephant (17) and zebra (2), with single records on
hooked-lipped (black) rhinoceros and hippopotamus dung.
Temporal activity: Diel flight activity in darkness (crepuscular: dusk)
(Caveney et al. 1989), primarily in the summer rainy season (Sept. to
June); in Gauteng bushveld: peak activity in the early (Oct., Nov.) and
late summer (Jan., Feb.); also tolerant of drier conditions (Sept.; May,
June); bias to colonisation of fresh dung (0–2 days old, 50%; 2–4 days,
35%; N=175); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Arid:
Kalahari Xeric Savanna (AT1309), Namibian Savanna Woodlands
(AT1316); Mesic: Southern African Bushveld (AT0717), Zambezian
and Mopane Woodlands (AT0725), Angolan Mopane Woodlands
(AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambe-
zian Baikiaea Woodlands (AT0726), Southern Miombo Woodlands
(AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bush-
veld (SVcb), Lowland (SVl), Mopane (SVmp), Sub-Escarpment ♀ 2 mm

ONITINI
356 SURICATA 6 (2020)

Onitis alexis (continued)

Savanna (SVs) (Savanna Biome); Dry Highveld Grassland (Gh), drier

units of Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland
(Gs) (Grassland Biome); Albany Thicket Biome (AT); marginal in six
other bioregions/biomes.
Assessment rationale: EOO = 9 990 325 km2 (S of Sahara, gross esti-
mate); exceptionally widespread soil generalist in more open vegetation
of both reserves and farmland where it readily colonises the dung of
both indigenous mammals and domestic livestock; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on food associations; occurs in many
reserves including Kruger National Park (South Africa), Gorongosa
National Park (Mozambique), Serengeti National Park (Tanzania),
Yankari Game Reserve (Nigeria).

ONITINI
SURICATA 6 (2020) 357

Onitis aygulus
(Fabricius, 1781)

= Scarabaeus inuus Herbst, 1789

Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus aygulus: ‘India’ [cited type locality is an error];

S. inuus: Not stated.
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Dry climate bordering arid areas in winter, bimodal and
late summer rainfall regions of SW Africa; mainly S coastal and central ♂
South Africa but also S margins of Namibia and Botswana with scat-
tered records in arid regions.
Locality data (mean ± SD, range): Altitude: 971 ± 508, 0–2 410 m;
annual rainfall: 356 ± 133, 67–741 mm; annual temperature (max. +
min. /2): 16.9 ± 2.2, 8.9–21.2°C (N=157).
Habitat: No quantitative assessment; DBRU collection records from sand
(25), sandy loam (32), sandy clay loam (22) and clay (3) on pasture or
grassland (28), scrub/shrubland (26), open woodland (6); also in fallow
crop fields (6).
Food types: No quantitative assessment; DBRU collection records mostly
from cattle dung (88) with one from buffalo dung.
Temporal activity: Diel flight activity in darkness (crepuscular: dusk/
dawn and nocturnal) (Caveney et al. 1989) in the warmer summer in
all regions (Oct. to May), irrespective of rainfall periodicity.
Ecoregions Namibia, Botswana: S Kalahari Xeric Savanna (AT1309),
Succulent Karoo (AT1322).
Bioregions South Africa: In the NW: bioregions along the boundaries
of the Savanna (Eastern Kalahari Bushveld (SVk)), Grassland (S Dry
Highveld Grassland (Gd)) and Nama Karoo (Upper Karoo (NKu))
biomes; in the S: Albany Thicket (AT) and Fynbos (F) biomes. 5 mm
Assessment rationale: EOO = 756 700 km2; widespread on different soil
types with vegetation associations reflecting its range across both dry
grassland and shrubland-dominated bioregions; DBRU collection re-
cords in Fynbos Biome (34oS 19–25oE) are all in pastures or crop fields
and suggest possible past range expansion in response to transformation
of natural shrubland vegetation; along S coast variable transformation
(AT, F: 4–80%); NW part of range used primarily for grazing of farm
livestock and little transformed (SV, NK, G: 0–4%); assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Addo Elephant National Park.

♀
5 mm

ONITINI
358 SURICATA 6 (2020)

Onitis bilobatus
Ferreira, 1976a

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘South West Africa : Opuwo 18o 03’ S, 13o 50’ E, Kaoko-
land’ [N Namibia].
Taxonomy: Accepted Group XVIII species known only by the male holo-
type; should be compared with Group XVIII relative, O. mniszechi van
Lansberge, 1886, described from Damaraland, but appears to differ.
♂ Distribution: Type locality in dry N Namibian savanna.
Locality data: Altitude: 1264 m; annual rainfall: 367 mm; annual tem-
perature (max. + min. /2): 17.0oC (N=1).
Habitat: No quantitative assessment; unknown.
Food types: No quantitative assessment; association unknown, but rela-
tives in Group XVIII have been recorded partly or entirely from dung
of monogastric herbivores.
Temporal activity: Probable flight activity in darkness in the late summer
rainy season.
Ecoregions Namibia: N Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO unknown; known by a single specimen from
a dry savanna region; currently assessed as Data Deficient (DD).
Conservation measures: A quantitative survey at the E edge of the arid
Kaokoveld and surrounding regions is required to determine the EOO,
AOO, and ecological associations and how these differ from O. mnisze-
chi; not currently known to occur in any protected area.

5 mm

ONITINI
SURICATA 6 (2020) 359

Onitis caffer
Boheman, 1857

= Onitis nicanor LeConte, 1861

Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: O. caffer: ‘Caffraria tota’ [inexact, all of SE South Africa];

O. nicanor: ‘Southern States’ [described in error as showing very rare
occurrence in southern USA].
Taxonomy: Accepted Group VII species; limited sexual dimorphism
(legs), prominence of characters varies with body size; very close relative
♂
of O. perpunctatus Balthasar, 1963, but differs in seasonality; isolated
highland population in central Namibia needs to be compared with
South African individuals when more reference material is available.
Distribution: Moister regions in E half of South Africa as well as S and
SW coastal regions; outlier population in central uplands, Namibia;
also cited from Lesotho; Mozambique citation requires validation, pos-
sibly confused with O. autumnalis Davis, 1986.
Locality data (mean ± SD, range): Altitude: 990 ± 277, 0–2 274 m; an-
nual rainfall: 585 ± 182, 120–1 047 mm; annual temperature (max. +
min. /2): 17.0 ± 2.0, 10.6–22.6°C (N=275).
Habitat: No adequate quantitative assessment; at warmer N limit of
range in Gauteng bushveld: sampled on both deep sand (35) and
sandy clay loam (18); on sand: bias to shaded thicket (29) compared to
open woodland (5) and grassland (1), but, on sandy clay loam: bias to
grassland (12) compared to open woodland (4) and shaded ticket (2);
DBRU collection records biased to finer-grained soils: sand (31), sandy
loam (60), sandy clay loam (71), clay (5), with a bias to open vege-
tation: grassland/pasture (96), scrub/shrubland (17), open woodland
(17), forest (1), also fallow or harvested crop fields (15).
Food types: No quantitative assessment; DBRU collection records pri- 2 mm
marily from cattle dung (210), but also dung of buffalo (3), horse (1),
elephant (1), rhinoceros (1) and human/cattle dung mix (2).
Temporal activity: Diel flight activity in darkness (crepuscular: dusk)
(Caveney et al. 1989); records from every month of the year, but
primarily active during cooler months in autumn, winter and spring;
seasonal bias varies regionally; on Darling Hills in cooler Western Cape,
winter rainfall region: sampled Mar. to Sept.; on warmer Gauteng
bushveld: sampled at cooler end of summer rainy season and beginning
of dry season (Apr. to June); only 6% (14/228) of DBRU collection
records during Nov. to Jan.
Ecoregions Namibia: NW Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bushveld
(SVcb), margins of Lowveld (SVl) (Savanna Biome); Dry Highveld
Grassland (Gh), Mesic Highveld Grassland (Gm), Sub-Escarpment
Grassland (Gs) (Grassland Biome); moister E Upper Nama Karoo
(NKu) (Nama Karoo Biome); Albany Thicket Biome (AT); Fynbos
Biome in areas cleared of natural shrubland and replaced with pasture
or crop fields; marginal in Succulent Karoo Biome (SK).
Assessment rationale: EOO = 576 190 km2; widespread across natural
grassland and transformed pasture vegetation, primarily in South Af-
rican farmland; records from fallow and harvested crop fields suggest
tolerance of extreme habitat transformation; presence in transformed ♀ 2 mm

ONITINI
360 SURICATA 6 (2020)

Onitis caffer (continued)

pastures and arable lands along S and SW South Africa coast may or
may not represent range expansion in response to clearance of natural
shrubland; owing to known associations and tolerance of habitat mod-
ification, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative ecological data from different climatic
regions of South Africa and central Namibia; protected in several na-
tional parks including Bontebok and Addo Elephant.

ONITINI
SURICATA 6 (2020) 361

Onitis confusus
Boheman, 1860

= Scarabaeus inuus Fabricius, 1781

Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: O. confusus: ‘juxta fluvium Svakop’ [near Swakop River,

Namibia]; S. inuus: ‘Sierra Leon’ [cited synonymy undoubtedly an
error if Sierra Leone type locality is correct].
Taxonomy: Accepted Group VII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: SW African endemic; primarily South Africa; Namibian re- ♂
cords represented by only the type locality; citations from tropical Afri-
ca are considered as errors (Sierra Leone, Kenya, Tanzania, Democratic
Republic of the Congo (DRC), Malawi, Zimbabwe, Mozambique).
Locality data (mean ± SD, range): Altitude: 603 ± 567, 0–1 536 m; an-
nual rainfall: 298 ± 126, 9–666 mm; annual temperature (max. + min.
/2): 16.6 ± 1.6, 14.0–20.3°C (N=34).
Habitat: No quantitative assessment; DBRU collection records domi-
nated by sand (18), mainly from SW coast, but also sandy clay loam
(3), sandy loam (3); primarily from low profile vegetation, either nat-
ural scrub (6), or disturbed by clearance of fynbos shrubs = pasture/
grassland (4), cleared patches (7), fallow crop fields (2); few in shrub/
woodland (3).
Food types: No quantitative assessment; DBRU collection records from
cattle (16), horse (5) and donkey (1) dung.
Temporal activity: Flight activity in darkness during autumn, winter and
spring (Mar. to Oct.), which is consistent with the seasonal rainfall
across the depicted range in the winter, bimodal and late summer rain-
fall regions of southern Africa.
Ecoregions Namibia: Cited type locality in the Namib Desert (AT1315).
Bioregions South Africa: Records scattered in various bioregions; SW of
5 mm
Savanna (SV) and Grassland (G) biomes; also Nama Karoo (NK), Al-
bany Thicket (AT), Fynbos (F) and S of Succulent Karoo (SK) biomes.
Assessment rationale: EOO = 529 675 km2; as many SW records are
from disturbed habitats, current range may or may not represent that
occupied in the past; data inadequate to determine natural associations,
but probably a soil generalist in low profile vegetation; widespread and
tolerant of disturbance.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and a survey to
better determine the EOO and AOO; possibly protected in West Coast
National Park, but most past records from sparse grassland are in areas
of the park now restored to shrubland.

♀
5 mm

ONITINI
362 SURICATA 6 (2020)

Onitis cribratus
van Lansberge, 1875b

No synonyms
Global: DD
Endemic: RSA, Botswana (?two different species)

J A S O N D J F M A M J

Type locality: ‘Caffrerie intérieure (N’Gami)’ [Lake Ngami, N Botswana].

Taxonomy: Accepted Group III species, but type locality either incorrect
or species cited as O. cribratus by Ferreira (1978) differs from that de-
scribed from two individuals (♂ + ♀) by van Lansberge (1875b), who
cites it as being very rare and larger than O. pecuarius van Lansberge,
♂ 1875b; limited sexual dimorphism (head, legs).
Distribution: Species cited as O. cribratus by Ferreira (1978) (see pho-
tograph) is hardly larger than O. pecuarius and occupies a small, wet,
highland grassland range restricted to South Africa, E of Lesotho,
where it is uncommon (red squares); Lake Ngami type locality (black
square) is climatically quite different.
Locality data (mean ± SD, range): Highland data only: average alti-
tude: 1 430 ± 131, 1 305–1 582 m; average annual rainfall: 762 ± 16,
744–775 mm; average annual temperature (max. + min. /2): 14.6 ±
0.4, 14.3–15.0°C (N=3).
Habitat: No quantitative assessment; limited DBRU collection records on
sandy loam (1), sandy clay loam (1) in grassland (1), pasture (1).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Dec. to Feb.).
Ecoregions Botswana: Type locality cited as Lake Ngami: Kalahari Acacia-
Baikiaea Woodlands (AT0709).
Bioregions South Africa: Central Sub-Escarpment grassland (Gs) (Grass-
land Biome).
2 mm
Assessment rationale: EOO = 265 km2 (grossly underestimated); high-
land individuals likely represent an undescribed species confused with
O. cribratus, which is known from only two specimens at the type lo-
cality; the highland species is uncommonly recorded on cattle dung in
grassland/pasture on farmland; although habitat transformation or loss
of particular dung types from ranges are possible influences on rarity of
both species, these remain unknown; both assessed as DD.
Conservation measures: A comparison between South African highland
individuals and the type specimens of O. cribratus is required to resolve
probable taxonomic confusion; depending on the results, a quantitative
survey of the type locality and the highlands E of Lesotho needs to
be conducted to determine EOO, AOO and ecological associations of
what is likely two different species; neither is currently known to occur
in protected areas.

♀
2 mm

ONITINI
SURICATA 6 (2020) 363

Onitis curvipes
van Lansberge, 1875b

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cap de Bonne-Espérance’ [Cape of Good Hope, South

Africa].
Taxonomy: Accepted Group XVIII species; limited sexual dimorphism
(head, legs), prominence of characters varies with body size.
Distribution: Highveld of South Africa; further records from the S coast,
one from pasture after clearance of natural shrubland, others without ♂
citations of vegetation type; records from Angola, Zimbabwe, Mozam-
bique, Botswana, Namibia, probably errors.
Locality data (mean ± SD, range): Altitude: 914 ± 621, 33–1 632 m;
annual rainfall: 458 ± 156, 256–787 mm; annual temperature (max. +
min. /2): 16.7 ± 1.3, 13.4–18.8°C (N=14).
Habitat: No quantitative assessment; single DBRU collection records
from clay, sandy clay loam, sandy loam, sand on limestone, natural
grassland and pasture.
Food types: No quantitative assessment; DBRU collection records from
horse (3), zebra (1), donkey (1) and cattle (2) dung.
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.).
Bioregions South Africa: In NW: Dry Highveld Grassland (Gd), Mesic
Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland
Biome), E Upper Karoo (NKu) (Nama Karoo Biome), Eastern Kalahari
Bushveld (SVk) (Savanna Biome); In SW: Little Karoo (NKl) (Nama
Karoo Biome), Shale Renosterveld (FRs), Sandstone Fynbos (FFs)
(Fynbos Biome).
Assessment rationale: EOO = 150 365 km2; cited as very rare in original 5 mm

description, possibly because of association with the dung of monogas-

tric herbivores at a time when population density of such indigenous
mammals was already depleted across its range; because of the wide
range, would not qualify for an IUCN threat category; poorly known
and assessed as Data Deficient (DD).
Conservation measures: Quantitative data on ecological associations is
required before an assessment may be made of conservation status, par-
ticularly, tests for specialisation to monogastric herbivore dung, which
may be the reason for continuing low population density across its
grassland/pasture range; not currently known from any protected area.

♀ 5 mm

ONITINI
364 SURICATA 6 (2020)

Onitis deceptor
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Southern Rhodesia (Enkeldorn, Buluwayo)’ [Zimbabwe:

Chivhu, Bulawayo].
Taxonomy: Accepted Group XVIII species; limited sexual dimorphism
(head, legs), prominence of characters varies with body size.
Distribution: Dry to moist savannas of southern and southern central
Africa: South Africa, Botswana, Zimbabwe, Mozambique, Angola, Ma-
♂ lawi; patchy occurrence in Kalahari outliers and other pockets of deep
sands, including SE coast; also reported from Democratic Republic of
the Congo (DRC), Tanzania, but requires validation.
Locality data (mean ± SD, range): Altitude: 961 ± 510, 0–1 698 m; an-
nual rainfall: 625 ± 219, 235–1 220 mm; annual temperature (max. +
min. /2): 19.9 ± 1.9, 16.3–23.4°C (N=62).
Habitat: Extreme deep sand specialist; in Gauteng bushveld and Bushlands
Halt: respectively, deep sand (238, 230), sandy clay loam/clay loam
(0, 4); in Gauteng bushveld: bias to woody vegetation, grassland (37),
open woodland (133), shaded thickets (68); at Bushlands Halt: bias
to open rather than shaded vegetation, grassland (189), dense shrub-
land (45).
Food types: No adequate quantitative assessment; in Botswana: sampled
from pig (1), elephant (3), cattle (1) and sheep (2) dung; DBRU collec-
tion records entirely from cattle dung (21).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.); strong bias to colonisation of fresh dung (0–1
days old, 71%; 1–2 days, 21%; N=34).
Ecoregions Botswana, Zimbabwe, Mozambique: Deep sand: Zambe-
zian and Mopane Woodlands (AT0725), Southern African Bushveld
5 mm (AT0717), Southern Miombo Woodlands (AT0719), Maputaland
Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Deep sand: W East Kalahari Bushveld (SVk),
Central Bushveld (SVcb) (Savanna Biome), N Indian Ocean Coastal
Belt Biome (CB).
Assessment rationale: EOO = 1 264 130 km2 (probably underestimated);
widespread EOO, but smaller, patchy AOO in a number of discrete
population centres owing to soil type specialisation; readily attracted to
cattle dung in farmland; population density could be markedly reduced
by clearance of woodland on sand in SVcb, but also fairly abundant in
deep sand grassland in CB; transformation: 0–2% (W SVk), 2–49%
(SVcb); assessed as Least Concern (LC).
Conservation measures: Conflicting habitat results suggest assessment of
conservation would be improved by further quantitative data on veg-
etation associations and data on dung type associations; protected in
Chobe National Park (Botswana), Tembe Elephant Park (South Africa).

5 mm
♀

ONITINI
SURICATA 6 (2020) 365

Onitis fabricii
Roth, 1851

= Onitis perplexus Boheman, 1857

= Onitis vethi van Lansberge, 1886
= Onitis obscuratus Fairmaire, 1893
Global: LC

J A S O N D J F M A M J

Type localities: O. fabricii: ‘Abyssinien’ [Ethiopia]; O. perplexus: ‘Caffraria

tota’ [all SE South Africa]; O. vethi: ‘Humpata’ [Angola]; O. obscura-
tus: ‘du Choa…sans indication de la localité précise’ [no exact locality,
Choa, Ethiopia].
Taxonomy: Accepted Group IX species; limited sexual dimorphism (legs), ♂
prominence of characters varies with body size; unless Boheman used
Caffraria in a geographically wide sense, type locality of synonym,
O. perplexus, lies outside of known modern range of O. fabricii.
Distribution: Dry to moist savanna from southern to NE Africa: N South
Africa, Zimbabwe, Mozambique, Angola, Kenya, Ethiopia; also report-
ed from Democratic Republic of the Congo (DRC).
Locality data (mean ± SD, range): Altitude: 999 ± 441, 59–1 698 m;
annual rainfall: 713 ± 174, 562–1 180 mm; annual temperature (max.
+ min. /2): 20.1 ± 2.1, 16.0–25.2°C (N=22).
Habitat: No quantitative assessment; DBRU collection records on finer-
grained soils: clay (1), sandy clay loam (1), sandy loam (3), in both
pasture (4) and open woodland (2).
Food types: No quantitative assessment; DBRU collection records on
both cattle (6) and elephant (2) dung.
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.).
Ecoregions Zimbabwe: Zambezian Baikiaea Woodlands (AT0726),
Zambezian and Mopane Woodlands (AT0725), Southern Miombo
Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa- 2 mm

vanna Biome).
Assessment rationale: EOO = 2 595 525 km2 (underestimated); ecologi-
cal data insufficient to assess conservation status of this uncommon spe-
cies; widespread occurrence suggests that it is not currently threatened
despite apparently low population density, assessed as Least Concern
(LC) rather than Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected
in Kruger (South Africa), Bicuar (Angola), Gorongosa (Mozambique)
national parks.

♀ 2 mm

ONITINI
366 SURICATA 6 (2020)

Onitis fulgidus
Klug, 1855

= Onitis klugii Harold, 1859 nom nov.

= Onitis cupreus Castelnau, 1840, sensu Harold, 1871b (error)
= Onitis aerarius Harold, 1878, sensu Péringuey, 1901 (?pars)
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: Onitis fulgidus: ‘Sena und Tette’ [central Mozambique:

Sena and Tete]; O. klugii is the unrecognised nom nov. for O. fulgidus
Klug, 1855; O. cupreus sensu Harold: ‘Keren’ [Eritrea – synonymy
undoubtedly erroneous]; O. aerarius sensu Péringuey: ‘Southern
♂ Rhodesia (Victoria Falls; Salisbury, Mazoe, Buluwayo)’ [Zimbabwe:
Victoria Falls; Harare, Mazoe, Bulawayo]. Occurs also in Nyassaland
and the Galla country’ [cited range in Zimbabwe, Malawi and Ethio-
pia suggests confusion between several species, one of which may be
O. fulgidus Klug, 1855].
Taxonomy: Accepted Group XVII species, but name pre-occupied by
O. fulgidus Castelnau, 1840, from Senegal, which is most unlikely to
be the same species unless the type locality is an error; validation of
nomenclature required; limited sexual dimorphism (legs), prominence
of characters varies with body size.
Distribution: Owing to confusion between species, not possible to de-
termine true range, but probably restricted to low-altitude, wooded
savanna of SE southern Africa; N South Africa, N Botswana, NE Na-
mibia, Zimbabwe to no further N than hot dry climates of the Zambezi
Valley in S Zambia, S Malawi and central Mozambique; reports from
Ethiopia, Kenya, Tanzania and Angola are errors.
Locality data (mean ± SD, range): Altitude: 832 ± 401, 52–1 529 m;
annual rainfall: 633 ± 124, 286–1 145 mm; annual temperature (max.
+ min. /2): 20.4 ± 2.0, 15.4–25.9°C (N=112).
Habitat: Strong bias to finer-grained soils in both Gauteng bushveld and
uMkhuze Game Reserve: sandy clay loam (80), deep sand (1) (Gauteng),
clay (2.8), sandy clay loam (21.5), deep sand (5.7) (uMkhuze);
in Gauteng bushveld: strong bias to shady woodland, thickets (46),
open woodland (28), grassland (7); all DBRU collection records from
5 mm shrub/woodland (10) on sand (5), sandy loam (5) or clay (1).
Food types: Limited data at Phalaborwa shows clear dung type bias: pig
(12), elephant (6), cattle (2); DBRU collection records from various
dung types: cattle (11), elephant (5), single records from wildebeest,
hooked-lipped (black) rhinoceros and horse dung.
Temporal activity: Diel flight activity in darkness (crepuscular: dusk)
(Caveney et al. 1989) in the summer rainy season (Nov. to Apr.); in
Gauteng bushveld: seasonal peaks in Nov. to Dec. and Feb.; attracted
to light.
Ecoregions Botswana, Zimbabwe: Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725); marginal occurrence in
two adjoining ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); marginal occurrence in two adjoining bioregions.
Assessment rationale: EOO = circa 366 000 km2; widespread, but AOO
smaller due to strong association with finer-grained soils in woodland
whose clearance would represent a threat to population density of
this species; transformation is 0–58% (SVl), 2–49% (SVcb); readily

ONITINI
SURICATA 6 (2020) 367

colonises cattle dung in farm rangeland despite an apparent bias to

other dung types; assessed as Least Concern (LC).
Conservation measures: Problems with nomenclature need to be resolved.
Conservation status would be dependent on preservation of dense
woody vegetation on finer-grained soils in SVcb and SVl; also useful ♀
to test possible effects of reduced dung type diversity on population
density in farmland; protected in Kruger National Park, Ithala Game
Reserve, Leeuwfontein and Tswaing nature reserves (South Africa).

2 mm

ONITINI
368 SURICATA 6 (2020)

Onitis inversidens
van Lansberge, 1875b

= Onitis inversus van Lansberge, 1875b

= Onitis aeruscator Gillet, 1918
Global: LC

J A S O N D J F M A M J

Type localities: O. inversidens: ‘Afrique australe’ [southern Africa] nom

nov. for O. inversus van Lansberge, a pre-occupied name; O. aeruscator:
‘Abyssinie’ [Ethiopia].
Taxonomy: Accepted Group XIX species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
♂ Distribution: Patchy distribution in hot, mostly dry, savanna regions from
SE to NE Africa according to patterns of range contraction by large
monogastric herbivores to whose dung types O. inversidens is special-
ised; Brits locality (black square) possibly represents relict population,
but occurrence needs confirmation; NE South Africa, NE Botswana,
Zimbabwe, Mozambique, Tanzania, Kenya, Ethiopia; also cited from
Malawi.
Locality data (mean ± SD, range): Average altitude: 545 ± 323,
59–1 131 m; average annual rainfall: 608 ± 143, 387–1 136 mm; av-
erage annual temperature (max. + min. /2): 22.4 ± 1.2, 19.2–25.2°C
(N=24).
Habitat: No quantitative assessment; limited DBRU collection records
from sand (1), sandy loam (3), sandy clay loam (1) in open woodland
(5), dry forest (1).
Food types: No quantitative assessment; at Phalaborwa: limited quanti-
tative data on dung of elephant (3), pig (1); DBRU collection records
entirely from coarse-fibred dung of large monogastric herbivores: ele-
phant (12), hooked-lipped (black) rhinoceros (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Mar.); attracted to light.
Ecoregions Botswana, Zimbabwe: Red squares: Zambezian and Mopane
Woodlands (AT0725).
5 mm Bioregions South Africa: Red squares: Mopane (SVmp), N Lowveld
(SVl) (Savanna Biome).
Assessment rationale: EOO = 1 704 640 km2 (underestimated); AOO
much smaller and patchy according to range contraction of monogas-
tric herbivores; all southern African, DBRU records (15) are from game
reserves or protected areas containing elephants; in addition to dung
type specialisation, there is possible further reduction in AOO due to
bias to finer-grained soils in wooded areas, but this requires testing;
however, present in various protected reserves over a very wide range;
therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations; observational
data suggests continuing protection of large monogastric herbivores
would be essential for conservation of this species; protected in various
national parks: Kruger (South Africa), Hwange (Zimbabwe), Manyara
(Tanzania), Tsavo East, Tsavo West (Kenya).

5 mm
♀

ONITINI
SURICATA 6 (2020) 369

Onitis licitus
Péringuey, 1901

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Colony (Burghersdorp)’ [Burgersdorp, Eastern Cape,

South Africa].
Taxonomy: Accepted Group VII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: Known from few localities on the S Highveld, Eastern
Cape, South Africa. ♂
Locality data (mean ± SD, range): Altitude: 1 619 ± 258, 1 436–1801 m;
annual rainfall: 528 ± 75, 475–581 mm; annual temperature (max. +
min. /2): 13.7 ± 1.6, 12.6–14.8°C (N=2).
Habitat: One DBRU collection record from sandy clay loam in grassland.
Food types: One DBRU collection record from cattle dung.
Temporal activity: Observed active by DBRU personnel in bright sun-
shine during daytime (mid-morning) in the early winter dry season
(May).
Bioregions South Africa: S Dry Highveld Grassland: Besemkaree Kop-
pies Shrubland (Gh 4); S Drakensberg Grassland: Stormberg Plateau
Grassland (Gd 3) (both Grassland Biome).
Assessment rationale: EOO = 105 km2 (underestimated); may be poorly
represented in collections due to restricted range and seasonal activity
at a time of the year when little sampling is conducted for dung beetles;
assessment of conservation status not possible owing to severely limited
information; transformation in occupied vegetation units: 3% (Gh 4),
9% (Gd 3); currently would qualify for an IUCN threat category, but
small range would be a collecting artefact; assessed as Data Deficient
(DD). 2 mm
Conservation measures: To assess conservation status, it is necessary to
conduct a survey in the highlands of the Eastern Cape during early win-
ter to determine its EOO, AOO and ecological associations; although
not currently known to be protected in any reserve, the extremely small
known range has been little transformed.

2 mm

ONITINI
370 SURICATA 6 (2020)

Onitis longitibialis
Ferreira, 1977

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Province: No 1024, Hopetown (30 km N)’ [South

Africa].
Taxonomy: Accepted Group XI species; known only by the holotype male.
Distribution: Known only from the type locality in an area of arid shrub/
grassland in central South Africa.
♂ Locality data: Altitude: 1 222 m; annual rainfall: 353 mm; annual tem-
perature (max. + min. /2): 17.6°C (N=1).
Habitat: No quantitative assessment; DBRU collection record: a single
known individual on sand (1) in pasture (1).
Food types: No quantitative assessment; DBRU collection record: one
individual collected from either cattle or horse dung.
Temporal activity: Flight activity in darkness during the late summer
rainy season (Mar.).
Bioregions South Africa: Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO unknown; represented in museums by only
the holotype specimen; as other Group XI members show a bias to
omnivore or monogastric herbivore dung, recent discovery and rari-
ty of collection may be related to dung type specialisation in an area
where monogastric herbivores are now uncommon; however, EOO and
ecological associations essentially unknown; assessed as Data Deficient
(DD).
Conservation measures: To assess conservation status, a quantitative
survey is required to determine the EOO, AOO and ecological asso-
ciations; not currently known to be protected in any reserve although
the type locality lies in an area in which the vegetation cover is little
transformed.

5 mm

ONITINI
SURICATA 6 (2020) 371

Onitis mendax
Gillet, 1918

= Onitis cerrutii Müller, 1949

= Onitis janssenii Gomes Alves, 1954
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. mendax: ‘Afrique Orientale: Sud-Ouest du Volcan

Gurui’ [SW of Gurui Volcano, Tanzania]; O. cerrutii: ‘Rhodesia meridi-
onali. Loc. class.: Gatooma’ [Zimbabwe]; O. janssenii: ‘Moçambique:...
Mutualí’ [Mozambique].
Taxonomy: Accepted Group XIX species; limited sexual dimorphism
(legs), prominence of characters varies with body size. ♂
Distribution: Widespread, but patchy occurrence in moist savanna re-
gions from southern to E Africa due to soil and dung type specialisa-
tion: N South Africa, Botswana, Namibia; Zimbabwe, Mozambique,
Tanzania, Kenya.
Locality data (mean ± SD, range): Altitude: 657 ± 449, 110–1 850 m;
annual rainfall: 628 ± 145, 374–1 042 mm; annual temperature (max.
+ min. /2): 21.5 ± 1.8, 13.4–23.2°C (N=39).
Habitat: No quantitative assessment; DBRU collection records entirely
from finer-grained soils: clay (1), sandy clay loam (7), sandy loam (6);
primarily in open shrub/woodland (8), grassland (1).
Food types: No quantitative assessment; DBRU collection records strong-
ly biased to monogastric herbivore dung: elephant (9), square-lipped
(white) rhinoceros (4); one record on ruminant herbivore dung: buf-
falo.
Temporal activity: Diel flight activity in darkness (crepuscular: dusk/
dawn and nocturnal) (Caveney et al. 1989) in the summer rainy season
(Oct. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Zambezian Baikiaea Woodlands (AT0726), Southern
Miombo Woodlands (AT0719).
5 mm
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 2 759 790 km2 (gross estimate); AOO
much smaller; range fragmented due to range contraction of mono-
gastric herbivores, particularly elephant and rhinoceros; occurrence
centred on game reserves or unproclaimed areas in which elephants still
occur on finer-grained soils (67% of available records from reserves,
n=37/55); recent records from dung of re-introduced elephant or rhi-
noceros (South Africa: Thabazimbi area, Loskop Dam Nature Reserve)
suggest low density relict populations still occurred in these areas; as-
sessed as Least Concern (LC) owing to wide range and occurrence in
many reserves.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; observations
suggest conservation status would be dependent on the continuing
protection of elephants and rhinoceros in reserves; protected in Kruger
National Park, Hluhluwe–iMfolozi Game Reserve (South Africa), Lake
Mutirikwe Game Reserve (Zimbabwe), Etosha National Park (Namib-
ia), Meru National Reserve, Masai Mara Game Reserve (Kenya).

♀
5 mm

ONITINI
372 SURICATA 6 (2020)

Onitis minutus
van Lansberge, 1875b

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘l’Afrique méridionale…..le sud de l’Afrique’ [southern

Africa].
Taxonomy: Accepted Group VII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: Lowlands in the winter and bimodal rainfall regions along
♂ the S coastal region, South Africa; also one old record from Loeriesfon-
tein to the arid N; only in Western and Eastern Cape; citations from
the old provinces of Transvaal and Natal are errors.
Locality data (mean ± SD, range): Altitude: 393 ± 353, 26–1 100 m;
annual rainfall: 386 ± 98, 148–484 mm; annual temperature (max. +
min. /2): 16.9 ± 1.1, 14.8–18.2°C (N=11).
Habitat: No quantitative assessment; DBRU collection records from tuft
grassland, scrub and shrubland in the Eastern Cape (single records),
but only in disturbed habitats in the Western Cape (grass and lucerne
pasture, fallow crop fields); recorded on sandy loam (4) with single
records on sand, sandy clay loam and clay.
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (7).
Temporal activity: In Western Cape: diurnal flight activity peaking in
early afternoon (during May at Gouda); primarily active during au-
tumn (May, June), but also recorded in spring (Oct.).
Bioregions South Africa: In the W: Shale Renosterveld (FRs) (Fynbos
Biome); in the E: Sandstone Fynbos (FFs) (Fynbos Biome), Albany
Thicket Biome (AT); in the N: Hantam Karoo (SKt 2) (Succulent
Karoo Biome) (Loeriesfontein).
2 mm Assessment rationale: EOO = 70 125 km2; widespread, but not com-
monly collected; assessment of conservation status difficult owing to
limited information; readily attracted to cattle dung in farmland and
apparently tolerant of habitat disturbance; significance, if any, of re-
gional differences in shrubland vs disturbed habitat association is un-
clear, but transformation in W: 80–90% (FRs); in E: 8–28% (FFs);
assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, it is necessary to
conduct a survey along the southern coastal lowlands of the Western
and Eastern Cape during autumn to determine its EOO, AOO and
ecological associations; also, if these differ between the Western and
Eastern Cape; not currently known to be protected in any reserve.

2 mm
♀

ONITINI
SURICATA 6 (2020) 373

Onitis mniszechi
van Lansberge, 1875b

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Damara’ [Damaraland, N Namibia].

Taxonomy: Accepted Group XVIII species; limited sexual dimorphism
(head, legs), prominence of characters varies with body size.
Distribution: Arid savanna in N Namibia, mostly in uplands; one record
on cool coastline.
Locality data (mean ± SD, range): Altitude: 1 236 ± 707, 58–1 805 m; ♂
annual rainfall: 265 ± 147, 14–391 mm; annual temperature (max. +
min. /2): 18.7 ± 2.6, 15.9–22.2°C (N=5).
Habitat: No quantitative assessment; one DBRU record from sand, one
from sandy clay loam in scrub or low shrubland.
Food types: No quantitative assessment; one DBRU record from cattle
dung, one from zebra dung.
Temporal activity: Flight activity in darkness during the late summer
rainy season (Feb. to Apr.).
Ecoregions Namibia: N Kalahari Xeric Savanna (AT1309), Namibian Sa-
vanna Woodlands (AT1316), Angolan Mopane Woodlands (AT0702);
also Namib Desert (AT1315).
Assessment rationale: EOO = 67 800 km2; fairly wide range, but rare
in collections, possibly due to specialisation on dung of monogastric
herbivores whose ranges have contracted or whose dung types are not
well represented by domestic livestock in farm rangeland resulting in
low population density; assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, it is necessary
to conduct a survey in dry upland savanna and coastal regions of N
Namibia to determine its EOO, AOO and ecological associations; sus- 5 mm
pected dung specialisation may mean presence of equids is essential for
its conservation; protected in Etosha National Park (Namibia).

♀
5 mm

ONITINI
374 SURICATA 6 (2020)

Onitis obenbergeri
Balthasar, 1942

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Transvaal, Linokana’ [Linokana, North-West Province,

South Africa].
Taxonomy: Accepted Group XVII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: Hot, dry lowlands below the E escarpment and up dry river
valleys of the Limpopo and Sabi; N South Africa, SE Zimbabwe, E
♂ Botswana.
Locality data (mean ± SD, range): Altitude: 533 ± 228, 274–1 370 m;
annual rainfall: 431 ± 104, 304–656 mm; annual temperature (max. +
min. /2): 22.3 ± 1.1, 18.1–23.6°C (N=26).
Habitat: No quantitative assessment; DBRU collection records suggest a
bias to coarse-grained soils: sandy loam (11), sand (5), in open wood-
land (15), especially mopane.
Food types: At Phalaborwa: strong bias to monogastric herbivore dung;
much lower attraction to dung of ruminant herbivores and omnivores,
elephant (196), pig (35), cattle (14); DBRU collection records: mainly
cattle dung (16), also elephant (1), donkey (2), horse (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); at Phalaborwa: sensitive to rainfall events, warm
evenings after heavy rain (128), after light rain (96), warmer evening
under dry conditions (24).
Ecoregions Botswana, Zimbabwe: Dry S Zambezian and Mopane Wood-
lands (AT0725); also margins of Southern African Bushveld (AT0717).
Bioregions South Africa: Mopane (SVmp); marginal in Central Bushveld
(SVcb), Lowveld (SVl) (Savanna Biome).
2 mm Assessment rationale: EOO = 59 050 km2; strong association with wood-
lands and monogastric herbivore dung could result in lower popula-
tion density in farmland; particularly in South Africa where its range
is centred on a bioregion that has been 1–20% transformed, mainly
by cultivation; however, not currently threatened due to widespread
persistence of woodland across a wide range; assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations and by measure-
ments of population density in response to woodland clearance and
differences in dung type diversity between reserves and farmland; pro-
tected in Kruger National Park (South Africa).

2 mm ♀

ONITINI
SURICATA 6 (2020) 375

Onitis obscurus
van Lansberge, 1886

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Humpata’ [SW Angola].

Taxonomy: Accepted Group XVII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: N Namibian plateau; moister upland areas at E edge of
Kaokoveld (Namibia); SW Bié Plateau, SW Angola.
Locality data (mean ± SD, range): Altitude: 1 299 ± 203, 831–1 627 m;
annual rainfall: 478 ± 143, 282–813 mm; annual temperature (max. + ♂
min. /2): 20.4 ± 1.7, 17.1–23.0°C (N=21).
Habitat: No quantitative assessment; DBRU collection records from finer-
grained soils: clay (2), sandy clay loam (4), sandy loam (4), primarily
in open shrub-woodland (8); one record from a harvested crop field.
Food types: No quantitative assessment; DBRU collection records from
various dung types in farmland: cattle (7), donkey (1), and reserves:
elephant (1), zebra (1).
Temporal activity: Flight activity in darkness during the late summer
rainy season (Dec. to Apr.).
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702), N Na-
mibian Savanna Woodlands (AT1316); marginal occurrence in three
other ecoregions.
Assessment rationale: EOO = 157 030 km2; AOO possibly smaller if
associated with woodland; not recently re-recorded (1974) at its type
locality, which has been widely converted to grassland; however, not
currently threatened as wooded farmland is widespread across its range
where it is attracted to the dung of domestic livestock; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
2 mm
proved by quantitative data on ecological associations; it should also be
determined if population density is adversely influenced by clearance of
woodland; protected in Etosha National Park (Namibia).

♀ 2 mm

ONITINI
376 SURICATA 6 (2020)

Onitis orthopus
van Lansberge, 1875b

= Onitis orthopus var. tanganyikensis Janssens, 1937

Global: LC

J A S O N D J F M A M J

Type localities: O. orthopus: ‘Caffrerie intérieure (N’Gami)’ [SE African

interior: Lake Ngami, Botswana]; var. tanganyikensis: ‘Tanganyika’
[Tanzania].
Taxonomy: Accepted Group XIX species; limited sexual dimorphism
(legs), prominence of characters varies with body size; taxonomic status
of var. tanganyikensis should be re-examined; assessment below only for
♂ southern African O. orthopus.
Distribution: Probably range restricted to E Angola, NE Namibia, N Bot­
swana and W Zimbabwe due to specialisation to deep sands; possibly
also northeastwards to Tanzania if var. tanganyikensis does not deserve
species status.
Locality data (mean ± SD, range): Altitude: 1 056 ± 177, 871–1 362 m;
annual rainfall: 618 ± 117, 414–813 mm; annual temperature (max. +
min. /2): 21.4 ± 1.4, 18.5–22.4°C (N=7).
Habitat: No adequate quantitative assessment; on deep sand in N Botswa-
na: primarily sampled from open woodland (92), few from open shrub­
land on lacustrine sands (4); DBRU collection records on sand (3) and
sandy loam (1) in open shrub/woodland (3) and riverine grass (1).
Food types: In Botswana: strong bias to monogastric herbivore dung:
elephant (80); also dung of pig (15); few on sheep dung (1); absent
from cattle dung (0) and carrion (chicken livers) (0); DBRU collection
records on elephant (3) and cattle (1) dung.
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov., Dec.).
Ecoregions Namibia, Botswana, Zimbabwe: Zambezian Baikiaea Wood-
lands (AT0726); margins of three adjacent woodland ecoregions.
2 mm Assessment rationale: EOO = 281 070 km2 (possibly underestimated);
measured EOO and AOO restricted to moist sands of the central
mega-Kalahari system where it shows a strong bias to elephant dung
and possibly to woodland; large parts of the region officially under
protection although elephant population density was recently radically
reduced in SE Angola, but is now on the increase; currently assessed as
Least Concern (LC) on basis of protection in Botswana and Zimbabwe.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations, particularly if there
are any effects of woodland clearance; observations and available data
suggest conservation status would be dependent on the continuing pro-
tection of elephants in woodland on deep sand; a survey is necessary to
determine its present conservation status in Bicuar National Park and
SE Angola (see notes under Platyonitis bicuariensis Ferreira, 1976); pro-
tected in Chobe (Botswana) and Hwange (Zimbabwe) national parks.

2 mm ♀

ONITINI
SURICATA 6 (2020) 377

Onitis paraconfusus
Ferreira, 1976a

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal : no 1046, KNP, Skukuza’ [Skukuza, Kruger Na-

tional Park, South Africa].
Taxonomy: Accepted Group XIX species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: Currently known only from dry, lowland savanna of the
Kruger National Park in NE South Africa, but likely also occurring on ♂
the S Mozambique Coastal Plain, possibly also SE Zimbabwe.
Locality data (mean ± SD, range): Average altitude: 377 ± 139, 278–
475 m; average annual rainfall: 578 ± 110, 500–655 mm; average
annual temperature (max. + min. /2): 22.4 ± 0.6, 21.9–22.8°C (N=2).
Habitat: No quantitative assessment; not known, but one record from
Nyandu Bush that is characterised by open woodland on deep sand.
Food types: No quantitative assessment; one DBRU collection record
from elephant dung.
Temporal activity: Flight activity during darkness in the summer rainy
season (Nov., Mar.).
Bioregions South Africa: Mopane (SVmp), Lowveld (SVl) (Savanna
Biome).
Assessment rationale: EOO = 1 950 km2 (underestimated); AOO would
be smaller than EOO if sand specialisation is supported by quantitative
data; distribution would be susceptible to elephant range contraction if
it is a coarse-dung specialist like its closest relatives; however, currently,
ecologically very poorly known and assessed as Data Deficient (DD).
Conservation measures: A quantitative survey is required to determine
the EOO, AOO and ecological associations; continuing conservation
of monogastric herbivores such as elephant and rhinoceros would be
essential for the conservation of this species if it proves to be a coarse-
dung specialist; both known localities protected in the Kruger National
Park, South Africa.

2 mm

ONITINI
378 SURICATA 6 (2020)

Onitis parainflaticollis
Ferreira, 1977

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: ‘South Africa. Orange Free State: Ficksburg, Farm Op die
Wal’ [Free State, South Africa].
Taxonomy: Accepted Group XII species; limited sexual dimorphism
(prothoracic disc, legs), prominence of characters varies with body size.
Distribution: Scattered occurrence over a wide range in South Africa,
♂ mostly on drier, NW, upland savanna, but also on moist coastal hills;
also central uplands of Namibia.
Locality data (mean ± SD, range): Altitude: 1 115 ± 497, 88–1 726 m;
annual rainfall: 631 ± 228, 314–979 mm; annual temperature (max. +
min. /2): 18.3 ± 1.2, 15.3–19.7°C (N=12).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam in pasture around a granite outcrop; possible bias to sandy
soils in open vegetation requires quantitative support.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung, but rarity may indicate specialisation to less common
dung types; two recent records from fresh dung in hyrax middens in
Namibia.
Temporal activity: Diel periodicity unknown, but, presumably, flies
during darkness like most other warm temperature Onitis species; re-
corded in the summer rainy season (Dec. to Apr.).
Ecoregions Namibia: NW Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Primarily Central Bushveld (SVcb), Eastern
Kalahari Bushveld (SVk) (Savanna Biome); also Mesic Highveld Grass-
land (Gm) (Grassland Biome) and Indian Ocean Coastal Biome (CB).
5 mm Assessment rationale: EOO = 116 085 km2; ecologically poorly known;
AOO possibly restricted due to dung specialisation, like some other
Onitis species; no observations to support specialisation to dung of mo-
nogastric herbivores whose ranges have contracted (zebra, elephants,
rhinoceros) with scattered replacement by domestic livestock (horses,
donkeys) at low population density; recent observations from hyrax
dung may suggest an alternative type of dung specialisation; rarity in
collections suggests O. parainflaticollis is uncommon across its entire
wide range; however, there are no data to support local and regional
threats; currently assessed as Data Deficient (DD).
Conservation measures: Quantitative data on ecological associations,
particularly dung type associations, is required to assess conservation
status as these may strongly influence both EOO and AOO, more so
than soil and vegetation type; protected in several nature reserves, in-
cluding Lekgalameetse, Rustenburg and Vernon Crookes.

♀
5 mm

ONITINI
SURICATA 6 (2020) 379

Onitis pecuarius
van Lansberge, 1875b

= Onitis viridulus Boheman,1857, sensu Péringuey, 1901 (pars South Africa)

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: Onitis pecuarius: ‘Cap de Bonne-Espérance, Caffrerie’

[Cape of Good Hope and SE South Africa]; O. viridulus sensu
Péringuey: ‘Natal (Durban), Cape Colony (Seymour)...’ [South Africa].
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size; synonym requires
re-examination as cited type localities in ‘...Southern Rhodesia (Victo- ♂
ria Falls, Buluwayo)’ [Zimbabwe], undoubtedly relate to one or more
species other than O. pecuarius.
Distribution: Moist E regions of the Highveld and coast of South Africa
from Limpopo through KwaZulu-Natal to the Eastern Cape; records
from Zimbabwe are erroneous.
Locality data (mean ± SD, range): Altitude: 876 ± 669, 0–3 218 m; an-
nual rainfall: 730 ± 181, 390–1 065 mm; annual temperature (max. +
min. /2): 16.7 ± 2.4, 4.8–21.5°C (N=116).
Habitat: Little quantitative data; DBRU collection records primarily on
finer-grained soils, sand (2), sandy loam (10), sandy clay loam (26),
clay (1), primarily in pasture/grassland (35), open woodland (1); pop-
ulation density greater in modified habitats at Carolina (Jan. 2001),
natural Themeda grassland (16), fallow crop fields (34), improved Ki-
kuyu pasture (68).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (53), horse dung (1).
Temporal activity: Diel flight activity in darkness (crepuscular: dusk/
dawn) (Caveney et al. 1989) during the summer rainy season (Oct. 5 mm
to May).
Bioregions South Africa: In the E: E Mesic Highveld Grassland (Gm), E
Drakensberg Grassland (Gd), N and E Sub-Escarpment Grassland (Gs)
(Grassland Biome); Indian Ocean Coastal Belt Biome (CB); in the S:
pastures in Albany Thicket (AT), Afrotemperate Forest (FO) Biomes,
Shale Renosterveld (FRs) (Fynbos Biome).
Assessment rationale: EOO = 198 180 km2; abundant in both natural
and modified grasslands; restriction of collection records to moist pas-
ture along the S coast may indicate range expansion in response to
clearance of natural woody vegetation that characterises the regional
biomes (Albany Thicket; Fynbos, Afrotemperate Forest); one observa-
tion (black square) in isolated irrigated pasture in the Western Cape;
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; however, appar-
ently resilient to vegetation transformation within an agro-ecosytem;
protected in Ithala and Hluhluwe–iMfolozi game reserves.

♀
5 mm

ONITINI
380 SURICATA 6 (2020)

Onitis perpunctatus
Balthasar, 1963c

No synonyms
Global: LC
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type locality: ‘Natal, Distr. Richmond, Indaleni’ [Indaleni, KwaZulu-

Natal, South Africa].
Taxonomy: Accepted Group VII species; limited sexual dimorphism
(legs), prominence of characters varies with body size; very closely re-
lated to O. caffer Boheman, 1857, but differs in seasonality.
♂ Distribution: Primarily open vegetation in moist uplands along Sout-
pansberg, NW edge of Highveld and edge of E escarpment in the N
descending to coastal lowlands in the S; unclear if disjunction between
E escarpment and NW edge of Highveld is real or a collection artefact;
South Africa, Eswatini.
Locality data (mean ± SD, range): Altitude: 1 268 ± 503, 7–1 958 m;
annual rainfall: 741 ± 136, 390–1 157 mm; annual temperature (max.
+ min. /2): 16.5 ± 2.0, 13.1–22.1°C (N=57).
Habitat: No quantitative soil assessment; DBRU collection records from
finer-grained soils: sandy loam (4), sandy clay loam (6); on the Wolk-
berg: primarily in forest at ± 900 m and ± 1 375 m (3.9 per sample),
few in grassland at ± 1 350 m (0.1); however, DBRU collection records
primarily from pasture (8), but also forest (1).
Food types: On the Wolkberg: slight bias to pig dung at ± 900 m to ±
1 375 m (4.8 per sample), compared to cattle dung (3.0); however,
DBRU collection records entirely from cattle dung (11).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May); the only summer-active member of Group VII
in South Africa.
Bioregions South Africa: In the N: primarily, N and E edge of Mesic
5 mm Highveld Grassland (Gm), N Sub-Escarpment Grassland (Gs) (Grass-
land Biome); in the S: primarily Albany Thicket Biome (AT); marginal
occurrence in four other bioregions.
Assessment rationale: EOO = 96 210 km2; although quantitative data
from the Wolkberg at ± 1 400 m show much greater abundance in
forest compared to grassland, nevertheless, widely collected in pasture
and grassland across a large EOO and AOO; therefore, not considered
to be threatened and assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations to resolve appar-
ent contradictions between qualitative and current quantitative data;
protected in various game and nature reserves, including Abe Bailey,
Suikerbosrand, Ithala, Hluhluwei–iMfolozi.

♀
5 mm

ONITINI
SURICATA 6 (2020) 381

Onitis picticollis
Boheman, 1857

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘juxta fluvium Limpopo’ [near Limpopo River, southern

Africa].
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Southern African savannas and grassland; N South Africa,
N Namibia, E Botswana, Zimbabwe, Mozambique; report from Tan-
zania requires validation. ♂
Locality data (mean ± SD, range): Altitude: 871 ± 534, 55–1 783 m;
annual rainfall: 657 ± 163, 298–1 136 mm; annual temperature (max.
+ min. /2): 19.7 ± 2.9, 13.8–25.2°C (N=65).
Habitat: No quantitative assessment; DBRU collection records on sand
(4), sandy loam (5), sandy clay loam (5) and clay (2) in pasture/grass-
land (5) and open shrub/woodland (11).
Food types: No quantitative assessment; DBRU collection records dom-
inated by dung of monogastric herbivores (13): elephant (9), hooked-
lipped (black) (1) and square-lipped (white) rhinoceros (2), horse (1);
fewer on dung of ruminant herbivores (7): cattle (6), buffalo (1).
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season (Oct. to May); attracted to light.
Ecoregions Namibia, Zimbabwe: Namibian Savanna Woodlands
(AT1316); Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Savanna (SV) and N of Grassland (G) biomes:
Lowland (SVl), Mopane (SVmp); also straddling the margins of Cen-
tral Bushveld (SVcb), Dry Highveld Grassland (Gh), Mesic Highveld
Grassland (Gm); also N Sub-Escarpment Grassland (Gs).
Assessment rationale: EOO = 327 800 km2 (underestimated); found
on a range of soil types in open and partially shaded vegetation types; 2 mm
however, a large number of available records are from game and nature
reserves (60%, n=44/73), presumably due to bias to dung of monogas-
tric herbivores; smaller number of records on cattle dung in farmland
may reflect lower population density; however widespread in reserves
and farmland, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; particularly
a study to determine relative population densities where monogastric
herbivores are either present (many reserves) or absent (often in farm-
land); protected in Kruger National Park (South Africa).

♀ 2 mm

ONITINI
382 SURICATA 6 (2020)

Onitis pseudosetosus
Ferreira, 1976a

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Rhodesia : no 1310, Wankie N.P. (Nr. Sinamatella)’

[Hwange National Park, Zimbabwe].
Taxonomy: Accepted Group XI species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Moist savanna along the Waterberg and N edge of High-
veld, South Africa; further scattered records in lowveld savanna of
♂ South Africa N to Zimbabwe; also cited from Zambia, Kenya.
Locality data (mean ± SD, range): Altitude: 1 063 ± 312, 226–1 448 m;
annual rainfall: 639 ± 64, 503–736 mm; annual temperature (max. +
min. /2): 19.4 ± 1.9, 17.1–23.1°C (N=18).
Habitat: No adequate quantitative assessment; in open woodland at Leeuw-
fontein Nature Reserve: sampled on loamy sand (40), sandy loam (48),
stony sandy loam (29); one DBRU collection record on sandy loam.
Food types: No quantitative assessment; sampled in relative abundance to
composite pig/cattle bait in Leeuwfontein Nature Reserve (117) where
a zebra population was present.
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Feb.).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcm), Lowveld (SVl) (Sa-
vanna Biome).
Assessment rationale: EOO = 191 355 km2 (southern Africa only; un-
derestimated); data insufficient to determine if ecological separation
from the sympatric Group XI relative, O. aeruginosus, in Hwange
National Park is related to edaphic, climatic or some other ecological
factor; assessed as Least Concern (LC) owing to wide distribution and
2 mm
protection in various reserves.
Conservation measures: Assessment of conservation status would be im-
proved by a quantitative survey to determine the full EOO and the ef-
fect on AOO of possible ecological associations, particularly those with
soil and dung type; protected in Hwange National Park (Zimbabwe),
Leeuwfontein Nature Reserve (South Africa).

2 mm ♀

ONITINI
SURICATA 6 (2020) 383

Onitis robustus
Boheman, 1857

= Onitis zambesianus Péringuey, 1896

= Onitis lunaris Kolbe, 1897
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. robustus: ‘prope fluvium Limpopo’ [close to Limpopo

River, southern Africa]; O. zambesianus: ‘Zambezia (Manica)’ [central
Mozambique]; O. lunaris: ‘Im westlichen Theile der Issango-Ebene
nordlich vom Albert-Edward See’ [in the western parts of the Sem-
liki Valley north of Lake Albert, Democratic Republic of the Congo
(DRC); Semliki = Issango river of Emin Pasha]. ♂
Taxonomy: Accepted Group IX species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: S to E to W African moist savanna: patchy occurrence, pri-
marily in game reserves due, especially, to dung type specialisation: N
South Africa, N Botswana, N Namibia, Angola, Zimbabwe, Mozam-
bique, Democratic Republic of the Congo (DRC), Tanzania, Kenya,
Côte d’Ivoire; also reported from Rwanda, Uganda.
Locality data (mean ± SD, range): Altitude: 691 ± 401, 59–1 377 m;
annual rainfall: 697 ± 235, 329–1 136 mm; annual temperature (max.
+ min. /2): 22.2 ± 1.9, 17.3–25.2°C (N=15).
Habitat: No quantitative assessment; DBRU collection records entirely
from finer-grained soils: clay (2), sandy clay loam (2), sandy loam (3),
in open shrub/woodland (6).
Food types: No quantitative assessment; DBRU collection records dom-
inated by monogastric herbivore dung: elephant (10), square-lipped
(white) rhinoceros (3); one record from ruminant herbivore dung:
buffalo.
Temporal activity: Flight activity in darkness, primarily in the summer
rainy season (Oct. to May); attracted to light. 5 mm
Ecoregions Namibia, Botswana, Zimbabwe: Primarily Angolan Mopane
Woodlands (AT0702), Zambezian and Mopane Woodlands (AT0725),
Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 3 525 050 km2 (gross estimate); AOO
much smaller due to dung type and possibly finer-grained soil and
woody vegetation specialisation; low population density across a frag-
mented but widespread range centred on game reserves owing to range
contraction of monogastric herbivores, particularly elephant and rhi-
noceros; some old records lie outside of current reserves, e.g. Christmas
Pass, E Zimbabwe; assessed as Least Concern (LC) owing to wide range
and protection in various reserves..
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data to support qualitative observations; status
probably dependent on continuing protection of elephants, rhinoceros,
and woody vegetation within reserves with finer-grained soils; tests of
population density in nearby farmland may reveal threats outside of
reserves; protected in Hluhluwe–iMfolozi Game Reserve (South Af-
rica) and various national parks: Gorongosa (Mozambique), Etosha
(Namibia), Hwange (Zimbabwe), Serengeti (Tanzania).

5 mm
♀

ONITINI
384 SURICATA 6 (2020)

Onitis setosus
van Lansberge, 1875b

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Benguela’ [SW Angola].

Taxonomy: Accepted Group XI species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Dry to moist savanna in S central Africa possibly with a W
bias: Angola, N Namibia, N Zimbabwe; report from Tanzania requires
validation.
♂ Locality data (mean ± SD, range): Average altitude: 1 135 ± 295,
683–1 504 m; average annual rainfall: 417 ± 137, 298–745 mm; av-
erage annual temperature (max. + min. /2): 19.2 ± 2.6, 15.0–22.5°C
(N=8).
Habitat: No quantitative assessment; limited DBRU collection records
from finer-grained soils: sandy clay loam (1), clay (1) in open shrub/
woodland (2).
Food types: No quantitative assessment; limited DBRU collection records
from dung of large monogastric herbivores: elephant (1), donkey (1).
Temporal activity: Flight activity in darkness during the late summer
rainy season (Jan. to Apr.).
Ecoregions Namibia: N Namibian Savanna Woodlands (AT1316), Ango-
lan Mopane Woodlands (AT0702).
Assessment rationale: EOO = 119 160 km2; AOO possibly smaller ac-
cording to coincidence of monogastric herbivores with finer-grained
soils although quantitative support required for such specialisations;
specialisation would, however, account for patchy distribution in An-
gola, N Namibia and at the moist upper S edge of the Zambezi Valley;
assessed as Least Concern (LC) on basis of wide distribution although
loss of monogastric herbivores might be detrimental to population
2 mm density.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative survey data to determine the full EOO and the
effect on AOO of ecological associations, particularly those with soil
and dung type; protected in Etosha National Park, N Namibia.

2 mm

ONITINI
SURICATA 6 (2020) 385

Onitis tortuosus
Houston, 1983

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘ERMELO T[rans]v[aa]l/ (16 km S)’ [Mpumalanga, South

Africa].
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: Highlands, E South Africa, E Zimbabwe; coastline, E South
Africa, SE Mozambique.
Locality data (mean ± SD, range): Altitude: 1 248 ± 449, 0–2 392 m; ♂
annual rainfall: 751 ± 138, 416–1 136 mm; annual temperature (max.
+ min. /2): 17.0 ± 2.3, 9.9–23.4°C (N=204).
Habitat: Little quantitative data; at Carolina: no clear effect of grassland
modification, natural Themeda grassland (3), fallow crop field (6), im-
proved Kikuyu pasture (6); DBRU collection records primarily from
finer-grained soils: clay (10), sandy clay loam (48), sandy loam (33),
sand (8) on pasture/grassland (80) or crop fields (6); few in shrubland
(3), open woodland (1), forest (1).
Food types: Little quantitative data; in the Wolkberg: dung type bias to
ruminant herbivore dung, cattle (6), rather than omnivore dung, pig
(0); DBRU collection records primarily from cattle dung (135); single
records from buffalo, rhinoceros and zebra dung;.
Temporal activity: Diel flight activity in darkness (crepuscular: dusk)
(Caveney et al. 1989) during the summer rainy season (Oct. to May)
with cool season activity on the warmer coastline (June); attracted to
light.
Ecoregions Zimbabwe: Eastern Zimbabwean Montane Forest Grassland
Mosaic (AT1006), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Dry Highveld Grassland (Gh), Mesic Highveld
5 mm
Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland Biome);
Central Bushveld (SVcb) (Savanna Biome) (grassy uplands: Waterberg,
Soutpansberg); Indian Ocean Coastal Belt Biome (CB); marginal in
three other bioregions.
Assessment rationale: EOO = 348 320 km2; widespread in dry to moist
highland grassland where it is apparently little influenced by pasture
improvement and is readily attracted to the dung of cattle; therefore,
well-adapted to an agro-ecosytem where transformation varies from:
4–60% (Gh), 6–69% (Gm), and 2–50% (Gs) in South Africa; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tected in Abe Bailey, Suikerbosrand and Loskop Dam nature reserves;
also Ithala Game Reserve.

♀
5 mm

ONITINI
386 SURICATA 6 (2020)

Onitis uncinatus
Klug, 1855

= Onitis fodiens Boheman, 1857

Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. uncinatus: ‘Sena’ [central Mozambique]; O. fodiens:

‘Caffraria tota’ [all SE South Africa].
Taxonomy: Accepted Group XVIII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: Arid and moist savanna, S to E Africa: N South Africa,
Eswatini, N Namibian plateau, N Botswana, Angola, Zimbabwe, Mo-
♂ zambique, Malawi, Tanzania, Kenya; also reported from Democratic
Republic of the Congo (DRC), Rwanda, Burundi, Uganda, Ethiopia.
Locality data (mean ± SD, range): Altitude: 1 015 ± 420, 0–1 998 m;
annual rainfall: 599 ± 185, 203–1 198 mm; annual temperature (max.
+ min. /2): 19.8 ± 2.1, 13.7–25.7°C (N=313).
Habitat: In Gauteng bushveld: extreme bias to finer-grained soils, sandy
clay loam (138), deep sand (0), in open vegetation, grassland (47), open
woodland (73), shaded thickets (18); DBRU collection records largely
supportive: clay (14), sandy clay loam (33), sandy loam (56), sand (39)
in forest (2), open shrub/woodland (85), pasture/grassland (39).
Food types: At Phalaborwa: strong bias to both ruminant and monogas-
tric herbivore compared to omnivore dung types, cattle (30), elephant
(27), pig (2); DBRU collection records supportive: cattle (174), buffalo
(6), wildebeest (2), elephant (8), rhinoceros (3), zebra (2), horse (1),
donkey (1).
Temporal activity: Diel flight activity in darkness (crepuscular: dusk/
dawn and nocturnal) (Caveney et al. 1989) in the summer rainy season
(Oct. to Apr.); in Gauteng bushveld: early summer peak (Nov. to Jan.)
tailing off into late summer (Feb. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: In the E: Southern African
2 mm
Bushveld (AT0717), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719); in the W: Angolan Mopane
Woodlands (AT0726), N Namibian Savanna Woodlands (AT1316), N
Kalahari Xeric Woodlands (AT1309).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome);
also N Sub-Escarpment Grassland (Gs) (Grassland Biome); margins of
four other bioregions.
Assessment rationale: EOO = 4 665 140 km2 (gross estimate); AOO re-
duced by soil type bias; in southern Africa, centred on the finer-grained
soils of the E and W savanna regions separated by the deep sands of the
southern Kalahari basin; extremely widespread; also readily colonises
cattle dung in open vegetation on farms; assessed as Least Concern
(LC).
Conservation measures: None required; widely distributed in farmland,
national parks and game reserves; protected in the Kruger NP (South
Africa), Lake Mutirikwe GR (Zimbabwe), Gorongosa NP (Mozam-
bique), Etosha NP (Namibia), Manyara NP (Tanzania), Masai Mara
GR (Kenya).

2 mm
♀

ONITINI
SURICATA 6 (2020) 387

Onitis viridulus
Boheman, 1857

= Onitis consanguineus, Felsche, 1911

= Onitis insulanus, Kolbe, 1914
= Onitis viridulus Boheman, 1857, var. laevissimus Gillet, 1911 (pars)
Global: LC

J A S O N D J F M A M J

Type localities: O. viridulus: ‘Caffraria tota’ [all SE South Africa]; O. consan-

guineus: ‘Harrar’ [Ethiopia]; O. insulanus: ‘Insel Wau im Kivu-See (Ruan-
da)’ [Wau Island, Lake Kivu, Rwanda]; var. laevissimus: ‘d’Afrique orien-
tale anglaise (Naivasha, Rift Valley);.....d’Abyssinie’ [Kenya, Ethiopia].
♂
Taxonomy: Accepted Group III species; limited sexual dimorphism (legs),
prominence of characters varies with body size.
Distribution: S to NE African savanna; N South Africa, Eswatini, NE
Namibia, N Botswana, Angola, Zimbabwe, Mozambique, Malawi,
Rwanda, Tanzania, Kenya, Ethiopia; also reported from Democratic
Republic of the Congo (DRC), Burundi, Uganda; report from S Arabia
requires validation; Gambia record presumably an error.
Locality data (mean ± SD, range): Altitude: 803 ± 517, 0–1 958 m; an-
nual rainfall: 715 ± 167, 377–1 275 mm; annual temperature (max. +
min. /2): 20.2 ± 2.2, 14.3–25.2°C (N=206).
Habitat: In Gauteng bushveld: bias to finer-grained soils, sandy clay loam
(27), deep sand (7) in various vegetation types, grassland (12), open
woodland (7), shaded thicket (15); supported by DBRU collection re-
cords: clay (18), sandy clay loam (25), sandy loam (28), sand (23) on
pasture/grassland (39), open shrub/woodland (43), thicket/forest (7).
Food types: No quantitative assessment; DBRU collection records from
various ruminant and monogastric dung types: cattle (93), buffalo (4),
wildebeest (1), elephant (26), hooked-lipped (black) (1) and square-
lipped (white) (5) rhinoceros, hippopotamus (1), warthog (1), zebra
(2), horse (1).
Temporal activity: Diel flight activity in darkness (crepuscular: dusk/
2 mm
dawn) (Caveney et al. 1989) in the summer rainy season (Oct. to May);
also in the cooler season at the warmer coastline (June to Aug.); bias to
colonisation of older dung (0–2 day old, 34%; 2–4 day, 53%; n=66);
attracted to light.
Ecoregions Botswana, Zimbabwe, Mozambique: Zambezian Baikiaea
Woodlands (AT0726), Southern African Bushveld (AT0717), Zam-
bezian and Mopane Woodlands (AT0725), Southern Miombo Wood-
lands (AT0719), Maputaland Coastal Forest Mosaic (AT0119), mar-
gins of two other ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); also Mopane (SVmp) (Savanna Biome), N Indian
Ocean Coastal Belt Biome (CB); marginal in three other bioregions.
Assessment rationale: EOO = 4 084 025 km2 (gross estimate); AOO
would probably be more restricted due soil type specialisation; howev-
er, extremely widespread on finer-grained soils in both savanna reserves
and farmland owing to generalist vegetation and dung type associa-
tions; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations, partic-
ularly food associations; protected in many national parks and game
reserves: Kruger, Hluhluwe–iMfolozi (South Africa), Gorongosa (Mo-
zambique), Bicuar (Angola), Manyara (Tanzania). ♀ 2 mm

ONITINI
388 SURICATA 6 (2020)

Onitis westermanni
van Lansberge, 1886

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Zambèze’, lectotype ‘Zambesi’ [inexact, southern Africa].

Taxonomy: Accepted Group XVII species; limited sexual dimorphism
(legs), prominence of characters varies with body size.
Distribution: SE and E African savanna: NE South Africa, NE Botswana,
Zimbabwe, Zambia, Tanzania, Kenya; reports from Ethiopia, Namibia
require validation.
♂ Locality data (mean ± SD, range): Altitude: 954 ± 378, 268–1 521 m;
annual rainfall: 655 ± 129, 333–1 021 mm; annual temperature (max.
+ min. /2): 20.5 ± 1.8, 16.9–25.6°C (N=48).
Habitat: No quantitative assessment; DBRU collection records on sandy
clay loam (9), sandy loam (7), and sand (5) in pasture/grassland (8),
open woodland (7), and fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records pri-
marily on cattle dung (31); also buffalo (2), elephant (1), hooked-
lipped (black) (1) and square-lipped (white) (1) rhinoceros and human
dung (1).
Temporal activity: Diel flight activity in darkness (crepuscular: dusk)
(Caveney et al. 1989) in the summer rainy season (Nov. to Apr.); at-
tracted to light.
Ecoregions Botswana, Zimbabwe: Southern Miombo Woodlands
(AT0719); also moister areas Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725); marginal in two other
ecoregions.
Bioregions South Africa: Moister, warmer areas Lowveld (SVl) (Savanna
Biome).
2 mm Assessment rationale: EOO = 944 160 km2 (gross estimate); no quan-
titative data on ecological associations, but widespread and recorded
equally in woodland and grassland, primarily on finer-grained soils,
with many records from cattle dung in farmland; in South Africa, pri-
marily restricted to a protected area in the NE lowlands; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Kruger National Park (South Africa), Lake Mutirikwe Game Reserve
(Zimbabwe), Meru National Reserve (Kenya).

♀
2 mm

ONITINI
SURICATA 6 (2020) 389

Genus Platyonitis Janssens, 1942

Type species and designation: Platyonitis oberthuri Janssens, 1942, by monotypy.
Synonyms: = Epionitis Balthasar, 1942: Type species: Epionitis tarsatus Balthasar, 1942, by mono-
typy.
Last review: Genus created by Janssens (1942); valid new species added by Krikken, (1974) and
Ferreira (1976b, plus key).

Platyonitis Janssens, 1942, is currently represented by three Afrotropical species found in moist savanna of SW Angola
plus N Namibia (1) with a disjunction to moist or dry savanna of E Africa (Kenya, N Tanzania) (2). Mozambique record
likely an error due to a mislabeled specimen (Bezděk & Sladecek 2012). All species are apparently associated with elephant
dung, which is why the single species in SW Africa is assessed as Vulnerable (VU) since its range in N Namibia remains
unprotected and that in Angola was subject to decimation of elephant populations.

ONITINI
390 SURICATA 6 (2020)

Platyonitis bicuariensis
Ferreira, 1976b

No synonyms
Global: VU (see IUCN Red List – VU)

J A S O N D J F M A M J

Type locality: ‘Angola: noo 1674 Bicuari N.P. (Nr. Camp)’ [Bicuar Na-
tional Park, Angola].
Taxonomy: Accepted species; limited sexual dimorphism (legs).
Distribution: Deep sands in open woodland; SE Angola and Ovambo-
land, N Namibia.
Locality data (mean ± SD, range): Altitude: 1 092 ± 129, 952–1 207 m;
♂ annual rainfall: 609 ± 180, 473–813 mm; annual temperature (max. +
min. /2): 22.2 ± 0.8, 21.4–23.0°C (N=3).
Habitat: No quantitative assessment; DBRU collection records on sand
(1), sandy loam (1) in shrub/woodland (1), riverine grassland (1).
Food types: No quantitative assessment; DBRU collection records only
from monogastric herbivore (elephant) dung (2).
Temporal activity: Flight activity during darkness in the summer rainy
season (Dec.).
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702), Zambe-
zian Baikiaea Woodlands (AT0726).
Assessment rationale: EOO = 88 760 km2; probably a soil and dung type
specialist as the type series of 14 specimens were all taken from deep
sands on elephant dung in 1974; however, the meagre collection data
are inconclusive; in 2007, following decimation of elephant popula-
tions during the Angolan civil war, the occurrence of elephants around
the type locality was assessed only as ‘possible’ by the IUCN although
news reports from 2013 indicate that elephants have returned; in view
of recent range contraction and fragmentation history for elephants in
both Namibia and Angola, the conservation status of P. bicuariensis is
currently assessed as Vulnerable (VU), since, despite a large EOO, it is
5 mm
known from only three localities.
Conservation measures: To determine its conservation status more pre-
cisely, a quantitative survey is required to accurately assess its EOO,
AOO, ecological associations and population density; particular atten-
tion needs to be paid to (1) the impact of possible temporary disap-
pearance of elephants from around the type locality in Bicuar National
Park and other parts of SE Angola and (2) the effects of interaction with
humans resulting in only possible or occasional elephant occurrence in
parts of Ovamboland where P. bicuariensis has been recorded.

5 mm
♀

ONITINI
SURICATA 6 (2020) 391

Genus Tropidonitis Janssens, 1937

Type species and designation: Onitis paradoxus Boheman, 1857, by subsequent designation (Janssens 1937).
Synonyms: None.
Last review: Genus created and reviewed by Janssens (1937).

Tropidonitis Janssens, 1937, is a monotypic genus associated with elephant dung showing a small EOO on coastal sands in
SE Africa where it is known only from the Maputaland Centre of Endemism. Although recently recorded from protected
parts of this range, it is assessed as Vulnerable (VU).

ONITINI
392 SURICATA 6 (2020)

Tropidonitis paradoxus
(Boheman, 1857)

No synonyms
Global: VU

J A S O N D J F M A M J

Type locality: As Onitis paradoxus: ‘Caffraria’ [SE South Africa].

Taxonomy: Accepted species; limited sexual dimorphism (legs).
Distribution: Past and present records all from the coastal Maputaland
Centre of Endemism, SE Mozambique and NE KwaZulu-Natal, South
Africa.
Locality data (mean ± SD, range): Altitude: 36 ± 21, 0–60 m; annual
♂ rainfall: 720 ± 64, 654–808 mm; annual temperature (max. + min. /2):
22.5 ± 0.1, 22.3–22.6°C (N=6).
Habitat: No quantitative assessment; occurs in an area of wooded deep
sands; one DBRU collection record in shrubland.
Food types: No quantitative assessment; one DBRU collection record on
elephant dung; in extensive sampling: not recorded in Maputo Ele-
phant Reserve using pig dung as bait, only one recorded in Sand Forest
in Sileza Nature Reserve using elephant dung as bait.
Temporal activity: Unknown; probably active in darkness in the summer
rainy season (Nov. to Feb.).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Straddling the edge of the Lowland (SVl) Biore-
gion (Savanna Biome) and the N of the Indian Ocean Coastal Belt
Biome (CB).
Assessment rationale: EOO = 2 455 km2; noted as rare by the species
author (Boheman 1857); possibly always restricted to the Maputaland
Centre of Endemism owing to specialised habitat (deep sand, wood-
land) and dung type associations (monogastric herbivore, especially el-
ephant); occupies a small range, so qualifies for Vulnerable (VU) (EOO
< 20 000 km2 and < 10 locations) or even Endangered (EN) status
5 mm (EOO < 5 000 km2 and < 5 locations).
Conservation measures: To determine its conservation status more pre-
cisely a quantitative survey is required to accurately assess its EOO,
AOO, ecological associations and population density; it is probable
that continuing protection of woodland and elephants on deep sands
in Tembe Elephant Park are essential to conserve this species.

♀
5 mm

ONITINI
394 SURICATA 6 (2020)

TRIBE ONTHOPHAGINI
Burmeister, 1846

As Onthophagidae; Type genus: Onthophagus Latreille, 1802.

Synonyms:
Alloscelides Janssens, 1946; Type genus Alloscelus Boucomont, 1923b.

Whereas morphological phylogeny indicates monophyly for the tribe Onthophagini (Philips 2016), mo-
lecular phylogeny suggests paraphyly (Monaghan et al. 2007). The tribe currently comprises 37 genera
although others are being added, primarily by separating species groups from the largest genus, Onthopha­
gus Latreille, 1802, and raising them to generic level. Many genera are endemic to the Afrotropical (24) or
Oriental regions (4). Several are shared between the (A) Afrotropical and Palaearctic regions (Euonthophagus
Balthasar, 1959); (B) Afrotropical and Oriental (Digitonthophagus Balthasar, 1959; Haroldius Boucomont,
1914); (C) Oriental and Palaearctic (1); or, the (D) Afrotropical, Oriental and Palaearctic or S Palaearctic
(Caccobius Thomson, 1859; Cleptocaccobius Cambefort, 1984; Phalops Erichson, 1847b; Proagoderus van
Lansberge, 1883); whereas one, (E) Onthophagus s. str. Latreille, 1802, is found globally in warmer regions
of the Afrotropical, Oriental, Palaearctic, Australasian, Nearctic and Neotropical regions including Mada-
gascar.

Many of the endemic genera (at least 19) and one Afrotropical/Oriental genus are suspected to be ant- or
termite-associated although evidence is limited. Genera with such putative habits mostly occupy terminal
positions on partial morphological phylogenies (Philips 2016). The remainder mainly comprise tunnelling
genera although at least three comprise or include kleptocoprid species.

A total of 16 onthophagine genera are recorded from South Africa, Botswana and Namibia as of Novem-
ber 2017, although two others have since been added (see Preamble, Dierkens et al. (2017), Roggero et
al. (2019)). Below, genera are ranked from most basal to most derived according to the morphological
phylogeny of Philips (2016) with the further insertion of one recently described genus. They reflect the
behavioural patterns shown by the entire tribe. Most are probably dominated by tunnelling habits (10:
one with some kleptocoprid species). The more terminally derived remainder are probably dominated by
kleptocoprid (2) or ant or termite associations (4: one with no species accounts prepared).

In South Africa, Botswana and Namibia (Nov. 2017):

(1) Proagoderus van Lansberge, 1883: Represented by 15 tunnelling species centred on savanna (12),
upland grassland (2) or Kalahari deep sands (1) with six specialised on monogastric herbivore dung,
four with distributions extending into the E tropics.
(2) Digitonthophagus Balthasar, 1959: Represented by three tunnelling species centred on savanna (1) or
SW Arid climate (2).
(3) Phalops Erichson, 1847b: Represented by 13 tunnelling species centred on savanna (9), Kalahari
deep sands (1) or variously across savanna, upland grassland and Karoo (3).
(4) Kurtops Roggero, Barbero & Palestrini, 2016: Represented by three species with distributions cen-
tred on Kalahari sands (2) or unknown range (1).
SURICATA 6 (2020) 395

(5) Onthophagus s. str. Latreille, 1802: Represented by aproximately 120 named, but not necessarily
validated, tunnelling and kleptocoprid species with a large, but unknown number of additional
unnamed species; 82 species in 78 accounts show large or small ranges centred on savanna (47),
upland grassland (16), savanna and upland grassland (1), Kalahari deep sands (5), Upper Karoo (3),
SW Arid late summer rainfall region (6), E forest (1), or winter and bimodal rainfall regions (7).
(6) Hamonthophagus Roggero, Dierkens, Barbero & Palestrini, 2016: Represented by three species dis-
tributed in the SW Arid late summer rainfall region (1) or savanna (2).
(7) Euonthophagus Balthasar, 1959: Represented by nine species for which only four names could be
validated with distributions centred on savanna (2), Kalahari deep sands (1) and Upper Karoo (1);
one a monogastric dung specialist.
(8) Hyalonthophagus Palestrini & Giacone, 1988: Represented by two species centred on savanna (1) or
Kalahari deep sands (1).
(9) Milichus Péringuey, 1901: Represented by two species, of which only one named savanna monogas-
tric dung specialist could be validated.
(10) Caccobius Thomson, 1859: Represented by over 20 possibly kleptocoprid species; habits mostly
based on small body size; names could be validated for only six species, mostly centred on savanna
(5) or upland grassland (1).
(11) Mimonthophagus Balthasar, 1963b: Represented by only three savanna species; one a monogastric
dung specialist.
(12) Cleptocaccobius Cambefort, 1984: Represented by ± six kleptocoprid species, for which names could
be validated for only three savanna taxa.
(13) Haroldius Boucomont, 1914: Represented by a single ant-associated species from savanna; in the re-
cent partial revision of tribal classification (Tarasov & Dimitrov 2016), this genus was classified with
‘Basal Scarabaeinae’ on molecular grounds not in the Onthophagini on morphological grounds
(Philips 2016).
(14) Caccobiomorphus Balthasar, 1964b: Monotypic, associated with ponerine ants in savanna and, pos-
sibly, also forest in the tropics.
(15) Heteroclitopus Péringuey, 1901: Represented by a single, possibly termite-associated species in savanna.

Onthophagine genera show a strong bias to greater diversity in savanna. Few onthophagine species are
regarded as threatened.

ONTHOPHAGINI
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Genus Caccobiomorphus Balthasar, 1964b

Type species and designation: Caccobiomorphus brevisetosus Balthasar, 1964b, by original designation.
Synonyms: = Megaponerophilus Ferreira, 1972: Type species: Caccobius megaponerae Brauns, 1914,
by original designation.
Last review: Reviewed as Megaponerophilus Janssens, 1949, by Cambefort (1979).

According to Branco (2011), rules of nomenclature cause the authorship of the earlier generic name, Megaponerophilus
Janssens, 1949 (name only), to shift to Ferreira, 1972 (first description). This reduces Megaponerophilus Ferreira, 1972, to
a synonym as it would post-date the description of Caccobiomorphus Balthasar, 1964b. The small-bodied genus, Caccobio­
morphus, is now monotypic as species described from different localities under various different names are considered to
be synonyms that are associated with the same widespread species of predatory ant.

ONTHOPHAGINI
SURICATA 6 (2020) 397

Caccobiomorphus megaponerae
(Brauns, 1914)

= Caccobiomorphus brevisetosus Balthasar, 1964b

= Caccobiomorphus congolanus Balthasar, 1964b
Global: LC

J A S O N D J F M A M J

Type localities: As Caccobius (Caccophilus) megaponerae: ‘Bulawayo, S.

Rhodesia’ [Bulawayo, Zimbabwe]; C. brevisetosus: ‘Ostafrika, Mons
Martii’ [untraced locality, East Africa]; C. congolanus: ‘Congo, in der
Umgebung von Stanleyville’ [environs of Kisangani, Democratic Re-
public of the Congo (DRC)].
Taxonomy: Accepted species; limited sexual dimorphism (head, protho-
racic disc).
Distribution: Centred in lower-altitude savanna, SE Africa: South Africa,
Zimbabwe, Zambia, Mozambique; one synonym in tropical forest area
of Central Africa: Democratic Republic of the Congo (DRC).
Locality data (mean ± SD, range): Altitude: 703 ± 502, 76–1 388 m;
annual rainfall: 600 ± 187, 391–961 mm; annual temperature (max. +
min. /2): 22.1 ± 2.5, 18.7–23.6°C (N=7).
Habitat: No quantitative assessment; holotype recorded from a nest of the
predatory, termitophagous ponerine ant, Megaponera analis (Latreille,
1802) that is widespread in dry to moist savanna and dry forest of
sub-Saharan Africa; also seen moving in columns of ponerine ants.
Food types: No quantitative assessment; cited as possibly feeding on the
termite prey of Megaponera brought into the nests.
Temporal activity: Diurnal ambulatory activity in ant columns during the
summer rainy season (Dec. to May).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725),
Southern African Bushveld (AT0717), Zambezian Baikiaea Woodlands
(AT0726).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 299 670 km2 (southern Africa only; un-
derestimated); putative range extends from dry savanna to rainforest at 2 mm
Kisangani; synonymy of C. congolanus supported by records of Mega­
ponera host from forest localities in Cameroon; rarity in collections
related to infrequent discovery due to highly specialised habits; assessed
as Least Concern (LC) on basis of large range.
Conservation measures: Conservation status is linked to that of its Mega­
ponera host, which is not considered at risk although the tropical dry
forest part of its range is fragmented; assessment of status would be
improved by further distributional and quantitative ecological data;
protected in Kruger National Park (South Africa).

ONTHOPHAGINI
398 SURICATA 6 (2020)

Genus Caccobius Thomson, 1859

Type species and designation: Scarabaeus schreberi Linnaeus, 1767, by original designation.
Synonyms: = Histeridium Motschulsky, 1860: Type species: Scarabaeus schreberi Linnaeus, 1767,
by monotypy.

Subgenera:

Caccobius s. str. Thomson, 1859: Type species: Scarabaeus schreberi Linnaeus, 1767, by original designation.
Cacconemus Jekel, 1872: Type species: Onthophagus castaneus Klug, 1855, by subsequent designation (Branco
2011).
Caccophilus Jekel, 1872: Type species: Caccophilus himalayanus Jekel, 1872, by subequent designation (Cam-
befort 1979: Group V).
Last review: Genus divided into 12 species groups by Cambefort (1979); Group IV = subgenus,
Tomogonus d’Orbigny, 1902, now raised to generic status; Group XI now transferred
to genus, Cleptocaccobius Cambefort, 1984; Group XII = Caccobiomorphus Balthasar,
1964b; new Afrotropical Caccobius species added after 1979 by Cambefort (1980,
1984), Walter (2014).

The long-established genus, Caccobius Thomson, 1859, currently comprises 103 small-bodied species in the Afrotropical
(58), Oriental (30) and Palaearctic regions (15). These species are currently classified within three subgenera (Caccobius
s. str.; Cacconemus Jekel, 1872; Caccophilus Jekel, 1872). They are also classified into nine groups (Cambefort 1979) after
removal of three further groups into other genera. Cambefort’s Group I is cited as Caccobius s. str. (Palaearctic/Oriental)
and his Group III as Cacconemus (currently, two Afrotropical species). However, membership of Caccophilus, as defined
from 10 species by Jekel (1872), is spread across four different groups (V (Oriental), VIII (Palaearctic/Oriental), IX
(Afrotropical/Oriental), X (Afrotropical). Furthermore, the three remaining groups (Cambefort 1979) are also classified as
Caccophilus: Group II (Palaearctic/Oriental), VI (two E African species), VII (one W African species).

Species pages are presented for only six Caccobius species classified in four different groups. They represent just a small
proportion of the Caccobius species recognised in collections from South Africa, Botswana and Namibia. Of five species
described from South Africa, only three could be matched to reference material. However, a further three species described
from elsewhere were more-or-less validated as also occurring in southern Africa. Even though > 15 other Caccobius species
were recognised in collections from South Africa, Botswana and Namibia (mainly from South Africa), none could be
matched to described species with confidence. Some are probably undescribed.

The six species for which species pages are provided are all widespread, either along the E seaboard (four) or across Africa
(two) (Group III (1), Group VI (1), Group IX (2), Group X (2)).

ONTHOPHAGINI
SURICATA 6 (2020) 399

Caccobius castaneus
(Klug, 1855)

= Caccobius medioniger Ancey, 1883

Global: DD

J A S O N D J F M A M J

Type localities: As Onthophagus castaneus ‘Sena’ [central Mozambique];

C. medioniger: ‘Abyssinie’ [inexact: Ethiopia].
Taxonomy: Accepted Group III species in subgenus, Cacconemus Jekel,
1872; species cited, here, as C. castaneus requires validation in compari-
son to Caccobius (Cacconemus) pseudolaevis d’Orbigny, 1908.
Distribution: Dry to moist savanna woodlands from S to E to NE Africa;
South Africa, Botswana, Namibia, Mozambique, Zimbabwe, Kenya;
also cited from Eritrea, Ethiopia; citation from Chad may represent
a close relative, Caccobius (Cacconemus) pseudolaevis, described from
Chad.
Locality data (mean ± SD, range): Altitude: 863 ± 390, 286–1 530 m;
annual rainfall: 494 ± 158, 305–754 mm; annual temperature (max. +
min. /2): 21.2 ± 2.2, 16.3–23.4°C (N=12).
Habitat: No quantitative assessment; recorded on clay, sandy loam and
sand at localities in Kenya; no recorded vegetation data.
Food types: No quantitative assessment; recorded on cattle, buffalo and
elephant dung at localities in Kenya.
Temporal activity: Diel flight periodicity unknown; recorded during the
summer rainy season (Nov. to Feb.).
Ecoregions Namibia, Botswana, Zimbabwe: Southern Miombo Wood-
lands (AT0719), Zambezian and Mopane Woodlands (AT0725), Ka-
lahari Acacia-Baikiaea Woodlands (AT0709), Angolan Mopane Wood-
land (AT0702), Namibian Savanna Woodlands (AT1316).
Bioregions South Africa: N Central Bushveld (SVcb), Mopane (SVmp),
N Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 2 314 560 km2; only recorded sporadical-
2 mm
ly in low numbers, but over a wide range; poorly known ecologically
with validation of taxonomic identity required; thus, assessed as Data
Deficient (DD).
Conservation measures: Assessment of conservation status would benefit
from validation of differences between C. castaneus and C. pseudolaevis;
quantitative data on ecological associations is also required to generate
an assessment; protected in various reserves, including Kruger National
Park (South Africa), Etosha National Park (Namibia), Meru National
Reserve (Kenya).

ONTHOPHAGINI
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Caccobius cavatus
d’Orbigny, 1908

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: ‘Sud du lac Tchad: mont des Niellims sur le moyen
Chari...,Kiao-Kala sur les rives de moyen Chari..., Rhodesia: Salisbury’
[mid-reaches, Chari River, Chad; Harare, Zimbabwe].
Taxonomy: Accepted Group X species in subgenus, Caccophilus Jekel,
1872; sexual dimorphism (head, prothoracic disc); prominence of
characters varies with body size.
♂ Distribution: Widespread on deep sands across dry to moist savanna of
W and southern Africa: South Africa, Zimbabwe, Botswana, NE Na-
mibia, Chad, Côte d’Ivoire; scattered records presumably a collection
artefact related to small body size; disjunction between W and S Africa
may also be a collection artefact.
Locality data (mean ± SD, range): Altitude: 539 ± 477, 0–1 065 m; an-
nual rainfall: 693 ± 202, 320–1 006 mm; annual temperature (max. +
min. /2): 21.4 ± 1.2, 17.9–23.0°C (N=26).
Habitat: In Botswana: strong bias to moister NE (3 615) compared to
arid SW (202); in uMkhuze Game Reserve: extreme bias to deep sand
(5.5) compared to sand on clay (0.4), sandy clay loam (0.0), clay (0.0)
with 70% of abundance in grassland (cited as Caccobius sp. 5); limited
DBRU collection records on sand (2) in grassland (1), shrubland (1),
forest/thicket (2).
Food types: In Botswana: mainly sampled to dung (pig: 895; elephant:
592; cattle: 521; sheep: 1 795), few to carrion (chicken livers: 14); in
Côte d’Ivoire: similar bias to human dung (133) compared to that of
cattle (58); limited DBRU collection records on cattle dung (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Mar.).
1 mm
Ecoregions Botswana, Zimbabwe: Kalahari Xeric Savanna (AT1309),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea
Woodlands (AT0726), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl) (Savanna Biome); Indian Ocean
Coastal Belt Biome (CB).
Assessment rationale: EOO = 861 245 km2 (gross estimate); AOO would
be somewhat smaller owing to soil specialisation although sandy soils
are widespread across its range; locally abundant and attracted to a
range of dung types, but data on vegetation associations insufficient to
determine effects of habitat transformation; assessed as Least Concern
(LC) on basis of wide range and generalist dung association.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on vegetation associations and possible
effects of habitat transformation; protected in Chobe National Park
(Botswana), uMkhuze Game Reserve (South Africa).

1 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 401

Caccobius ferrugineus
(Fahraeus, 1857)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Onthophagus ferrugineus: ‘prope fluvium Limpopo’ [near

Limpopo River, southern Africa].
Taxonomy: Accepted Group IX species in subgenus, Caccophilus Jekel, 1872;
sexual dimorphism (prothoracic disc), characters vary with body size.
Distribution: Widespread in warm, dry to moist, sandy savanna from S
to E to W Africa: South Africa, Mozambique, Zimbabwe, Botswana,
Namibia, Angola, Zambia, Kenya, Côte d’Ivoire; also reported from
♀
Malawi, Tanzania, Democratic Republic of the Congo (DRC), Soma-
lia, Gabon, Nigeria, Benin, Senegal.
Locality data (mean ± SD, range): Altitude: 951 ± 337, 0–1 850 m; an-
nual rainfall: 510 ± 198, 85–1 254 mm; annual temperature (max. +
min. /2): 20.4 ± 1.9, 13.4–24.6°C (N=137).
Habitat: In Gauteng bushveld: strong bias to deep sand (363) compared
to sandy clay loam (8) and to grassland (265) compared to open wood-
land (61) or shaded thicket (45); on deep sand in Botswana: extreme
bias to moister NE (4 425) compared to arid SW (79); partly supported
by DBRU collection records on sand (17), sandy loam (3), sandy clay
loam (1) in grassland (8), shrubland (3), open woodland (10), fallow
crop field (1).
Food types: On deep sand in Botswana: bias to dung of omnivores (pig,
2 033), less on dung of monogastric herbivores (elephant, 1 211) and
ruminant herbivores (cattle, 385; sheep, 853), rare on carrion (chicken
livers, 22); similar on sandy loam at Phalaborwa: dung of pig (559),
elephant (110), cattle (74); DBRU collection records reflect commonly
sampled dung types: cattle (20), wildebeest (1), elephant (4), zebra (1),
human/cattle or buffalo mix (2).
Temporal activity: Flight activity during darkness in the summer rainy sea-
son (Oct. to Apr.); in Gauteng: flies from Oct. to Feb. at 18:00–20:00 in
cooler months (Nov., Feb.) and 18:00–22:00 in warmer mid-summer
1 mm
(Jan.); at Phalaborwa: equally abundant in warm weather after light
(256) or heavy rain (210), or in hot, dry weather (266); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna
(AT1309), Namibian Savanna Woodlands (AT1316), but more abun-
dant in Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian
Baikiaea Woodlands (AT0726), Zambezian and Mopane Woodlands
(AT0725), Southern African Bushveld (AT0717), Southern Miombo
Woodlands (AT0719); marginal in two other ecoregions.
Bioregions South Africa: Mostly, W Eastern Kalahari Bushveld (SVk),
Central Bushveld (SVcb), Mopane (SVmp), N Lowveld (SVl) (Savanna
Biome).
Assessment rationale: EOO = 4 823 540 km2 (gross estimate); AOO
would be limited by soil type bias although sandy soils are widespread
across its very large range; association with open vegetation would be
an advantage as clearance of woodland would presumably not be detri-
mental to population density; despite a bias to omnivore dung, found
in abundance on various dung types in both farmland and reserves;
assessed as Least Concern (LC).
Conservation measures: None recommended; protected in various na-
tional parks including Kruger (South Africa), Chobe (Botswana).

ONTHOPHAGINI
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Caccobius histerinus
(Fahraeus, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Onthophagus histerinus: ‘juxta fluvium Gariep’ [probably

inaccurate; near Orange River, South Africa].
Taxonomy: Accepted Group VI species in subgenus, Caccophilus Jekel,
1872; material cited here as C. histerinus requires validation; howev-
er, it is assumed to be identified correctly as it is uniformly similar
in length to the type (4.5 mm) whereas a very close, South African
♀ relative is uniformly smaller (± 3 mm); also needs to be compared with
C. longipennis d’Orbigny, 1904 (type: 4.3 mm long), described from
‘Abyssinie: environs de Diré Daoua’ [Dire Dawa, Ethiopia], but range
cited as far S as Zambia; minor sexual dimorphism (head).
Distribution: Mainly lowland, sandy savanna; possibly patchily distrib-
uted along the entire E seaboard from S to NE Africa: South Africa,
NE Namibia, N Botswana, Zimbabwe, Mozambique; also cited from
Democratic Republic of the Congo (DRC), Malawi, Tanzania, Kenya,
Eritrea.
Locality data (mean ± SD, range): Altitude: 474 ± 525, 0–1 503 m; an-
nual rainfall: 696 ± 148, 436–1 006 mm; annual temperature (max. +
min. /2): 21.3 ± 1.5, 18.3–23.1°C (N=38).
Habitat: In uMkhuze Game Reserve: extreme bias to deep sand (4.1)
compared to sand over clay, sandy clay loam, clay (all: 0) (cited as
Caccobius sp. 4); small, very close relative found on sandy clay loam
(0.1) and clay (0.2) (cited as Caccobius sp. 7); no adequate quantitative
assessment on vegetation associations: found in low numbers in both
forest and grassland on deep sands of Maputo Special Reserve.
Food types: No quantitative assessment; no data.
Temporal activity: Diel flight periodicity unknown; seasonal activity in
the summer rainy season (Nov. to Feb.).
Ecoregions Botswana, Zimbabwe, Mozambique: Zambezian and Mo-
pane Woodlands (AT0725), Southern Miombo Woodlands (AT0719),
Southern African Bushveld (AT0717), Maputaland Coastal Forest
1 mm Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); N Indian Ocean Coastal Belt Biome
(CB).
Assessment rationale: EOO = 1 133 180 km2 (gross estimate); AOO
would be somewhat smaller owing to soil specialisation although moist
sandy savanna is widespread across its large EOO; therefore, currently
assessed as Least Concern (LC).
Conservation measures: Type material needs to be examined to ensure
validity and status of this species in comparison to two close relatives;
assessment of conservation status would be improved by quantitative
data on vegetation and food associations; protected in uMkhuze and
Tembe game reserves (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 403

Caccobius nigritulus
(Klug, 1855)

= Onthophagus litigiosus Péringuey, 1901

Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Onthophagus nigritulus: ‘Sena’ [central Mozambique];

O. litigiosus: ‘Mozambique (Rikatla)’ [Ricatla].
Taxonomy: Accepted Group IX species in subgenus, Caccophilus Jekel,
1872; Caccobius upembanus Frey, 1958, described from ‘Kaswabilenga’
[SE Democratic Republic of the Congo (DRC)] should be re-examined
as a valid taxon or possible further synonym.
Distribution: Widespread in dry to moist sandy savanna from southern to
E Africa: South Africa, Mozambique, Zimbabwe, Botswana, Namibia,
Angola, Tanzania, Kenya.
Locality data (mean ± SD, range): Altitude: 695 ± 424, 0–1 444 m; an-
nual rainfall: 598 ± 190, 267–1 136 mm; annual temperature (max. +
min. /2): 21.1 ± 1.7, 16.8–25.7°C (N=102).
Habitat: In Gauteng bushveld: extreme bias to deep sand (112) compared
to sandy clay loam (4) and bias to open woody savanna (79) and shaded
thicket (30) compared to grassland (7); similar soil bias in uMkhuze
Game Reserve: sand (1.4), sand on clay (0.1), sandy clay loam (0), clay
(0); on deep sand in Botswana: extreme bias to moister wooded NE
(15 656) than open arid SW (12); largely supported by DBRU collec-
tion records: sand (9), sandy loam (4), sandy clay loam (1) in grassland/
pasture (4), shrubland (2), open woodland (11), forest (2).
Food types: In Botswana: strong bias to omnivore dung (pig, 10 888)
compared to herbivore dung (elephant, 1 783; sheep, 2 200; cattle,
713) or carrion (chicken livers, 72); same bias at Phalaborwa: pig (652),
elephant (175), cattle (41); similar bias in Gauteng bushveld: pig (353),
horse (2), cattle (40), carrion (1); DBRU collection records reflect com-
mon dung types: human/cattle mix (1), elephant (12), rhinoceros (2),
zebra (1), horse (1), donkey (1), cattle (9), buffalo (2), wildebeest (1).
Temporal activity: Diurnal flight activity during the summer rainy sea-
son (Oct. to Apr.); in Gauteng bushveld: activity commencing in early 1 mm
summer (Oct.), building to a peak in mid-summer (Dec., Jan.), tailing
off to late summer (Apr.); at Phalaborwa: most active during hot, dry
weather (653), less so on warm days following heavy rainfall (sunny,
126) or light rainfall (cloudy, 89).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Rare in Ka-
lahari Xeric Savanna (AT1309); more abundant in Kalahari Acacia-
Baikiaea Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726),
Zambezian and Mopane Woodlands (AT0725), Southern Miombo
Woodlands (AT0719); also Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), Lowveld
(SVl), also very low population density in W Eastern Kalahari Bushveld
(SVk) (Savanna Biome); also N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 2 013 940 km2 (gross estimate); AOO
would be somewhat smaller owing to soil specialisation although sandy
soils are widespread across its range; despite a measured bias to omni-
vore dung, recorded in abundance on other dung types including that of
farm livestock; bias to wooded savanna suggests clearance of trees would
be detrimental to population density; however, assessed as Least Con-
cern (LC) due to widespread range and protection in various reserves.
Conservation measures: None recommended; protected and abundant in
various national parks including: Kruger (South Africa), Chobe (Bot­
swana), Hwange (Zimbabwe).

ONTHOPHAGINI
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Caccobius obtusus
(Fahraeus, 1857)

= Onthophagus mastrucatus Péringuey, 1901

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus obtusus: ‘in terra Natalensi’ [KwaZulu-

Natal, South Africa]; O. mastrucatus: ‘Cape Colony (Graham’s Town),
Natal (Durban)’ [South Africa: Eastern Cape, KwaZulu-Natal].
Taxonomy: Accepted Group X species in subgenus, Caccophilus Jekel,
1872; sexual dimorphism (head), prominence of characters varies with
body size; habitus and aedeagus of Caccobius cuspidiger d’Orbigny,
♂ 1913 (described from Kenya: ‘Afrique orientale anglaise: Kamilito dans
le Nandi au nord-est du lac Victoria-Nyanza’) are extremely close to
C. obtusus; review required to investigate possible synonymy.
Distribution: In South Africa: widespread on the moister E parts of the
Highveld plus SE and S coasts; also uplands and wet coastal plain of
central Mozambique plus uplands of Tanzania, Kenya; n.b. C. cuspidi­
ger cited from Kenya, Malawi.
Locality data (mean ± SD, range): Altitude: 1 047 ± 634, 0–2 286 m;
annual rainfall: 796 ± 208, 255–1 659 mm; annual temperature (max.
+ min. /2): 16.6 ± 2.6, 10.6–25.2°C (N=75).
Habitat: No adequate quantitative data; across a 500–2 800 m grassland
gradsect in S KwaZulu-Natal: recorded only at 1 000 m (17.0) and
1 500 m (19.9); at Carolina: marginally more abundant in improved
Kikuyu pasture (274) and fallow crop field (266) than natural Themeda
grassland (165); DBRU collection records from sand (2), sandy loam
(9), sandy clay loam (10), clay (2), primarily in grassland/pasture (20),
also open woodland (2), fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records pri-
marily from dung of cattle (23), also elephant (1), horse (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
1 mm Bioregions South Africa: Centred on N and E Mesic Highveld Grassland
(Gm), Sub-Escarpment Grassland (Gs) (Grassland Biome) plus Indian
Ocean Coastal Belt Biome (CB); also S coastal localities in Fynbos
Biome (probably disturbed and replaced by pasture).
Assessment rationale: EOO = 781 120 km2 (gross estimate); AOO small-
er, centred on moist cool uplands and moist coastal regions; recorded
at forest localities, but presumably primarily in grassland although
quantitative support is lacking; readily attracted to dung of domestic
livestock and tolerant of habitat modification in farmland; assessed as
Least Concern (LC)
Conservation measures: Assessment of conservation status would be im-
proved by review of taxonomic status and EOO plus quantitative data
on ecological associations; protected in Ithala Game Reserve, Mt Sheba
Nature Reserve (South Africa).

♀
1 mm

ONTHOPHAGINI
SURICATA 6 (2020) 405

Genus Cleptocaccobius Cambefort, 1984

Type species and designation: Caccobius (Caccophilus) signaticollis d’Orbigny, 1902, by original designation.
Synonyms: None.
Last review: Entire genus created and reviewed by Cambefort (1984); new species added by Cam-
befort (1985b), Josso and Prévost (2006b).

Cleptocaccobius Cambefort, 1984, was created by raising Group XI of Caccobius Thomson, 1859 to generic level (Cam-
befort 1979). It currently comprises 23 small-bodied species found in the Afrotropical (15) and Oriental regions (7) with
one shared between the Afrotopical and Palaearctic regions (Arabian Peninsula). Species accounts are provided for three
species recorded from South Africa, Botswana and Namibia although three other species were recognised in collections
from southern Africa that could not be named with confidence.

ONTHOPHAGINI
406 SURICATA 6 (2020)

Cleptocaccobius convexifrons
(Raffray, 1877)

= Onthophagus pudens Péringuey, 1901

= Caccobius viridicollis Fahraeus, 1857, sensu d’Orbigny, 1902
= Caccobius viridicollis Fahraeus var. picipennis d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus convexifrons: ‘Ile de Zanzibar’ [Zanzibar,

Tanzania]; O. pudens: ‘Southern Rhodesia (Salisbury)’ [Harare, Zimbab-
we]; C. viridicollis sensu d’Orbigny: Not stated; var. picipennis: ‘l’Af-
rique orientale allemande: Bagamoyo’ [Bagamoyo, Tanzania].
Taxonomy: Accepted kleptocoprid species.
Distribution: Widespread in moist sandy savanna from S to E and E to
W Africa: South Africa, Botswana, Zimbabwe, Tanzania, Kenya; also
cited from Democratic Republic of the Congo (DRC), Eritrea, Côte
d’Ivoire, Mauritania.
Locality data (mean ± SD, range): Altitude: 806 ± 360, 55–1 272 m;
annual rainfall: 622 ± 124, 283–751 mm; annual temperature (max. +
min. /2): 21.0 ± 2.0, 16.8–24.4°C (N=12).
Habitat: In Gauteng bushveld: extreme specialisation to deep sand (524)
compared to sandy clay loam (6) with a bias to more open vegetation:
grassland (201), open woodland (241), shaded thicket (88); in Botswa-
na: common in moist NE (3 203), rare in arid SW (1).
Food types: In Botswana: abundant on various dung types although
showing clear bias: sheep (1 927), pig (820), cattle (281), elephant
(142), also chicken liver carrion (33).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); in Gauteng bushveld: primarily active early in
the rainy season (Oct., Nov.) with activity continuing at lower levels
until end of Mar.
Ecoregions Botswana: Zambezian and Mopane Woodlands (AT0725),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Kalahari Xeric Savan-
na (AT1309).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome).
1 mm
Assessment rationale: EOO = 3 973 335 km2; AOO would be much
smaller owing to soil specialisation on deep sands; however, widespread
and locally abundant; low number of localities would be a collection
artefact due to small body size; assessed as Least Concern (LC).
Conservation measures: No conservation measures necessary at present;
protected in Leeuwfontein Nature Reserve (South Africa), Chobe Na-
tional Park (Botswana).

ONTHOPHAGINI
SURICATA 6 (2020) 407

Cleptocaccobius postlutatus
(d’Orbigny, 1905)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Caccobius postlutatus: ‘Afrique orientale anglaise: Nairo-

bi’ [Nairobi, Kenya].
Taxonomy: Accepted kleptocoprid species, but material cited here as
C. postlutatus requires validation; highland and lowland populations
of southern Africa indistinguishable; minor sexual dimorphism (head).
Distribution: Putative widespread range in moist lowland woodland and
highland grassland or forest, from S to E Africa: South Africa, Zimbab­
we, Mozambique, Kenya; lowland range (red squares); highland range
(black squares).
Locality data (mean ± SD, range): Altitude: 885 ± 567, 0–2 274 m; an-
nual rainfall: 724 ± 212, 356–1 157 mm; annual temperature (max. +
min. /2): 18.9 ± 3.4, 10.6–25.2°C (N=51).
Habitat: No quantitative assessment; DBRU collection records from finer-
grained soils (sandy loam: 3; sandy clay loam: 3; clay: 1) in highland
pasture (2), lowland woodland (5), plantation (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (4), elephant (4), donkey (1), human (1).
Temporal activity: Diel flight periodicity unknown; active in the summer
rainy season (Oct. to Apr.).
Ecoregions Namibia, Zimbabwe: N Namibian Savanna Woodlands
(AT1316), Zambezian and Mopane Woodlands (AT0725), Southern
Miombo Woodlands (AT0719).
Bioregions South Africa: Centred on Lowveld (SVl) (Savanna Biome);
N and E edge Mesic Highveld Grassland (Gm), N Sub-Escarpment
Grassland (Gs) (Grassland Biome); Indian Ocean Coastal Belt Biome
(CB). 1 mm
Assessment rationale: EOO = 1 627 085 km2 (gross estimate); AOO
would be smaller assuming indications of soil (finer-grained) and veg-
etation associations (lowland woodland, highland grassland) are sup-
ported by quantitative data; however, putative widespread occurrence
justifies an assessment of Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by validation of species identity, further distribution data and
quantitative data on ecological associations; lowland population pro-
tected in Kruger National Park (South Africa), highland populations in
the Cathedral Peak area of N uKhahlamba Drakensberg Park (World
Heritage Site) (South Africa).

ONTHOPHAGINI
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Cleptocaccobius viridicollis
(Fahraeus, 1857)

= Caccobius minimus d’Orbigny, 1898

= Caccobius viridicollis Fahraeus, 1857, var. semicoeruleus d’Orbigny,1905
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus viridicollis: ‘prope fluvium Gariep’ [near

Orange River, South Africa]; C. minimus: ‘Arabie: Yemen’ [Yemen];
var. semicoeruleus: ‘Afrique orientale anglaise: Tavéta, Boura, Mwataté,
Voi....Sambouru’ [E Kenya].
Taxonomy: Accepted kleptocoprid species; synonym from Yemen as-
sumed valid.
Distribution: Mostly arid to dry savanna from S to NE Africa with a syn-
onym described from S Arabia (Yemen): South Africa, Botswana, Namib-
ia, Zimbabwe, E Kenya; also cited from Mozambique, Malawi, Tanzania,
Somalia, Ethiopia, Eritrea; disjunctions may be partly a collection artefact
related to very small body size and partly due to unsuitable conditions.
Locality data (mean ± SD, range): Altitude: 971 ± 417, 0–1 830 m; an-
nual rainfall: 448 ± 208, 88–1 004 mm; annual temperature (max. +
min. /2): 19.1 ± 2.5, 13.3–24.4°C (N=134).
Habitat: In Gauteng bushveld: bias to finer-grained soils in open woody
vegetation, deep sand (603), sandy clay loam (1 628), grassland (645),
open woodland (1 403), shaded thickets (173); in uMkhuze Game Re-
serve: strong bias to coarse-grained soils, deep sand (21.4), sand on clay
(4.8), sandy clay loam (0.3), clay (0.6); partly supported by DBRU
collection records from sand (5), sandy loam (8), sandy clay loam (4) in
grassland/pasture (3), scrub/shrubland (7), open woodland (7).
Food types: At Phalaborwa: bias to dung of omnivores (pig: 758), and rumi-
nant herbivores (cattle: 328), least to monogastric herbivores (elephant:
81); in Gauteng bushveld: somewhat similar trend (pig: 21, cattle: 9,
horse: 11); DBRU collection records from dung of cattle (12), donkey (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Apr.); in Gauteng bushveld: diel peak in flight in the late morn-
ing (10:00–12:00), declining after midday (12:00–14:00) and seasonal
peak early rainy season (Oct., Nov.) becoming lower in mid-summer
(Dec., Feb.) and very low in late summer (Mar., Apr.); at Phalaborwa:
most abundant on a hot, dry day (559), remaining relatively abundant
1 mm on a warm, cloudy day following light rainfall (288) or a warm, sunny
day soon after heavy rainfall (263).
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna Wood-
lands (AT1316), Kalahari Xeric Savanna (AT1309), Kalahari Acacia-
Baikiaea Woodlands (AT0709), Zambezian and Mopane Woodlands
(AT0725), marginal occurrence in three other ecoregions.
Bioregions South Africa: Primarily Eastern Kalahari Bushveld (SVk),
Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna
Biome); also E Upper Karoo (NKu) (Nama Karoo Biome); marginal
occurrence in three other bioregions.
Assessment rationale: EOO = 1 295 810 km2 (African mainland range
only; assumed disjunct); widespread with fairly generalist ecological
associations; much of range coincides with little-transformed areas; as-
sessed as Least Concern (LC).
Conservation measures: Assessment of conservation would be improved
by further quantitative data on soil and dung type associations to re-
solve slight conflicts in existing data; protected in Kruger National Park
(South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 409

Genus Digitonthophagus Balthasar, 1959

Type species and designation: Scarabaeus bonasus Fabricius, 1775, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Génier and Moretto (2017).

Until recently, Digitonthophagus Balthasar, 1959, has been considered to comprise only two species. However, a recent
revision has determined that one of the taxa comprises a species complex. Therefore, the number of species has now been
raised to 16 found in the Afrotropical (12) and Oriental regions (2) or shared between the W Oriental and S Palaearctic
(Afghanistan) (1), or, Afrotropical and S Palaearctic (Arabian Peninsula) (1). One of the Afrotropical species is also widely
represented in Madagascar probably due to introduction from E Africa.

Three species are represented in southern Africa. One is widespread in dry to moist savannas of South Africa, Botswana
and Namibia ranging into the E tropics. Two are restricted to the W, one to the N in dry NW Namibia, W Angola, and
W Democratic Republic of the Congo (DRC), one to the S in arid W South Africa and W Namibia, overlapping with the
other species in NW Namibia and SW Angola. All three are common tunnelling species that are assessed as Least Concern
(LC).

ONTHOPHAGINI
410 SURICATA 6 (2020)

Digitonthophagus gazella
(Fabricius, 1787) auctorum

= Scarabaeus dorcas Olivier, 1789

Global: LC

J A S O N D J F M A M J

Type localities: As Scararabaeus gazella: Not stated; S. dorcas: ‘Madagas-

car’.
Taxonomy: Accepted species, but recent revision (Génier & Moretto
2017) shows that in its original sense S. gazella [=D. gazella] is a junior
synonym of Scarabaeus catta Fabricius, 1787, described from ‘Coro-
mandel’ [E India]; as the species described as S. dorcas [=Digitonthopha­
♂ gus dorcas] has been widely introduced into the Americas, Australia and
Japan under the species name gazella, a case has been made to suppress
the original affiliation of S. gazella, conserve the modern name, Digit­
onthophagus gazella, for the introduced species (ICZN case no. 3722)
and make S. dorcas [=D. dorcas] a junior synonym; sexually dimorphic
(head, disc of prothorax); prominence of characters varies with body
size.
Distribution: Widespread in grasslands and grassland savannas of S Cen-
tral and SE Africa: South Africa, Botswana, NE Namibia, Zimbabwe,
Mozambique, SW Angola, Zambia, S Democratic Republic of the
Congo (DRC), Tanzania, SE Kenya; also widespread in Madagacar
where it was probably introduced by accident about 1 000 yrs ago (Ra-
hagala et al. 2009); widely introduced intentionally into Australia, S
USA (range expanded S through central America), South America and
Japan; also recently into New Zealand.
Locality data (mean ± SD, range): Altitude: 804 ± 478, 0–1 760 m; an-
nual rainfall: 626 ± 211, 149–1 659 mm; annual temperature (max. +
min. /2): 19.5 ± 2.3, 13.5–25.2°C (N=404).
Habitat: In Gauteng bushveld: bias to finer-grained soils (sandy clay
loam, 310; deep sand, 92) and open vegetation (grassland: 302; open
woodland: 73; shaded thicket: 27); limited soil bias in uMkhuze Game
Reserve: deep sand (1.6), sand on clay (3.1), sandy clay loam (3.0), clay
(1.8); DBRU collection records on sand (25), sandy loam (19), sandy
clay loam (34), clay (16) in grassland/pasture (47), scrub/shrubland
(13), open woodland (14), shaded thicket (2), also fallow crop fields
2 mm
(3).
Food types: On deep sand in Botswana: bias to herbivore dung (elephant,
25; cattle, 21; sheep, 48) compared to omnivore dung (pig, 7) and
carrion (chicken livers, 0); but on sandy loam at Phalaborwa: slight bias
to omnivore dung (pig, 182) compared to herbivore dung: monogastric
(elephant, 144), ruminant (cattle, 76); DBRU collection records on
dung of cattle (101), buffalo (2), elephant (5), zebra (4), horse (4),
donkey (1), human (1), human/buffalo mix (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); also Aug., May, mostly at warmer coastline; in
Gauteng bushveld: active early to late summer (Oct. to Feb.) tailing off
in Mar., rare in Apr.; at Phalaborwa: most active on a warm evening
soon after heavy rainfall (232), less so on warm evenings after cloudy
weather with light rain (89) or hot, dry weather (80); attracted to light.
Ecoregions Botswana, Zimbabwe, Mozambique: SE Kalahari Xeric
Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands (AT0709),
Zambezian and Mopane Woodlands (AT0725), Southern African
Bushveld (AT0717), Southern Miombo Woodlands (AT0719),

ONTHOPHAGINI
SURICATA 6 (2020) 411

Maputaland Coastal Forest Mosaic (AT0119), marginal records in one

other ecoregion.
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk), Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); Dry Highveld Grassland (Gh),
Sub-Escarpment Grassland (Gs) (Grassland Biome); Indian Ocean
Coastal Belt Biome (CB), Albany Thicket Biome (AT); occurrence in ♀
Fynbos Biome possibly due to range expansion in response to clearance
of natural vegetation.
Assessment rationale: EOO = 3 711 400 km2 (African mainland only);
widespread in open vegetation; apparent slight bias to finer-grained
soils and possibly herbivore dung; widely introduced into other conti-
nents; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data to resolve slight conflicts in soil
and dung type association; protected in many national parks, including
Kruger (South Africa), Gorongosa (Mozambique).

2 mm

ONTHOPHAGINI
412 SURICATA 6 (2020)

Digitonthophagus namaquensis
Génier, 2017

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Farm Klipkoppies (6) (28.87137 S 21.30944 E), 957 m,

Northern Cape, South Africa’.
Taxonomy: Accepted species; sexually dimorphic (head, disc of protho-
rax); prominence of characters varies with body size.
Distribution: Arid late summer rainfall region of SW Africa: South Afri-
ca, Namibia, SW Angola.
♂ Locality data (mean ± SD, range): Altitude: 984 ± 261, 249–1 730 m;
annual rainfall: 215 ± 89, 22–424 mm; annual temperature (max. +
min. /2): 18.3 ± 2.1, 14.9–22.2°C (N=69).
Habitat: No quantitative assessment; DBRU collection records from sand
(6), sandy loam (2), sandy clay loam (3) in scrub (9), open woodland
(2).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (5), buffalo (1), wildebeest (1), zebra (2), horse (1), don-
key (1).
Temporal activity: Flight activity in darkness during the late summer
rainy season (Dec. to May), attracted to light.
Ecoregions Namibia: Succulent Karoo (AT1322), Namib Desert
(AT1315), Namibian Savanna Woodlands (AT1316), SW Kalahari
Xeric Savanna (AT1309), Angolan Mopane Woodlands (AT0702).
Bioregions South Africa: Kalahari Duneveld (SVkd) (Savanna Biome);
Bushmanland (NKb) (Nama Karoo Biome); Namaqualand Hardeveld
(SKn), Trans-Escarpment Succulent Karoo (SKt), Rainshadow Valley
Karoo (SKv) (Succulent Karoo Biome); Albany Thicket Biome (AT).
Assessment rationale: EOO = 795 870 km2; widespread in little-
transformed arid regions of SW Africa used primarily for grazing of
domestic livestock or conservation of indigenous fauna; not, therefore,
facing any threats; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
2 mm
proved by quantitative ecological data; protected in Kgalagadi Trans-
frontier Park (South Africa), Etosha National Park (Namibia).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 413

Digitonthophagus viridicollis
Génier, 2017

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Moanda, Zaïre’ [Moanda, W Democratic Republic of the

Congo (DRC)].
Taxonomy: Accepted species although Génier and Moretto (2017) note
that separation from D. gazella (Fabricius, 1787) auctorum at species
level may not be justified as differences are geographically consistent,
but limited; sexually dimorphic (head, disc of prothorax); prominence
of characters varies with body size. ♂
Distribution: Dry savanna, W extremes of South central Africa: N Na-
mibia, W Angola, W margins of Democratic Republic of the Congo
(DRC).
Locality data (mean ± SD, range): Altitude: 1 222 ± 300, 683–1 998 m;
annual rainfall: 390 ± 117, 125–623 mm; annual temperature (max. +
min. /2): 19.6 ± 2.2, 16.3–23.0°C (N=29).
Habitat: No quantitative assessment; DBRU collection records from sand
(2), sandy loam (2), sandy clay loam (3) in grassland (2), shrubland (2),
open woodland (3).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (7).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.), attracted to light.
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316), Angolan
Mopane Woodlands (AT0702), W Kalahari Xeric Savanna (AT1309),
W Kalahari Acacia-Baikiaea Woodlands (AT0709), W Zambezian
Baikiaea Woodlands (AT0726).
Assessment rationale: EOO = 419 910 km2; widespread; possibly, gen-
eralist soil and vegetation associations; recorded on dung of domestic
livestock; probably not threatened by clearance of woodland, but quan- 2 mm

titative data required in support; currently assessed as Least Concern

(LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative ecological data; protected in Etosha National
Park (Namibia).

♀ 2 mm

ONTHOPHAGINI
414 SURICATA 6 (2020)

Genus Euonthophagus Balthasar, 1959

Type species and designation: Scarabaaeus amyntas Olivier, 1789, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Balthasar (1959); valid new Afrotropical species added
by Cambefort (1972).

Euonthophagus Balthasar, 1959, was created by raising Group 8 (d’Orbigny 1913) of Onthophagus Latreille, 1802, to ge-
neric level. It currently comprises 36 valid species in the Palaearctic (20) and Afrotropical regions (16). As the differences
between species are often subtle, some Afrotropical species undoubtedly remain undescribed whereas others have been
wrongly synonymised. A revision of the Afrotropical species is necessary.

Nine Euonthophagus species were recognised in collections from South Africa, Botswana and Namibia. Five of these species
remain unnamed and it is unclear which, if any, remain undescribed. It was not possible to match them to existing species
names although two have been described from South Africa in addition to those recognised below in species accounts, one
valid, one a synonym.

Species accounts are presented for only four species of which only two were described from South Africa. Of the four
species, three are valid whereas one is considered valid, but officially remains a synonym. These four species are centred on
the Upper Karoo, SW Kalahari, southern savanna, E savanna ranging into the E tropics. All are locally abundant and none
is considered threatened although one has been recorded only on monograstic herbivore dung in game reserves.

ONTHOPHAGINI
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Euonthophagus carbonarius
(Klug, 1855)

= Onthophagus aterrimus Gerstaecker, 1871

= Onthophagus interstitialis Fahraeus, 1857, sensu Gestro, 1895
= Onthophagus glaber Boheman, 1857 (nec Fahraeus), sensu Péringuey, 1901
= Onthophagus vicarius Péringuey, 1901 (synonymised in error)
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus carbonarius: ‘Sena’ [central Mozam-

bique]; O. aterrimus: ‘See Jipe und dem Bura-Bergen’ [Kenya: Lake
Jipe and Bura Mts]; O. interstitialis sensu Gestro: ‘Ganale Guddá’ [Ethi-
opia]; O. glaber sensu Péringuey: ‘Transvaal (Rustenburg)’ [North-West
Province, South Africa]; O. vicarius: ‘Cape Colony (Beaufort West)’
[Beaufort West, Western Cape, South Africa].
Taxonomy: Accepted species, but extensive history of confusion with very
close relatives; minor sexual dimorphism, varying with body size (head,
prothoracic disc); synonyms require validation as all have likely been
synonymised in error; O. vicarius synonymy certainly an error (see
species account: Euonthophagus vicarius (Péringuey, 1901)); citation
of ‘Cape Colony (Griqualand West)’ [central Northern Cape, South
Africa] as a second locality for O. glaber sensu Péringuey may represent
confusion with E. vicarius.
Distribution: Centred on moist, sandy, lowland savanna, S Central and
SE Africa: NE South Africa, E Botswana, S Angola, S Zambia, S and
N Zimbabwe, central and S Mozambique; following records are all er-
rors: Arabian Peninsula, Mauritania, Senegal, Sudan, Ethiopia, Eritrea,
Kenya, Tanzania and Cape Province of South Africa.
Locality data (mean ± SD, range): Altitude: 627 ± 417, 11–1 899 m;
annual rainfall: 619 ± 146, 304–1 121 mm; annual temperature (max.
+ min. /2): 21.3 ± 1.9, 13.1–25.7°C (N=115).
Habitat: No adequate quantitative assessment; DBRU collection records
suggest a bias to sandy soils: sand (35), sandy loam (21), sandy clay
loam (11), clay (1) in woodland: grassland/pasture (6), shrubland (9),
open woodland (42), forest/thickets (6). 2 mm
Food types: At Phalaborwa: primarily attracted to omnivore (pig: 3 930) or mo-
nogastric herbivore dung (elephant: 2 697), less to ruminant herbivore dung
(cattle: 625); DBRU collection records from various dung types: human or
human cattle mix (6), elephant (18), rhinoceros (5), horse (2), donkey (1),
zebra (3), warthog (1), cattle (33), buffalo (3), wildebeest (1), waterbuck (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season, but recorded in all months except Sept. reflecting its lowland centre of distribution; attracted to light; at
Phalaborwa: primarily active on evenings after a hot, dry day (3 954), less on an evening following a warm cloudy day with
light rainfall (2 319) or an evening following a warm day a few days after heavy rainfall (1 028).
Ecoregions Botswana, Zimbabwe, Mozambique: E Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian & Mopane
Woodlands (AT0725), Southern African Bushveld (AT0717), S Southern Miombo Woodlands (AT0719), Maputaland
Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 946 605 km2 (gross estimate); widespread EOO, but AOO may be restricted by soil, vegetation
and dung type specialisation so that clearance of woodland on sand in farmland may be detrimental to population density;
however, locally abundant, particularly in game reserves where 64% (68 out of 106) of DBRU records were made; observed
flying in large numbers to elephant dung at dusk in Bicuar National Park (Angola); assessed as Least Concern (LC) owing to
wide range and widespread protection.
Conservation measures: Assessment of conservation status would be improved by quantitative data on habitat associations;
protected in various national parks and game reserves: Kruger, uMkhuze, Hluhluwe–iMfolozi (South Africa); Hwange (Zim-
babwe), Bicuar (Angola), Moremi (Botswana).

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Euonthophagus flavimargo
(d’Orbigny, 1902)

No synonyms
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: as Onthophagus flavimargo: ‘Natal’ [KwaZulu-Natal, South

Africa].
Taxonomy: Accepted species, but cited type locality probably an error as
it lies far outside of the known range of the species.
Distribution: Deep sands of dry to arid SW Kalahari: Botswana, South
Africa.
Locality data (mean ± SD, range): Altitude: 1 080 ± 130, 764–1 467 m;
annual rainfall: 264 ± 60, 142–419 mm; annual temperature (max. +
min. /2): 19.3 ± 0.7, 17.6–21.2°C (N=114).
Habitat: No adequate quantitative data; in Northern Cape: high fre-
quency in SW Kalahari, mainly on deep sands (86.7%, 85 out of 98
samples), low frequency in uplands to S of Orange River (5.8%, 7/121)
and in NE Bushmanland (13.4%, 9/67); DBRU collection records al-
most exclusively on sand (12), also sandy clay loam (1) in grassland (5),
scrub/shrubland (6), open woodland (2).
Food types: In Botswana: bias to dung of omnivores (pig: 71) and mo-
nogastric herbivores (elephant: 55) compared to ruminant herbivores
(cattle: 11; sheep: 26), not in carrion (chicken livers: 0); DBRU collec-
tion records on dung of cattle (11), donkey (1).
Temporal activity: Flight activity in darkness, primarily during the late
summer rainy season (Dec. to May, also Oct.).
Ecoregions Botswana: SE Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Kalahari Duneveld (SVkd), W and S Eastern
Kalahari Bushveld (SVk) (Savanna Biome); extreme NE Bushmanland
(NKb) (Nama Karoo Biome).
Assessment rationale: EOO = 211 965 km2; although quantitative sup-
port is required, AOO probably limited by occurrence of deep sand,
2 mm
which nevertheless covers most of the large EOO; range restricted to a
little-transformed region used primarily for conservation and grazing of
farm livestock to which it is attracted despite a measured bias to omni-
vore and indigenous herbivore dung; assessed as Least Concern ((LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations; protected in
Kgalagadi Transfrontier Park (South Africa/Botswana).

ONTHOPHAGINI
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Euonthophagus jeanneli
(d’Orbigny, 1913)

= Onthophagus tibialis Frey, 1963

= Onthophagus tibialis diversiceps Frey, 1975b
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus jeanneli: ‘Afrique orientale anglaise: Tavé-

ta......; Afrique orientale allemande: rivière Himo sur le versant sud-est
du Kilimandjaro’ [Taveta, Kenya; Himo River, Tanzania]; O. tibialis:
‘Makindu, Kongo’ [possibly an error for Makindu, Kenya]; O. tibialis
diverciceps: ‘Krueger Nationalpark: Skukuza, Satara, Pundamilia’ [Kru-
ger National Park, South Africa].
Taxonomy: Accepted species; O. tibialis diversiceps not formally synony-
mised; status and origin of O. tibialis Frey, 1963 needs further examina-
tion; minor sexual dimorphism.
Distribution: Patchy occurrence owing to monogastric dung association
in hot, dry savanna on the E seaboard of Africa: South Africa, Kenya,
Tanzania.
Locality data (mean ± SD, range): Altitude: 371 ± 102, 153–576 m;
annual rainfall: 582 ± 100, 458–788 mm; annual temperature (max. +
min. /2): 22.4 ± 0.9, 20.7–23.5°C (N=19).
Habitat: No quantitative assessment; at South African Wildlife College:
recorded in very open savanna on coarser granite (2.6) and finer gabbro-
derived soils (3.8) within the Kruger National Park; much less abun-
dant in adjoining communal farm rangeland (0.8, 0.3); severely limited
DBRU collection records from sandy loam (3) in shrubland (1), open
woodland (1).
Food types: No quantitative assessment; all DBRU collection records
from coarse-fibred, monogastric herbivore dung; elephant (13), zebra
(1).
Temporal activity: Diel flight periodicity unknown, active during the
summer rainy season (Nov. to Mar.).
Bioregions South Africa: Mopane (SVmp), Lowveld (SVl) (Savanna 2 mm
Biome).
Assessment rationale: EOO = 409 200 km2 (gross estimate, possibly
larger); AOO would be much smaller and dependent on occurrence of
large monogastric herbivores from whose dung all known records have
been made; possibility of soil or vegetation specialisation not tested;
despite small known AOO, nearly all modern records are from two
large national parks; therefore, currently assessed as Least Concern
(LC), although this would change if elephant populations came under
increasing threat.
Conservation measures: Assessment of conservation status would be
much improved by a survey of elephant dung in savanna along the
E seaboard of Africa to determine if the disjunct pattern is real or a
collecting artefact; further quantitative data is required on ecological as-
sociations, and further work on the effects of range contraction of large
monogastric herbivores; protected in Kruger National Park (South
Africa), Tsavo East National Park (Kenya).

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Euonthophagus vicarius
(Péringuey, 1901)

No synonyms (but see Taxonomy)

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: as Onthophagus vicarius: ‘Cape Colony (Beaufort West)’

[Beaufort West, Western Cape, South Africa].
Taxonomy: Requires formal revalidation; currently synonymised in error
with Euonthophagus carbonarius (Klug, 1855) described from ‘Sena’
[central Mozambique].
Distribution: Centred on upland Karoo in the S of the arid, late summer
rainfall region (climates III1, II(III)a, II4a – Walter & Lieth 1964)
and dry parts of spring/autumn, bimodal rainfall region (climate III5):
South Africa.
Locality data (mean ± SD, range): Altitude: 1 236 ± 233, 564–1 598 m;
annual rainfall: 282 ± 51, 197–439 mm; annual temperature (max. +
min. /2): 15.8 ± 1.2, 14.0–18.9°C (N=64).
Habitat: No adequate quantitative assessment; in Northern Cape: record-
ed with high frequency in Upper Karoo (70.6%, 48 out of 68 sam-
ples), frequency much lower in uplands to S of Orange River (19.0%,
23/121), but abundance similar in samples with E. vicarius (3.1 or 3.6/
sample), rare in SW Kalahari, absent from low altitude in Bushman-
land; abundance/sample similar on sand (2.4), sandy loam (3.6), sandy
clay loam (3.4); DBRU collection records on sand (3), sandy loam (5),
sandy clay loam (6), clay (1) in grassland/pasture (2), scrub/shrubland
(10), open woodland (2).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (12), horse (2).
2 mm
Temporal activity: Diel flight periodicity unknown, but presumably
flies in darkness like other Euonthophagus spp.; recorded during spring
(Sept.) and late summer rainy seasons (Dec. to Mar.).
Bioregions South Africa: Upper Karoo (NKu) (Nama Karoo Biome),
Rainshadow Valley Karoo (SKv) (Succulent Karoo Biome), margins of
four adjoining bioregions.
Assessment rationale: EOO = 97 405 km2; widespread across a little-
transformed range used primarily for grazing of farm livestock; record-
ed with high frequency during a survey of the Upper Karoo; assessed as
Least Concern (LC) despite absence of known protection.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; currently
not known from any protected area; a survey of upland areas in Karoo
National Park would be worthwhile.

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Genus Hamonthophagus Roggero,

Dierkens, Barbero & Palestrini, 2016
Type species and designation: Onthophagus bituberculatus Olivier, 1789, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Roggero et al. (2016).

Hamonthophagus Roggero, Dierkens, Barbero & Palestrini, 2016, comprises five species recently excised from Group 32
of Afrotropical Onthophagus Latreille, 1803, as defined by d’Orbigny (1913). These species are distributed throughout
arid to dry savannas from S to E to W Africa. Three species are represented in southern Africa: one in the arid SW; two in
savanna with representation also in the E tropics.

ONTHOPHAGINI
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Hamonthophagus acutus
(d’Orbigny, 1908)

= Onthophagus bituberculatus Boheman, 1860

Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Onthophagus acutus: ‘lac Ngami..... Okahandya....

Salem’ [Botswana: Lake Ngami; Namibia: Okahandja and Salem];
O. bituberculatus [pre-occupied name] ‘in vicinitate fluviorum Svakop
et Kuisip’ [Namibia: vicinity of Swakop and Kuiseb rivers].
Taxonomy: Accepted species recently transferred from Onthophagus
Group 32 of d’Orbigny (1913); some type localities would, presum-
ably, relate to a close relative (H. fallax d’Orbigny, 1913): i.e. ‘région de
Mpala.....Nyassa’ [Mpala region, Tanzania; Lake Malawi region]; this
type material was not examined in the recent review of Roggero et al.
(2016) as it could not be located.
Distribution: Widespread in the S and central arid late summer rainfall
region: central SW South Africa, W Botswana, Namibia; records from
Malawi, Tanzania, Democratic Republic of the Congo (DRC) remain
unvalidated and likely represent confusion with H. fallax.
Locality data (mean ± SD, range): Altitude: 1 005 ± 315, 58–1 840 m;
annual rainfall: 219 ± 109, 14–532 mm; annual temperature (max. +
min. /2): 19.1 ± 1.9, 13.4–22.9°C (N=50).
Habitat: No quantitative assessment; limited DBRU collection records on
sand (6), sandy loam (1), sandy clay loam (1) in scrub/shrubland (6);
also in pasture (1), open woodland (1).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (7), wildebeest (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Sept. to May).
Ecoregions Namibia, Botswana: Succulent Karoo (AT1322), central
Namib Desert (AT1315), S Namibian Savanna Woodlands (AT1316),
2 mm
SW Kalahari Xeric Savanna (AT1309), margins of one other ecoregion.
Bioregions South Africa: Kalahari Duneveld (SVkd), W Eastern Kalahari
Bushveld (SVk) (Savanna Biome); Upper Karoo (NKu) (Nama Karoo
Biome); marginal in two other bioregions.
Assessment rationale: EOO = 390 575 km2; AOO may be biased to arid
regions of sandy soils; however, widespread in little-transformed re-
gions used primarily for grazing of domestic livestock or conservation;
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations and more pre-
cise assessment of EOO and AOO after taxonomic revision; protected
in Kgalagadi Transfrontier Park (South Africa, Botswana), Namib-
Naukluft National Park (Namibia).

ONTHOPHAGINI
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Hamonthophagus depressus
(Harold, 1871c)

= Onthophagus laceratus Gerstaecker sensu Péringuey, 1901

= Onthophagus carteri Blackburn, 1904
= Onthophagus depressus var. marmoreus d’Orbigny, 1904
Global: LC (see IUCN Red List – LC as Onthophagus)

J A S O N D J F M A M J

Type localities: As Onthophagus depressus: ‘Caffraria’ [SE South Africa],

lectotype: ‘SOUTH AFRICA: [Caffraria]’; O. laceratus sensu Pé-
ringuey: ‘Natal (Durban, Malvern, Maritzburg)’ [KwaZulu-Natal,
South Africa]; O. carteri: ‘N. S. Wales; Sydney’ [Australia: accidental
introduction]; var. marmoreus: Not stated.
Taxonomy: Accepted species recently transferred from Onthophagus
Group 32 of d’Orbigny (1913); record for O. laceratus Gerstaeck-
er sensu Péringuey, 1901, from ‘Salem, Kuisip’ [Namibia], would be
O. acutus d’Orbigny, 1908; the record is marked, ‘teste Boheman’ [=
witnessed by Boheman].
Distribution: Mostly known from sandy savanna and E coastal sands,
southern Africa; NW and SE centres possibly separated by Kalahari
deep sands; SW RSA outlier in Orange River Valley; Botswana, Namib-
ia, South Africa, Mozambique, Malawi, Zimbabwe, Zambia, Angola,
Democratic Republic of the Congo (DRC), Burundi, Tanzania; also
recorded as a rarity on Kenyan coastal plain; accidental introduction
into E Australia, SE USA, Madagascar, Mauritius.
Locality data (mean ± SD, range): Altitude: 701 ± 476, 0–1 762 m; an-
nual rainfall: 676 ± 200, 190–1 198 mm; annual temperature (max. +
min. /2): 20.1 ± 1.9, 15.5–24.4°C (N=128).
Habitat: No adequate quantitative data; across a post-mining vegetation
gradient at Richards Bay: primarily in early succession shrubland
(1.90), rare in grassland (0.03) and under shade in woodland (0.03); 2 mm
DBRU collection records from sand (12), sandy loam (5) in open
shrub/woodland (10), pasture (4).
Food types: No adequate quantitative data; at Phalaborwa: only sampled
to pig dung (3) not elephant or cattle dung (0); limited DBRU col-
lection records to dung of cattle (14), horse (1), elephant (1), human/
cattle mix (2).
Temporal activity: Flight activity in darkness; recorded during most
months of the year (Oct. to May; July, Aug.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: N Namib-
ian Savanna Woodlands (AT1316), Kalaharia Xeric Savanna (AT1309),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern Miombo
Woodlands (AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); Indian Ocean Coastal Belt Biome
(CB); marginal in four other bioregions.
Assessment rationale: EOO = 669 355 km2 (E African range underes-
timated); AOO may be restricted by soil and vegetation type special-
isation, but data inconclusive; not recorded in the Kalahari despite
an apparent bias to deep sands; usually recorded in low numbers but
found across a wide area; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by better quantitative data on ecological associations; protected
in Kruger National Park, uMkhuze Game Reserve (South Africa).

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Hamonthophagus fallax
(d’Orbigny, 1913)

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: ‘Afrique orientale allemande: Dar-es-Salam...; sud du

Nyassa: Zomba dans la région du haut Chiré’ [Dar es Salaam, Tanza-
nia; Zomba in the region of the upper Shire River, Malawi], lectotype:
‘MALAWI: [Nyassa Zomba haut Chiré]’.
Taxonomy: Accepted species recently transferred from Onthophagus
Group 32 of d’Orbigny (1913).
Distribution: Dry to moist, hot, lowland savanna from S central to E
Africa: Kenya, Tanzania, Burundi, Democratic Republic of the Congo
(DRC), Malawi, Zambia, N Botswana, N Namibia; also cited from
Angola; record from Eastern Cape, South Africa (Roggero et al. 2016),
requires further validation.
Locality data (mean ± SD, range): Altitude: 884 ± 184, 492–1 075 m;
annual rainfall: 666 ± 242, 443–1 108 mm; annual temperature (max.
+ min. /2): 22.5 ± 0.8, 21.4–23.9°C (N=7).
Habitat: No quantitative assessment; unknown.
Food types: No quantitative assessment; unknown.
Temporal activity: Diel flight periodicity unknown, but probably flies in
darkness like other Group 32 species; active in summer rainy season
(Nov. to Mar.).
Ecoregions Namibia, Botswana: Zambezian Baikiaea Woodlands
(AT0726), N margins Zambezian and Mopane Woodlands (AT0725).
Assessment rationale: EOO = 327 565 km2 (underestimated); range and
ecological associations poorly known; S part of range may be associated
with riverine systems and this needs to be investigated; currently as-
sessed as Data Deficient (DD).
Conservation measures: Further investigation of range and ecological as-
sociations required before an assessment may be made of conservation
status; protected in Liwonde National Park, Malawi.

2 mm

ONTHOPHAGINI
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Genus Haroldius Boucomont, 1914

Type species and designation: Haroldius rugatulus Boucomont, 1914, by subsequent designation (Arrow 1931).
Synonyms: = Cyclotrogus Wasman, 1918: Cyclotrogus heimi Wasman, 1918, by original designa-
tion.
= Ponerotrogus Silvestri, 1924: Ponerotrogus annandalei Silvestri, 1924, by original des-
ignation.
= Afroharoldius Janssens, 1949: Afroharoldius ennearthrus Janssens, 1949, by original
designation.
= Formicdubius Philips & Scholtz, 2000: Type species: Formicdubius convexus Philips
& Scholtz, 2000, by original designation.
Last review: Entire genus and relatives considered by Krikken and Huijbrechts (2006), new Orien-
tal species added subsequently.

The tribal level placement of Haroldius Boucomont, 1914, has been questioned. Although formerly placed in the Can-
thonini (Krikken & Huijbrechts 2006), the genus has most recently been assigned to the Onthophagini (Krell & Philips
2010; Philips 2016). However, the most recent molecular phylogeny (Tarasov & Dimitrov 2017) indicates affinities
to basal Scarabaeinae currently classified with the Canthonini (Deltochilini) although bootstrap support varies from
<50–59% using different methods.

Haroldius currently comprises 37 small-bodied species recorded in the Afrotropical (4) and Oriental regions (33). Various
species have been cited as showing ant or termite associations including all those classified in the synonymous genera (Was-
man 1918; Silvestri 1924; Janssens 1949; Philips & Scholtz 2000). However, as some species lack the trichomes (Krell
& Philips 2010) that are indicators of myrmecophily, validity of the current generic membership of Haroldius requires
further investigation.

Apparently, the four African species all possess trichomes, which are regions of setae associated with release of glandular
secretions used to appease ants (Krell & Philips 2010). As some morphologically similar Oriental species also show ant
associations, suggestions for revalidation of the past generic-level separation from Oriental Haroldius may or may not be
justified.

A single Haroldius species has been recently recorded in association with ants at a single locality in South Africa (Philips &
Scholtz 2000; Krell & Philips 2010). Ant or termite associations probably account for such rarity of collection and further
African species may await discovery.

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Haroldius convexus
(Philips & Scholtz, 2000)

= Formicdubius convexus Philips & Scholtz, 2000

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: H. convexus, nom. nov. for Formicdubius convexus: ‘South

Africa, Pretoria, The Willows, 25°45’S, 28°21’E’.
Taxonomy: Accepted species, but difficult to distinguish from the larger-
bodied H. leleupi Janssens, 1953, described from Katanga [Lualaba
Province], Democratic Republic of the Congo (DRC).
Distribution: Known only from the type locality in the suburbs of Preto-
ria [Tshwane], Gauteng, South Africa, within a region of moist, upland
savanna.
Locality data (average): Altitude: 1 452 m; annual rainfall: 699 mm;
annual temperature (max. + min. /2): 17.2°C (N=1).
Habitat: No quantitative assessment; type series of 16 individuals found
under a stone together with ants (Pheidole megacephala (F.)).
Food types: No quantitative assessment; like all other African Haroldius
species, apparently possesses trichomes (setae arranged singly or in clus-
ters near the articulation between the abdomen and prothoracic disc),
which would be indicative of ant association if involved in secretion of
allomones.
Temporal activity: Diel periodicity unknown; recorded in the mid-
summer rainy season (Jan.).
Bioregions South Africa: S edge of Central Bushveld (SVcb) (Savanna
Biome).
Assessment rationale: EOO not known; poorly represented in museum
collections, probably owing to ecological specialisation and very small
body size; although ecologically poorly known, found in association
with ants like two of the other three known species (one with termites);
0.5 mm possible host, Pheidole megacephala, is thought to be native to south-
ern Africa, but is widespread across various vegetation types in various
African countries; unknown if H. convexus shares this range; currently
assessed as Data Deficient DD).
Conservation measures: It is necessary to resolve relationships with
H. leleupi and mount a quantitative survey to determine EOO, AOO
and ecological associations before an assessment of conservation status
may be made; not currently known from any protected region.

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Genus Heteroclitopus Péringuey, 1901

Type species and designation: Heteroclitopus remipes Péringuey, 1901, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Branco (1991); new species added by Branco and Josso
(2007).

Heteroclitopus Péringuey, 1901, currently comprises 10 poorly known species recorded in savanna and forest areas of the
Afrotropical region. Rarity of collection despite widespread occurrence may indicate specialist habits although these are
mostly unknown. Heteroclitopus are often sampled to light after nightfall. The single species described and recorded from
E savanna of South Africa and Zimbabwe is the southernmost member of the genus. Péringuey (1901) speculates that it
may be associated with termites.

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Heteroclitopus remipes
Péringuey, 1901

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: ‘Natal (Durban), Southern Rhodesia (Upper Hanyani

River)’ [Durban, KwaZulu-Natal, South Africa; inexact, Zimbabwe].
Taxonomy: Accepted species.
Distribution: Scattered records suggest a widespread savanna and SE
coastal distribution, but records are too few and/or inexact to deter-
mine the precise range; South Africa, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 809 ± 597, 52–1 452 m;
annual rainfall: 756 ± 209, 413–966 mm; annual temperature (max. +
min. /2): 19.0 ± 1.4, 17.2–20.5°C (N=6).
Habitat: No quantitative assessment; unknown.
Food types: No quantitative assessment; unknown although Péringuey
(1901) and Branco (1992) speculate on termite associations.
Temporal activity: Flight activity in darkness; seasonal activity unknown.
Bioregions South Africa: East Kalahari Bushveld (SVk), Central Bush-
veld (SVcb), Lowveld (SVl) (Savanna Biome); Indian Ocean Coastal
Belt Biome (CB).
Assessment rationale: EOO = 42 635 km2 (gross underestimate; South
African range only); ecologically poorly known and rarely encountered,
perhaps due to specialised association with termites; exact range uncer-
tain; assessed as Data Deficient (DD).
Conservation measures: To adequately assess conservation status, quan-
titative data on ecological associations is required, particularly data to
determine whether or not it is ant- or termite-associated; protected in
Dronfield Nature Reserve (South Africa).

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 427

Genus Hyalonthophagus Palestrini & Giacone, 1988

Type species and designation: Onthophagus (Proagoderus) viridiceps d’Orbigny, 1910, by original designation.
Synonyms: None.
Last review: Created as a subgenus of Onthophagus Latreille, 1802 (Palestrini & Giacone 1988),
reviewed by Palestrini and Giacone (1989); one new species (Deschodt et al. 2019).

Although recently separated from Proagoderus van Lansberge, 1883, and described as a subgenus of Onthophagus Latreille,
1802, Hyalonthophagus Palestrini & Giacone, 1988 has been subsequently treated at either subgeneric (Palestrini & Gi-
acone 1989) or generic level (Hanski & Cambefort 1991b). Although not formally raised to generic status by 2017, it was
treated as a genus since it differs distinctly from other onthophagines. Subsequent to completion of this book, Hyalontho­
phagus has been formally raised to genus level with description of one new species (Deschodt et al. (2019); no species
account, but see Preamble).

At the end of 2017, Hyalonthophagus comprised ten very closely related Afrotropical species distributed in the S, E and
W within the drier or hotter savannas of the continent. Two described species, then, occurred within the savannas of Na-
mibia, Botswana and South Africa: one in E savannas mainly in South Africa; one on the sands at the N and NE edge of
the Kalahari (but see Preamble). Both are recorded with reasonable frequency and are therefore assessed as Least Concern
(LC).

ONTHOPHAGINI
428 SURICATA 6 (2020)

Hyalonthophagus alcyon
(Klug, 1855)

No synonyms
Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type localities: As Onthophagus alcyon: ‘Tette und Sena’ [central Mozam-

bique: Tete and Sena], lectotype: ‘Tete and Sena’.
Taxonomy: Accepted species within a complex of very closely related taxa
that are difficult to separate; limited sexual dimorphism (head); appar-
ently characterised by metallic blue/black colouration.
Distribution: Patchy records from savanna of southern central Africa: N
♂ and NE edges of S Kalahari and Zambezi River Valley; Mozambique,
N South Africa, W Zimbabwe, N Botswana, S Angola; citations from
Kenya, Tanzania require validation.
Locality data (mean ± SD, range): Altitude: 938 ± 315, 61–1 229 m;
annual rainfall: 635 ± 95, 472–813 mm; annual temperature (max. +
min. /2): 21.9 ± 1.7, 19.6–25.7°C (N=12).
Habitat: No quantitative assessment; suspected to be a deep sand special-
ist, but not well supported by available data; limited DBRU collection
records on sand (2), sandy loam (2), sandy clay loam (1) in grassland
(1), open woodland (3).
Food types: No quantitative assessment; DBRU collection records biased
to dung of monogastric herbivores (elephant, 8) compared to ruminant
herbivores (cattle, 3).
Temporal activity: Diurnal flight activity during the summer rainy season
(Nov. to Feb.).
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Zambezian Baikiaea Woodlands (AT0726), Zambe-
zian and Mopane Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 474 810 km2; EOO a gross estimate based
2 mm on limited available data; poorly known and possibly confused with
other species; AOO and population density may be influenced by soil
(sand) and dung type specialisation (dung of monogastric herbivores
and their occurrence), but quantitative data is required in support;
currently assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation would be improved
by survey and accurate identification to determine full EOO and by
quantitative data on ecological associations to determine AOO; pro-
tected in several national parks: Chobe (Botswana), Hwange (Zimbab­
we), Bicuar (Angola).

ONTHOPHAGINI
SURICATA 6 (2020) 429

Hyalonthophagus alcyonides
(d’Orbigny, 1913)

= Oniticellus splendens Wallengren, 1881

= Onthophagus (Proagoderus) alcyon var. viridiceps d’Orbigny, 1902
= Onthophagus (Proagoderus) alcyon var. virens d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: as Onthophagus alcyonides: ‘Lac Ngami......; Transvaal:

Shilouvane près de Leydsdorp....; Natal: Ladysmith....; Colonie du
Cap: Port-EIizabeth’ [Botswana: Lake Ngami; South Africa: Shilu-
vane, Ladysmith, Port Elizabeth], lectotype: ‘Transvaal: Shilouvane’;
O. splendens: ‘Transvaalia’ [South Africa]; var. viridiceps and var. virens: ♂
‘Mozambique’.
Taxonomy: Accepted species within a complex of very closely related taxa
that are difficult to separate; limited sexual dimorphism (head); me-
tallic colouration varies from cupreous to green; status of Wallengren
synonym requires further study.
Distribution: Mostly moist savanna centred on NE South Africa with
scattered outlier records in Botswana, central and N Mozambique, SE
South Africa.
Locality data (mean ± SD, range): Altitude: 576 ± 473, 0–1 628 m; an-
nual rainfall: 694 ± 177, 414–1 220 mm; annual temperature (max. +
min. /2): 20.9 ± 2.1, 15.1–25.2°C (N=62).
Habitat: In Gauteng bushveld: strong bias to deep sand (134) compared
to sandy clay loam (3) and to open woodland (107) compared to shad-
ed thicket (5) or grassland (25) (wrongly cited as H. alcedo d’Orbigny,
1913); in uMkhuze Game Reserve: no consistent soil bias, sand (5.4),
sand on clay (3.2), sandy clay loam (2.5), clay (4.9); DBRU collection
records suggest a bias to finer-grained soils, sand (6), sandy loam (6),
sandy clay loam (12), clay (1) in grassland/pasture (8), open shrub/
woodland (16), forest/thickets (3).
Food types: In Gauteng bushveld: strong bias to omnivore dung (pig:
75) compared to herbivore dung (horse: 11; cattle: 6) (wrongly cited
as H. alcedo d’Orbigny, 1913); DBRU collection records from dung
of cattle (13), buffalo (1), wildebeest (1), elephant (8), rhinoceros (3),
2 mm
donkey (1), baboon (2), human/cattle mix (1).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.).
Ecoregions Botswana: Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl), moister edges of Mopane (SVmp) (Savanna Biome); also N
margins of two other bioregions with a few scattered records in Sub-
Escarpment Grassland (Gs) (Grassland Biome).
Assessment rationale: EOO = 232 285 km2; widespread in moist woody
savanna of NE South Africa in both farm rangeland and reserves; data
suggest that clearance of woodland would reduce population density;
however, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data to resolve conflicting results on soil as-
sociations; protected in uMkhuze Game Reserve, Kruger National Park
(South Africa).

ONTHOPHAGINI
430 SURICATA 6 (2020)

Genus Kurtops Roggero, Barbero & Palestrini, 2016

Type species and designation: Onthophagus signatus Fahraeus, 1857, by original designation.
Synonyms: None.
Last review: Genus created and reviewed by Roggero et al. (2016).

Kurtops Roggero, Barbero & Palestrini, 2016, was recently created to accommodate the species comprising Group 21
(d’Orbigny 1913) of Onthophagus Latreille, 1802. Strong evidence is presented by Roggero et al. (2016) to support this
decision. Kurtops currently comprises three endemic Afrotropical species with distributions centred in southern Africa.
The two better known species are common on sandy soils in the Kalahari or the Kalahari and E coast. The third is ex-
tremely poorly known.

ONTHOPHAGINI
SURICATA 6 (2020) 431

Kurtops caffrarius
(d’Orbigny, 1902)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Caffrérie’ [inexact, ‘Caffraria’; somewhere in southern Af-

rica].
Taxonomy: Accepted species known by single holotype specimen; former-
ly classified in Onthophagus Group 21 and recently transferred to the
new genus, Kurtops Roggero, Barbero & Palestrini, 2016; one unla-
belled specimen seen in Ditsong Museum.
Distribution: No precise known locality; type locality of ‘Caffraria’ var-
iously used by different collectors and authors to refer to SE Africa or
even the entire subcontinent of southern Africa.
Locality data: None available.
Habitat: Not known.
Food types: Not known.
Temporal activity: Diel flight periodicity and seasonal activity unknown.
Bioregions South Africa: Unknown.
Assessment rationale: EOO unknown; as a very close relative of the
Kalahari-centred K. quadraticeps (Harold, 1867), K. caffrarius might be
expected to show similar specialist habitat associations on deep sand
in open shrub/grassland, but there is no support for this hypothesis;
known by extremely few old museum records; given the absence of
precise locality data and lack of ecological data, assessed as Data Defi-
cient (DD); however, paucity of records presumably indicates specialist
association and/or highly restricted range.
Conservation measures: Impossible to recommend any conservation
measures considering the paucity of distribution and ecological data;
no obvious focal area for a survey to rediscover this species; occurrence
within the shaded area on the map considered unlikely; occurrence in
large area around question mark, possibly, more likely.

2 mm

ONTHOPHAGINI
432 SURICATA 6 (2020)

Kurtops quadraticeps
(Harold, 1867c)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Freistaat Oranje’ [Free State, South Africa].

Taxonomy: Accepted species formerly classified in Onthophagus Group 21
and recently transferred to the new genus, Kurtops Roggero, Barbero &
Palestrini, 2016.
Distribution: Arid to dry deep sands of the Kalahari, E Kalahari outliers
and deep sands in Kaokoveld; South Africa, Botswana, Namibia, Zim-
babwe.
Locality data (mean ± SD, range): Altitude: 1 079 ± 158, 703–1 684 m;
annual rainfall: 321 ± 122, 157–659 mm; annual temperature (max. +
min. /2): 19.8 ± 1.2, 16.3–23.1°C (N=115).
Habitat: No quantitative assessment; on deep sand in Botswana: much
more abundant in the open shrub/grassland of the arid SW (1 242)
than the moister NE (46); DBRU collection records from sand (23),
sandy loam (4), sandy clay loam (1) in open woodland (5), scrub/
shrub­land (14), grassland (5).
Food types: On deep sand in Botswana: strong bias to omnivore dung
(pig: 794), but also attracted to herbivore dung types (elephant: 207,
cattle: 97, sheep: 184), poorly attracted to carrion (chicken livers: 3);
DBRU collection records from the dung of cattle (22), buffalo (1),
waterbuck (1), elephant (1), horse (1), donkey (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: N Namibian Savanna
Woodlands (AT1316), Kalahari Xeric Savanna (AT1309), Kalahari
Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea Wood-
lands (AT0726), NW Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Kalahari Duneveld (SVkd), Eastern Kalahari
2 mm Bushveld (SVk), sand outliers: Central Bushveld (SVcb), Mopane
(SVl), (Savanna Biome); extreme N margins Nama Karoo Biome.
Assessment rationale: EOO = 799 195 km2; observations suggest that
the AOO is restricted by soil specialisation although deep sand covers
> 80% of the widespread EOO; range is little transformed, particularly
in the SW, which is used primarily for the grazing of domestic livestock
to whose dung it is readily attracted despite a bias to omnivore dung;
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and vegetation associations; pro-
tected in Central Kalahari Game Reserve and Kgalagadi Transfrontier
Park (Botswana, South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 433

Kurtops signatus
(Fahraeus, 1857)

= Onthophagus junodi d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: K. signatus: ‘prope fluvium Limpopo’ [close to Limpopo

River, southern Africa]; O. junodi: ‘Mozambique: district de Delagoa...,
Delagoa-Bay’ [Maputo, Mozambique].
Taxonomy: Accepted species formerly classified in Onthophagus Group 21
and recently transferred to the new genus, Kurtops Roggero, Barbero &
Palestrini, 2016; synonymy of O. junodi should be re-examined; elytral
colour change from entirely cream to increasing amounts of melanic
patternation, currently considered to represent intraspecific variation.
Distribution: Widespread on sandy soils in dry inland and moist SE
coastal savanna of southern Africa: Angola, Namibia, Botswana, South
Africa, Zimbabwe, Mozambique.
Locality data (mean ± SD, range): Altitude: 935 ± 347, 0–1 958 m;
annual rainfall: 362 ± 193, 133–971 mm; annual temperature (max. +
min. /2): 19.8 ± 1.8, 14.3–23.7°C (N=293).
Habitat: In uMkhuze Game Reserve: extreme bias to sand (132.0) com-
pared to sand over clay (3.6), sandy clay loam (0.1), clay (0.3); sup-
ported in Gauteng bushveld: sand (74), sandy clay loam (1) with a bias
to open woodland (50) and grassland (25), not in shaded thickets (0);
supported by DBRU collection records on sand (33), sandy loam (13)
in grassland/pasture (10), scrub/shrubland (13), open woodland (22).
Food types: In Botswana: attracted in abundance to various dung types:
pig (3 727), elephant (1 705), cattle (960), sheep (2 221) and rarely
to carrion (46); at Phalaborwa: similar bias to dung of pig (238), but
more to cattle (160) than elephant (57); DBRU collection records on
1 mm
various dung types: cattle (32), wildebeest (1), impala (1), elephant (6),
rhinoceros (1), zebra (1), horse (1), donkey (4), human (2), human/
Melanic elytra
cattle mix (1)
Temporal activity: Diurnal flight activity during most months of the year
(Aug., Oct. to May); but, in Gauteng bushveld: primarily active early
in the rainy season (Oct. to Dec.) tailing off into later summer (Jan.,
Feb.); at Phalaborwa: moderately affected by weather conditions: warm
cloudy day following light rainfall (148), hot dry day (104), warm day
soon after very heavy rainfall (204).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: N Namib-
ian Savanna Woodlands (AT1316), Kalahari Xeric Savanna (AT1309),
Angolan Mopane Woodlands (AT0702), Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726),
Zambezian and Mopane Woodlands (AT0725), Maputaland Coastal
Forest Mosaic (AT0119).
Bioregions South Africa: N Bushmanland (NKb), N Upper Karoo (NKu)
(Nama Karoo Biome); Kalahari Duneveld (SVkd), Eastern Kalahari
Bushveld (SVk), Central Bushveld (SVcb), Mopane (SVmp), Lowveld
(SVl) (Savanna Biome); N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 133 835 km2 (underestimated); AOO
limited by occurrence of deep sands and other particularly sandy soils,
but these are widespread in southern Africa; recorded in abundance
1 mm
over a wide area on a variety of dung types; occurs in grassland, but bias
to tree savanna suggests population density might be adversely affected
Pale elytra

ONTHOPHAGINI
434 SURICATA 6 (2020)

Kurtops signatus (continued)

by clearance of woodland; transformation in NE range in South Africa:

0–58% (SVl), 2–49% (SVcb), 0–20% (SVmp); however, widespread
with SW range in little-transformed farmland used primarily for graz-
ing of domestic livestock; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by more detailed data on effects of shrubland and wood-
land clearance; protected in various national parks, including: Kruger
(South Africa), Hwange (Zimbabwe), Chobe (Botswana).

ONTHOPHAGINI
SURICATA 6 (2020) 435

Genus Milichus Péringuey, 1901

Type species and designation: Onthophagus apicalis Fahraeus, 1857, by monotypy.
Synonyms: None.
Last review: Entire genus reviewed by Cambefort (1996b).

Milichus Péringuey, 1901, currently comprises 15 species recorded from dry to moist savannas and forests in the lowlands
to uplands of the Afrotropical region. Two species have been recorded in the region encompassing South Africa, Botswana
and Namibia. One, recorded primarily from monogastric herbivore dung, occurs in dry to moist savanna throughout
Africa. The other from the E coast of South Africa remains unnamed so no species account is presented.

ONTHOPHAGINI
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Milichus apicalis
(Fahraeus, 1857)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Onthophagus apicalis: ‘in terra Natalensi’ [KwaZulu-

Natal, South Africa].
Taxonomy: Currently accepted as a single extremely widely distributed
species; minor sexual dimorphism (prothoracic disc).
Distribution: Widespread in dry to moist African savanna, but, now,
mostly localised in game reserves owing to dung specialisation: South
♂ Africa, Zimbabwe, Botswana, Namibia, Kenya, Côte d’Ivoire; also
cited from Angola, Mozambique, Somalia, Ethiopia, Uganda, Rwanda,
Burundi, Democratic Republic of the Congo (DRC), Gabon, Chad,
Nigeria, Guinea (Conakry).
Locality data (mean ± SD, range): Altitude: 509 ± 316, 55–1 227 m;
annual rainfall: 657 ± 175, 171–1 121 mm; annual temperature (max.
+ min. /2): 21.6 ± 1.9, 16.8–24.4°C (N=32).
Habitat: In uMkhuze Game Reserve: strong bias to finer-grained soils:
sand (1.6), sand on clay (10.0), sandy clay loam (29.4), clay (2.8) with
a 63% bias to woodland (N=407); at South African Wildlife College
(Kruger National Park) and adjacent communal rangeland: bias to
finer-grained gabbro-derived (53.0, 51.9) rather than coarser-grained
granite-derived soils (30.7, 29.8); DBRU collection records on sand
(1), sandy loam (5), clay (1) in shrubland (2), open woodland (5),
shaded thicket (1).
Food types: On deep sand in NE Botswana: sampled only to elephant
dung (18), zero (0) on dung of omnivores (pig), ruminant herbivores
(cattle, sheep) and carrion (chicken livers); at Phalaborwa: extreme bias
to elephant dung (40) compared to pig (2) and cattle dung (2); DBRU
collection records primarily from dung of monogastric herbivores: ele-
phant (8), rhinoceros (2), but also cattle dung (2).
Temporal activity: Flight activity in darkness during the summer rainy
2 mm
season (Oct. to Mar.); at Phalaborwa: primarily active in warm weather
after heavy rainfall (31) rather than in hot, dry weather (5) or warm
weather after light rainfall (8).
Ecoregions Namibia, Botswana, Zimbabwe: N Namibian Savanna
Woodlands (AT1316), Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Lowveld (SVl), Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 7 382 875 km2 (gross estimate); AOO
much smaller due to association with the dung of indigenous mono-
gastric herbivores whose ranges have now mostly contracted into game
reserves, hence the fragmented pattern on the map of southern Africa;
AOO and population density probably also influenced by soil and
vegetation bias; however, owing to extremely wide range and ample
protection in reserves, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on vegetation associations and possible
effects of woodland clearance; conservation status clearly linked to
continuing protection of large monogastric herbivores, particularly,
elephant and rhinoceros; protected in various national parks including
Kruger (South Africa), Chobe (Botswana), Tsavo East (Kenya).

ONTHOPHAGINI
SURICATA 6 (2020) 437

Genus Mimonthophagus Balthasar, 1963b

Type species and designation: Onthophagus apicehirtus d’Orbigny, 1915, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Moretto (2009).

Mimonthophagus Balthasar, 1963b, currently comprises eight species with Afrotropical (7) and Madagascar (1) distribu-
tions. The three species of S and S to E Africa are centred in savannas and this may also be true of four species centred in
W central Africa although available locality data is inexact. Of the three species found in Botswana and/or South Africa,
one appears to be a specialist on E coastal sands with a range extending into the E tropics. The other two may be associated
primarily with elephant dung on sandy soils although current evidence is weak. One is found only at the NE edge of the
Kalahari whereas the other has a range extending into the E tropics.

ONTHOPHAGINI
438 SURICATA 6 (2020)

Mimonthophagus ambiguus
(Péringuey, 1901)

= Onthophagus hinnulus Klug, 1832, sensu Gerstaecker, 1873

= Caccobius bicornutus Frey, 1955
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus ambiguus: ‘Natal (Isipongo)’ [Isipingo,

KwaZulu-Natal, South Africa]; O. hinnulus sensu Gerstaecker: ‘Insel
Sansibar’ [Zanzibar, Tanzania]; C. bicornutus: ‘Sansibar.......Tanganyika
Mhonda’ [NE Tanzania].
Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc).
♂ Distribution: E coastal sands from The Haven, South Africa, to at least,
Malindi, Kenya; also up Zambezi River Valley system to Rekommetjie/
Lake Malawi shores: South Africa, Mozambique, Zimbabwe, Tanzania,
Kenya; also cited from Malawi.
Locality data (mean ± SD, range): Altitude: 34 ± 100, 0–541 m; annual
rainfall: 857 ± 134, 681–1 275 mm; annual temperature (max. + min.
/2): 22.1 ± 1.3, 18.9–24.6°C (N=31).
Habitat: No quantitative data for soil association; on deep sand in Ma-
puto Special Reserve: strong bias to grassland on fossil lagoons (96.0)
compared to sand forest patches (large, 4.7; medium, 23.0; small, 7.9)
and cooler coastal dune forest (1.8); severely limited DBRU collection
records on sand (3) in pasture (1), open woodland (1).
Food types: No quantitative data: severely limited DBRU collection re-
cords on dung of cattle (3); sampled to pig dung.
Temporal activity: Flight activity in darkness (18:00–20:00) during the
summer rainy season (Nov. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128).
Bioregions South Africa: Indian Ocean Coastal Belt Biome (CB).
2 mm
Assessment rationale: EOO = 298 230 km2; range suggests a sand special-
ist although quantitative support is currently lacking; occurs in warm
forested areas although a strong bias to bias to open situations suggests
forest clearance would not be detrimental to population density; wide-
spread along the E coast and found in abundance in coastal reserves;
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and food associations; protected
in iSimangaliso Wetland Park (World Heritage Site) (South Africa),
Maputo Special Reserve (Mozambique).

2 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 439

Mimonthophagus anomalus
(Klug, 1855)

= Onthophagus moestus Fahraeus, 1857

= Onthophagus calvus d’Orbigny, 1904
= Onthophagus julianae Paulian, 1937
= Onthophagus muripedis Frey, 1975b
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus anomalus: ‘Sena’ [central Mozambique];

O. moestus: ‘in terra Natalensi’ [KwaZulu-Natal, South Africa]; O. cal­
vus: ‘Boran Galla: Haouacio’ [Ethiopia]; O. julianae: ‘Afrique: Uganda’;
O. muripedis: ‘Rhodesien: Wankie, Kariba, Dett, Kenmaur; Transvaal, ♂
Krueger Nationalpark: Pundamilia; Botswanaland: Chobe; Moçam-
bique: Gorongosa’ [Zimbabwe, South Africa, Botswana, Mozambique].
Taxonomy: Accepted species; minor sexual dimorphism (fore tibiae).
Distribution: Widespread across savanna from S to NE Africa, but shows
patchy occurrence centred mainly on game reserves: NE South Africa,
Mozambique, Zimbabwe, Botswana, N Namibia, Angola, Tanzania,
Uganda, Ethiopia; also cited from Kenya.
Locality data (mean ± SD, range): Altitude: 730 ± 350, 59–1 247 m;
annual rainfall: 647 ± 145, 466–1 136 mm; annual temperature (max.
+ min. /2): 22.7 ± 1.5, 19.3–25.7°C (N=22).
Habitat: No quantitative data; DBRU collection records primarily from
game reserves or protected areas on sand (1), sandy loam (3), sandy clay
loam (1), clay (1) in grassland (2), open woodland (4).
Food types: In Botswana: strong bias to omnivore dung (pig: 1 061) com-
pared to other dung types (elephant: 115; cattle: 19; sheep: 245) and
carrion (chicken livers: 7); however, DBRU collection records primarily
from dung of elephant (10), also zebra (1), cattle (2).
Temporal activity: Flight activity in darkness during the summer rainy 1 mm

season (Nov. to Apr.); attracted to light.

Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 2 658 380 km2 (gross estimate); despite a
measured bias to omnivore dung, recently found primarily on elephant
dung in game reserves (88.2% of available S records, N=15 out of 17);
despite this apparent range contraction to a much smaller AOO, it is
found in various reserves across a very large EOO; therefore, currently,
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations; protected in various
national parks including: Kruger (South Africa), Hwange (Zimbabwe),
Chobe (Botswana), Bicuar (Angola), Manyara (Tanzania).

1 mm

ONTHOPHAGINI
440 SURICATA 6 (2020)

Mimonthophagus limbibasis
(d’Orbigny, 1905)

No synonyms
Global: DD (see IUCN Red List – NT)

J A S O N D J F M A M J

Type locality: As Onthophagus limbibasis: ‘Lac Ngami’ [Lake Ngami, N

Botswana].
Taxonomy: Accepted species.
Distribution: Only known from savanna woodland on the sand/sandy
loam soil mosaic at the northeast edge of the S Kalahari: Botswana,
Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 044 ± 104, 924–1 106 m;
annual rainfall: 540 ± 109, 414–608 mm; annual temperature (max. +
min. /2): 21.6 ± 0.7, 21.1–22.4°C (N=3).
Habitat: No quantitative assessment; limited DBRU collection records
from sandy loam (2) in open woodland (2).
Food types: No quantitative assessment; limited DBRU collection records
from the dung of monogastric herbivores (elephant, 3).
Temporal activity: Diel flight periodicity unknown, but possibly flies in
darkness like other Mimonthophagus species; recorded during the sum-
mer rainy season (Nov., Feb.).
Ecoregions Botswana, Zimbabwe: Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian Baikiaea Woodlands (AT0726).
Assessment rationale: EOO = 2 790 km2 (possibly underestimated);
poorly known species represented by few individuals in museum col-
lections; on the basis of apparent rarity and possible ecological speciali-
sation, particularly to the dung of range-restricted elephant, this species
has been assessed as Near Threatened (NT) on the IUCN Red List;
however, assessed here as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative sur-
vey of N Botswana, W Zimbabwe and surrounding regions is required
to determine the EOO, AOO and ecological associations; in particu-
lar, the possibility of finer-grained soil, woodland and elephant dung
specialisation needs to be investigated; protected in Hwange National
Park, Zimbabwe.

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 441

Genus Onthophagus Latreille, 1802

Type species and designation: Scarabaeus taurus Schreber, 1759, by monotypy.
Synonyms: = Psilax Erichson, 1848: Type species: Onthophagus pronus Erichson, 1848, by monotypy.
= Monapus Erichson, 1848: Type species: none verified.
Last review: Afrotropical members of genus reviewed by d’Orbigny (1913); subsequent additions
of 239 new Afrotropical species currently classified in Onthophagus; some subsequent
synonymies and various changes in status as regards raising of species groups to subge-
nera within Onthophagus or separation from Onthophagus at generic level (see below).

The long-established genus, Onthophagus Latreille, 1802, is represented globally within tropical, warm temperate and
winter rainfall regions with limited penetration into temperate climates. It is currently represented by approximately
2 145 species, although numbers frequently change because of the addition of new species, subtraction of synonyms or
subtraction of species due to raising of species groups or subgenera to generic level. Numbers of regional species cited here
are essentially estimates owing to the frequency of additions and subtractions (Neotropical: 92; Nearctic: 76; Palaearctic:
192; Oriental: 666 (reduced by ~90 species after Parascatonomus Paulian, 1932, was raised from subgeneric to generic level
(Ochi et al. 2012)); Afrotropical: 796; Madagascar: 4; Australia: 201; New Guinea: 118).

Of the synonyms recently cited for the genus (Smith 2009), only two are cited above. Other names have been revalidated at sub-
generic level by Moretto (2009) (Gonocyphus van Lansberge, 1885b); have become a synonym of Proagoderus van Lansberge,
1883 (Tauronthophagus Shipp, 1895c); or, a possible synonym of Parascatonomus (Chalcoderus Erichson, 1848). If Chalcoderus
was awarded the status of nomen oblitum (=forgotten name) (Palestrini 1982), it would become a synonym of Paracatonomus
due to affiliations shown by the Chalcoderus type species, Onthophagus maculatus (Fabricius, 1801) (see Group 15 notes).

The review of d’Orbigny (1913) examined the entire membership of Onthophagus in the Afrotropical region. Four sub-
genera were recognised. All have subsequently been raised to generic level (Proagoderus; Phalops Erichson, 1847b; Diastel­
lopalpus van Lansberge, 1886; Onthophagus s. str.). A total of 32 species groups were recognised within Onthophagus s. str.
For those Afrotropical groups that remain classified within Onthophagus, many subsequent papers by 20 different authors
have added 239 new species to those described during or before 1913. The process of revision continues and it is likely
that all of d’Orbigny’s 32 core groups will ultimately be raised to subgeneric or generic level. Some have already received
formal subgeneric names or have been extracted to other genera.

The alpha taxonomy of southern African Onthophagus remains incomplete and requires much revision.
(1) No accounts were prepared for a total of 41 currently valid species described from South Africa, Botswana or Na-
mibia as the names could not be reliably matched to reference material.
(2) No accounts were prepared for another eight species described from elsewhere since claimed records from southern
Africa could not be verified.
(3) Many species from South Africa, Botswana and Namibia could not be included as names were unavailable or could not
be validated even, in some cases, for species that were well represented in collections with supporting ecological data.
(4) Undoubtedly, a number of species remain undescribed; some current valid species names are synonyms; some
synonyms deserve revalidation; whereas some have been synonymised with the wrong species.
(5) The 78 species accounts presented, here, may represent only 50–60% of the true diversity of Onthophagus in South
Africa, Botswana and Namibia. However, even some of the species names included in the accounts require valida-
tion or revision.

The group structure proposed by d’Orbigny (1913) is used as a guide to representation by Onthophagus and former
Onthophagus in South Africa, Botswana and Namibia on the understanding that it represents a stage in the revision of the
species complexes.
Group 1: See four probable ant or termite-associated genera (Pseudosaproecius Balthasar, 1941b; Eusaproecius Branco,
1989; Cambefortius Branco, 1989) including one represented in southern Africa: Stiptopodius Harold, 1871b
(but not validated).

ONTHOPHAGINI
442 SURICATA 6 (2020)

Group 2: Four validated widespread species; ranges into E tropics.

Group 3: Represented in southern Africa, but no validated species.
Group 4: One validated highland, endemic; South African species.
Group 5: Subgenera, Gonocyphus van Lansberge, 1885b, and Afrostrandius Moretto, 2009; in the S: one Afrostrandius
on monogastric herbivore dung in savanna; ranging into E tropics.
Group 6: See genus, Mimonthophagus Balthasar, 1963a.
Group 7: Subgenera, Eremonthophagus Zunino, 1979, and Bicornonthophagus Tagliaferri & Moretto, 2012: accounts
for seven widespread (1), southern African (1) or SE African (5) taxa.
Group 8: See genus, Euonthophagus Balthasar, 1959.
Group 9: Four validated species endemic to South Africa.
Group 10: Two southern African (1) or widespread (1) taxa.
Group 11: Kleptocoprid species; accounts for seven widespread (1), southern African (4), or South African endemic (2) taxa.
Group 12: Subgenus, Trichonthophagus Zunino, 1979: possibly kleptocoprid on Heliocopris species; accounts for seven
widespread (1) or southern African (6) taxa.
Group 13: Represented in southern Africa but no validated species; some species transferred to genera, Digitonthophagus
Balthasar, 1959, and Phalops Erichson, 1847b.
Group 14: Represented in South Africa and Namibia by two species that have been validated and transferred to the new genus,
Jossonthophagus Dierkens, Moretto & Génier, 2017, only after submission of this book, so no species accounts.
Group 15: One Group 15 member (Onthophagus maculatus) now in genus, Parascatonomus Paulian 1932; one south-
ern African group member (Onthophagus graphicus Wallengren, 1881) remains classified in Onthophagus;
O. graphicus is a carrion species in southern African savannas, ranging into the E tropics.
Group 16: Accounts for seven species endemic to South Africa (3), southern Africa (3), or widespread to E tropics (1).
Group 17: Accounts for four species endemic to South Africa (3) or southern Africa (1).
Group 18: Carrion-associated; accounts for five species endemic to South Africa (3), Namibia (1), or widespread to E
tropics (1).
Group 19: Carrion-associated; accounts for five taxa widespread to E tropics (2) or endemic to southern (1) or South
Africa (2).
Group 20: Accounts for two SW Arid (1) or Kalahari (1) species.
Group 21: See genus, Kurtops Roggero, Barbero & Palestrini, 2016.
Group 22: Account for one Kalahari-centred species.
Group 23: Accounts for six species widespread in the tropics (2) or endemic to South Africa (4).
Group 24: Accounts for five species widespread to E tropics (1) or centred in southern Africa (4) (one now transferred
to the new genus, Tiaronthophagus Roggero, Moretto, Palestrini & Barbero, 2019).
Group 25: Palaearctic species group.
Group 26: E African montane species group.
Group 27: Removed from Afrotropical fauna (Moretto 2013a).
Group 28: West African species group.
Group 29: Sahel species group.
Group 30: Accounts for two SW African centred species.
Group 31: Subgenus, Furconthophagus Zunino, 1979; accounts for seven species endemic to South Africa (4) or more
widely distributed (3); many others without validated names.
Group 32: See genera, Hamonthophagus Roggero, Dierkens, Barbero & Palestrini, 2016, or, Morettius Roggero, Dier-
kens, Barbero & Palestrini, 2016 (which is not represented in southern Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 443

Onthophagus absyrtus
Balthasar, 1946

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Africa mér., Johannesburg’ [South Africa: Johannesburg].

Taxonomy: Accepted species; sexually dimorphic (head, prothoracic disc),
characters vary in prominence with body size.
Distribution: Moist highland and montane grassland in the N Highveld,
South Africa.
Locality data (mean ± SD, range): Altitude: 1 556 ± 248, 1 135–1 836 m; ♂
annual rainfall: 759 ± 139, 638–1 157 mm; annual temperature (max.
+ min. /2): 15.5 ± 1.8, 13.1–18.9°C (N=11).
Habitat: No quantitative assessment; limited DBRU collection records all
from sandy clay loam (3) in pasture (2), fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (3), human/cattle mix (1).
Temporal activity: Diel flight periodicity unknown; recorded primarily in
the summer rainy season (Sept. to Jan.).
Bioregions South Africa: Primarily around the margins of Mesic High-
veld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 65 350 km2; AOO may be much smaller
due to specialisation to very fine-grained soils; restricted AOO in moist
highland grassland may also account for rarity in collections; however,
quantitative data is required to support severely limited observations;
currently assessed as Data Deficient (DD).
Conservation measures: Before an assessment of conservation status may
be made, it is necessary to make a more precise measurement of EOO/
AOO and to obtain quantitative data on ecological associations plus
2 mm
possible influence of habitat transformation; protected in Suikerbos-
rand and Telperion nature reserves, Gauteng.

♀ 2 mm

ONTHOPHAGINI
444 SURICATA 6 (2020)

Onthophagus aequepubens
d’Orbigny, 1905

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Nyassa’ [Lake Malawi region].

Taxonomy: Accepted Group 2 species; sexually dimorphic (head), charac-
ters vary in prominence with body size.
Distribution: Apparent disjunct distribution pattern within southern
African lowland savanna; possibly a collection artefact or possibly re-
lated to soil and vegetation type specialisation: South Africa, Namibia,
♂ Zimbabwe; type locality possibly Malawi.
Locality data (mean ± SD, range): Altitude: 780 ± 445, 55–1 730 m;
annual rainfall: 570 ± 192, 125–988 mm; annual temperature (max. +
min. /2): 20.5 ± 2.3, 17.2–24.5°C (N=25).
Habitat: In Gauteng bushveld: extreme specialisation to deep sand (142)
compared to sandy clay loam (0), extreme specialisation to shaded
thicket (141) compared to open woodland (1) and grassland (0); soil
association not supported in uMkhuze Game Reserve: sand (0.5), sand
over clay (0.3), sandy clay loam (0.5), clay (1.3) but 100% in shade
(12); severely limited DBRU collection records on sand (1), clay (1) in
woodland (1) and forest (2).
Food types: No quantitative assessment; DBRU collection records on
dung of baboon (1), human/cattle mix (3), horse (2), cattle (2).
Temporal activity: Diurnal flight activity; in Gauteng: peak activity early
in the summer rainy season (Oct. to Jan.), rare thereafter (Feb., March);
also recorded during Aug. in warmer coastal region (iMfolozi Game
Reserve).
Ecoregions Namibia, Zimbabwe: Southern Miombo Woodlands
(AT0719), riverine vegetation of Zambezian and Mopane Woodlands
(AT0725), Namibian Savanna Woodlands (AT1316).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
2 mm
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 245 690 km2 (underestimated); some sup-
port for soil specialisation; clearance of dense woody vegetation would
be detrimental owing to shade specialisation; transformation in South
Africa amounts to 0–58% (SVl), 2–49% (SVcb), 0–20% (SVmp);
however, owing to widespread range and occurrence in both farmland
and game reserves, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data to resolve conflict in soil association
data; quantitative data on food associations would also be useful; pro-
tected in Kruger National Park (South Africa), Lake Mutirikwe Game
Reserve (Zimbabwe).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 445

Onthophagus aeruginosus
Roth, 1851

= Onthophagus tenuicornis Klug, 1855

= Onthophagus chalcostomus Fahraeus, 1857
= Onthophagus aeruginosus var. janthinus d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: ‘Tigré in N. Abyssinien’ [Tigray, N Ethiopia]; O. tenuicor­

nis: ‘Sena’ [central Mozambique]; O. chalcostomus: ‘prope fluvium
Limpopo’ [close to Limpopo River; southern Africa]; var. janthinus:
‘Ouarsangueli’ [N Somalia].
♂
Taxonomy: Accepted Group 2 species; colour variation: usually iridescent
green, occasionally cupreous or blueish, but melanic on South African
Highveld; sexually dimorphic (head, prothoracic disc), characters vary
in prominence with body size.
Distribution: Mainly dry S to NE African savanna or grassland: South Af-
rica, Namibia, Botswana, Angola, Zimbabwe, Malawi, Mozambique,
Tanzania, Kenya, Ethiopia, Somalia; also reported from Rwanda,
Burundi, Democratic Republic of the Congo (DRC), Eritrea; report
from Guinea (Bissau) presumably an error; melanic var. (black squares),
green var. (red squares) with central South African outlier in Orange
River Valley.
Locality data (mean ± SD, range): Green var.: Altitude: 905 ± 530,
0–2 274 m; annual rainfall: 670 ± 175, 34–1 415 mm; annual tem-
perature (max. + min. /2): 19.6 ± 2.4, 10.1–25.2°C (N=208). Melanic
var.: Altitude: 1 406 ± 119, 1 240–1 721 m; annual rainfall: 569 ±
70, 452–703 mm; annual temperature (max. + min. /2): 16.6 ± 1.0,
14.4–18.3°C (N=14).
Habitat: Relative soil generalist with bias to woody vegetation; in Gauteng
bushveld; sand (1 467), sandy clay loam (1 065), grassland (371), open
2 mm
woodland (1 209), shaded thickets (952); in uMkhuze Game Reserve;
sand (14.8), duplex soil (sand over clay) (30.3), sandy clay loam (13.9),
clay (17.3); supported by DBRU collection records: sand (20), sandy
loam (18), sandy clay loam (16), clay (4) in grassland/pasture (19 –
including 10 for melanic var.), shrubland (9), open woodland (14),
forest/thickets (13).
Food types: At Phalaborwa: bias to omnivore dung (pig: 185) over her-
bivore dung (elephant: 84, cattle: 72); DBRU collection records from
various dung types: cattle (45), wildebeest (1), waterbuck (1), impala
(1), elephant (3), rhinoceros (3), zebra (1), horse (1), warthog (2), ba-
boon (1), human (1), human/cattle mixture (18).
Temporal activity: Diurnal flight activity throughout the year; in Gauteng
bushveld: peak flight activity late morning (10:00–12:00) with peak
seasonal activity early in the summer rainy season (Oct. to Jan.) de-
clining a little (Feb. to April) to low (May, June) and very low levels
in the cool dry season (July to Sept.); at Phalaborwa: equally active in
warm sunny conditions soon after heavy rainfall (178) or warm cloudy
conditions soon after light rainfall (171), less active under hot dry con-
ditions (92).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Namibian
Savanna Woodlands (AT1316), NW Kalahari Xeric Savanna (AT1309),
Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian and Mo-
pane Woodlands (AT0725), Southern African Bushveld (AT0717), ♀
2 mm

ONTHOPHAGINI
446 SURICATA 6 (2020)

Onthophagus aeruginosus (continued)

Southern Miombo Woodlands (AT0719), Maputaland Coastal Forest

Mosaic (AT0119), margins of one other ecoregion.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome); N Dry Highveld Grassland (Gh), N
Mesic Highveld Grassland (Gm), N Sub-Escarpment Grassland (Gs)
(Grassland Biome); N Indian Ocean Coastal Belt Biome (CB); mar-
ginal in two other bioregions; melanic var. in Dry Highveld Grassland
(Gh).
Assessment rationale: EOO = 4 968 210 km2 (underestimated gross es-
timate); widespread, soil and vegetation generalist found abundantly
on various dung types in both farmland and reserves; assessed as Least
Concern ( LC).
Conservation measures: None required; protected in many reserves in-
cluding Kruger National Park (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 447

Onthophagus albipennis
Péringuey, 1908

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Cape Colony (Uitenhage)’ [Eastern Cape, South Africa].

Taxonomy: Accepted Group 11 species.
Distribution: Arid to dry areas in S late summer rainfall region and drier
parts of bimodal rainfall region: Northern and Eastern Cape, South
Africa; absence of records from arid terrain between coastal hinterland
and Northern Cape uplands may or may not be a collecting artefact
(see habitat notes below).
Locality data (mean ± SD, range): Altitude: 1 158 ± 209, 121–1 598 m;
annual rainfall: 245 ± 53, 133–427 mm; annual temperature (max. +
min. /2): 17.0 ± 1.7, 14.0–19.4°C (N=113).
Habitat: No adequate quantitative assessment; in Northern Cape: primar-
ily in uplands to S of Orange River (76.7% = 66 out of 86 traps, 4.3/
trap) and in Upper Karoo (92.1% = 115 out of 126 traps, 18.0/trap),
few in arid lowlands of Bushmanland (4.5% = 3 out of 67 traps, 0.01/
trap); also in Northern Cape: recorded on sand (15.8/trap, N=72),
sandy loam (15.1, N=66), sandy clay loam (17.4, N=19); DBRU col-
lecting records from sand (4), sandy loam (3), sandy clay loam (1) in
grassland/pasture (2), scrub (4), open woodland (2).
Food types: No quantitative assessment; sampled to composite cattle/
sheep baits; DBRU collection records on cattle dung (5).
Temporal activity: Diurnal flight activity during the late summer rainy
season (Dec. to May).
Bioregions South Africa: Bushmanland (NKb), Upper Karoo (NKu)
(Nama Karoo Biome), also margins of both Rainshadow Valley Karoo
(SKv) (Succulent Karoo Biome) and Albany Thicket Biome (AT).
Assessment rationale: EOO = 126 235 km2; widespread soil generalist
with high frequency and high abundance in upland Karoo; not record-
2 mm
ed in more arid lower areas; N part of range little transformed and used
primarily for grazing domestic livestock that drop dung to which it is
readily attracted; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by a survey to determine the limits of the EOO, particularly if
the species is also characteristic of the arid Lower Karoo; formal assess-
ment of ecological associations would also assist; not currently recorded
from any reserve.

ONTHOPHAGINI
448 SURICATA 6 (2020)

Onthophagus albipodex
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Lake Ngami’ [Botswana].

Taxonomy: Accepted Group 24 species, but research required on relation-
ships to species of similar appearance described from further N and
further SW; sexually dimorphic (head, prothoracic disc), characters
vary in prominence with body size.
Distribution: Moist savanna regions of SE Africa; apparent disjunction
♂ across the hot, dry Limpopo Valley may be real or a collection artefact;
type locality is an outlier at W edge of range: South Africa, Zimbabwe,
Mozambique, Botswana; also cited from Malawi; citation from Angola
requires validation.
Locality data (mean ± SD, range): Altitude: 988 ± 379, 11–1 510 m;
annual rainfall: 686 ± 123, 414–1 100 mm; annual temperature (max.
+ min. /2): 19.6 ± 2.0, 16.7–24.4°C (N=37).
Habitat: In Gauteng bushveld: bias to deep sand (15) compared to sandy
clay loam (2), bias to grassland (14) compared to open woodland (3) or
shaded thicket (0); only partly supported by limited DBRU collection
records: sandy loam (3), sandy clay loam (4) in grassland/pasture (6),
shrubland (2).
Food types: No quantitative assessment; DBRU collection records: dung
of cattle (9), human (1), human/cattle mix (1).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Feb.); in Gauteng bushveld: only active in early rainy season
(Oct. to Dec.).
Ecoregions Botswana, Zimbabwe: Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian Baikiaea Woodlands (AT0726), Southern
Miombo Woodlands (AT0719), margins of one other ecoregion.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome); N margins Dry Highveld Grassland
2 mm
(Gh) (Grassland Biome).
Assessment rationale: EOO = 324 950 km2 (probably underestimated);
widespread, but recorded in relatively low population density through-
out its range; conflicting soil association data negate any conclusions on
specialisation or generalisation; may not be unduly influenced by clear-
ance of woodland owing to apparent association with open vegetation;
not common, but also not rare; currently assessed as Least Concern
(LC).
Conservation measures: Further quantitative data on ecological associ-
ations is required to improve assessment of conservation status, par-
ticularly to determine if AOO is influenced by soil type; protected in
Kruger National Park and Tswaing Nature Reserve (South Africa).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 449

Onthophagus apiciosus
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Afrique orientale allemande’ [Tanzania].

Taxonomy: Accepted Group 19 species, but distribution requires valida-
tion as often confused with other close relatives; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Moist savanna from southern to East Africa: South Africa,
Angola, Botswana, Zimbabwe, Mozambique, Malawi, Zambia, Tan-
zania; also reported from Democratic Republic of the Congo (DRC), ♂
Kenya.
Locality data (mean ± SD, range): Altitude: 1 038 ± 360, 266–1 698 m;
annual rainfall: 709 ± 151, 416–1 130 mm; annual temperature (max.
+ min. /2): 19.2 ± 1.9, 16.7–23.7°C (N=33).
Habitat: No quantitative assessment; limited DBRU collection records
from sand (1), sandy loam (1), clay (1) in pasture (1), open woodland
(1), forest (1), even an orchard (1).
Food types: No quantitative assessment; limited DBRU collection records
from human/cattle dung mixture (2); but see Assessment rationale.
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); attracted to light.
Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome); N Sub-Escarpment Grassland (Gs)
(Grassland Biome).
Assessment rationale: EOO = 2 057 540 km2; AOO unclear; ecologi-
cally poorly known owing to lack of quantitative data, but Group 19
is thought to be characterised primarily by carrion associations; easily
confused with putative close relatives, but probably widespread pos-
sibly with generalist habitat associations; therefore, assessed as Least 2 mm
Concern (LC).
Conservation measures: Assessment of conservation status needs to be
improved by quantitative ecological data to confirm generalist habitat
associations with omnivore dung and/or carrion specialisation; protect-
ed in Kruger National Park (South Africa).

♀ 2 mm

ONTHOPHAGINI
450 SURICATA 6 (2020)

Onthophagus asperulus
d’Orbigny, 1905

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Transvaal: Boksburg.....Johannesburg....Natal: Malvern’

[South Africa: Gauteng, KwaZulu-Natal].
Taxonomy: Accepted Group 23 species, but status needs to be validat-
ed or revised after comparison with O. impictus Fahraeus, 1857, and
O. scabrosus Péringuey, 1901 (both from type localities in KwaZulu-
♂ Natal) that are currently synonymised with O. hyaena (Fabricius, 1801:
S Eastern Cape and Free State range), but probably in error; sexually
dimorphic (head, prothoracic disc), characters vary in prominence with
body size.
Distribution: Coast and uplands along SE seaboard (E Eastern Cape,
KwaZulu-Natal), N Highveld (Gauteng, Mpumalanga, Limpopo);
South Africa.
Locality data (mean ± SD, range): Altitude: 1 241 ± 533, 46–2 392 m;
annual rainfall: 779 ± 117, 517–1 136 mm; annual temperature (max.
+ min. /2): 16.0 ± 1.9, 9.9–21.4°C (N=139).
Habitat: No adequate quantitative assessment; in Suikerbosrand Nature
Reserve: montane grassland 1 888–1 930 m (3.9 per sample), Highveld
Grassland 1 631–1 685 m (8.7), open woodland 1 637–1 689 m (1.4);
at 1 496–1 541 m in Abe Bailey Nature Reserve: grassland (16.3), open
woodland (2.9); at Carolina in Jan.: natural Themeda grassland (28.5),
rehabilitating crop field (25.2), improved Kikuyu pasture (109.0);
DBRU collection records: sand (2), sandy loam (31), sandy clay loam
(44), clay (5) in grassland/pasture (75), shrubland (1), open woodland
(2), fallow crop field (3).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (84), horse (1), human/cattle mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
2 mm rainy season (Aug. to May).
Bioregions South Africa: N Mesic Highveld Grassland (Gm), Sub-
Escarpment Grassland (Gs) (Grassland Biome); Indian Ocean Coastal
Belt Biome (CB); E Albany Thicket Biome (AT).
Assessment rationale: EOO = 170 285 km2; widespread in farmland
from the warmer, moister N and E Highveld to the SE coast where it
is readily attracted to the dung of farm livestock and is not negatively
influenced by pasture improvement; assessed as Least Concern (LC).
Conservation measures: Attention needs to be paid to taxonomic is-
sues with nomenclature; assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
various nature and game reserves including Suikerbosrand, Abe Bailey,
Ithala, Vernon Crookes.

2 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 451

Onthophagus axillaris
Boheman, 1860

Onthophagus criniger
d’Orbigny, 1904

= Onthophagus crinitus d’Orbigny, 1902

Global: DD

J A S O N D J F M A M J

Type localities: O. axillaris Boheman: ‘in regione fluvii Svakop’ [Swakop

River region, W Central Namibia]; O. criniger, nom. nov. for O. crinitus
(pre-occupied name): ‘Lac Ngami’ [Lake Ngami, N Botswana]. ♂
Taxonomy: Both cited as accepted Group 16 species, but very closely re-
lated; current status of Péringuey’s (1908) synonymy of O. criniger with
O. axillaris unclear; to validate species-level separation, types of both
need to be re-examined, perhaps together with NE African (O. pilicollis
d’Orbigny, 1902) and W Central African relatives (O. hirsutus d’Orbig-
ny, 1902), minor sexual dimorphism (head).
Distribution: O. axillaris: dry to moist savanna: NW Namibia (red
squares) and SW Angola; O. criniger: dry to moist savanna: N Botswana
(black square) and edge of Zambezi Valley in Zimbabwe and Malawi.
Locality data (mean ± SD, range): Combined: altitude: 1 249 ± 461,
1–1 998 m; annual rainfall: 415 ± 276, 9–1 256 mm; annual tempera-
ture (max. + min. /2): 18.8 ± 1.9, 15.8–22.4°C (N=20+3).
Habitat: No quantitative assessment; DBRU collection records on sand
(2), sandy loam (4) in grassland (2), open woodland (4).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (4), elephant (2), zebra (1).
Temporal activity: Diurnal flight activity, primarily during mid- and late
summer rainy seasons (Oct. to June).
Ecoregions Namibia, Botswana: O. axillaris: Namib Desert (AT1315), 2 mm

Namibian Savanna Woodlands (AT1316), NW Kalahari Xeric Savanna

(AT1309), Angolan Mopane Woodlands (AT0702); O. criniger: Kala-
hari Acacia-Baikiaea Woodlands (AT0709).
Assessment rationale: Combined EOO = 393 675 km2; despite close
similarities in appearance, taxonomic uncertainty and differences in
vegetation associations between the two ‘species’ renders it difficult to
assess conservation status; therefore, assessed as Data Deficient (DD),
but populations identified as O. axillaris probably deserve an assess-
ment as Least Concern (LC) as most records are from areas that are
little transformed.
Conservation measures: To generate an assessment of conservation status,
taxonomic issues need to be resolved; quantitative data on ecological
associations would also improve any assessment; populations identified
as O. axillaris protected in Bicuar National Park, Angola; Namibian
populations primarily recorded from farm rangeland.

♀
2 mm

ONTHOPHAGINI
452 SURICATA 6 (2020)

Onthophagus bayeri
Balthasar, 1942

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Süd-West Afrika, Okahandja’ [Okahandja, Namibia].

Taxonomy: Accepted Group 18 species; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size.
Distribution: Arid to dry uplands and highlands: central Namibia.
Locality data (mean ± SD, range): Altitude: 1 520 ± 252, 1 144–1 815 m;
♂ annual rainfall: 240 ± 92, 136–353 mm; annual temperature (max. +
min. /2): 17.6 ± 1.5, 15.8–19.8°C (N=8).
Habitat: Not known.
Food types: No quantitative assessment; reference material recorded from
baits of human dung and meat; a few from banana.
Temporal activity: Diel flight activity unknown; recorded primarily in the
summer rainy season (Oct. to Mar.).
Ecoregions Namibia: NW Kalahari Xeric Savanna (AT1309), S Central
Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 10 885 km2; known from only eight local-
ities across a range of < 20 000 km2; limited records may be related to
food specialisation as close relatives in Group 18 are carrion specialists;
although probably not threatened at present, supporting quantitative
survey data are required; currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, quantitative data
are required to determine the full EOO, AOO and ecological associa-
tions; W edge of range coincides with E edge of Namib-Naukluft Park,
but most records lay outside of protected areas.

2 mm

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 453

Onthophagus beiranus
Péringuey, 1908

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: ‘Mozambique (Beira; Lucitania)’ [Mozambique: Beira

and Búzi].
Taxonomy: Accepted Group 7 species belonging to the new subgenus,
Bicornonthophagus Tagliaferri & Moretto, 2012; needs to be compared
with O. auriculatus Klug, 1855 (type locality: Tete, Mozambique) and
O. insulsus Péringuey, 1901 (type locality: Eshowe, KwaZulu-Natal,
South Africa); sexually dimorphic (head, prothoracic disc), characters ♂
vary in prominence with body size.
Distribution: Disjunct and scattered occurrence on floodplains and in sa-
vanna woodlands and thicket of the Mozambique Coastal Plain owing
to soil type specialisation: South Africa, Mozambique.
Locality data (mean ± SD, range): Altitude: 154 ± 194, 0–1 005 m; an-
nual rainfall: 887 ± 225, 396–1 275 mm; annual temperature (max. +
min. /2): 22.3 ± 1.7, 17.8–25.2°C (N=47).
Habitat: In uMkhuze Game Reserve: strong bias to clay (46.7) compared
to sandy clay loam (2.3), sand over clay (7.3), and deep sand (0.1); sup-
ported by DBRU collection records: clay (11), sandy clay loam (12),
sand (3) on grassland/pasture (11), open woodland/dense woodland
(12), shaded thicket (3).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (13), buffalo (2), wildebeest (2), elephant (4), zebra (2),
waterbuck (2).
Temporal activity: Flight activity in darkness primarily during the rainy
season (Oct. to Apr.), but with some dry season activity across the
warm lowland range.
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
also Zambezian Coastal Flooded Savanna (AT0906), Southern 2 mm
Zanzibar-Inhambane Coastal Forest Mosaic (AT0128) (not shown on
map).
Bioregions South Africa: Lowveld (SVl) (Savanna Biome); N Indian
Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 98 720 km2; disjunct EOO based on
known records; AOO would be smaller coinciding with finer-grained
soils separated by large expanses of sandy soils; however, observed to be
abundant in cattle pads on Pungwe Flood Plain near Muda, Mozam-
bique; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further survey data to determine if known disjunct distri-
bution pattern is a collecting artefact; quantitative data on vegetation
and dung type associations would also be useful; protected in uMkhuze
Game Reserve.

♀
2 mm

ONTHOPHAGINI
454 SURICATA 6 (2020)

Onthophagus bicavifrons
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Afrique orientale allemande: Ouzambara....; lac Ngami..,

Zoulouland..; Natal: Durban’ [Usambara Mts, Tanzania; Lake Ngami,
Botswana; Zululand and Durban, South Africa].
Taxonomy: Accepted Group 18 species, but species cited here as O. bi­
cavifrons requires validation by comparison with type(s); sexually
dimorphic (head, prothoracic disc), characters vary in prominence
♂ with body size; SE coastal variety more densely punctate, but aedeagus
identical (black squares).
Distribution: Widespread in moist savanna from S to E Africa: South
Africa, Namibia, Botswana, Zimbabwe, Angola, Tanzania; also report-
ed from Mozambique, Malawi, Democratic Republic of the Congo
(DRC), Kenya.
Locality data (mean ± SD, range): Altitude: 928 ± 452, 0–1 733 m; an-
nual rainfall: 633 ± 206, 171–1 157 mm; annual temperature (max. +
min. /2): 19.6 ± 2.1, 14.0–25.2°C (N=149).
Habitat: No quantitative assessment; available DBRU records too limited
to determine trends: sand (3), sandy clay loam (1) in grassland/pasture
(3), shrubland (1), forest (1).
Food types: In Botswana: attracted only to carrion (chicken livers: 7), not
to dung (pig, elephant, cattle, sheep); limited DBRU collection records
on dead millipede (4), impala gut contents (1); also dung of horse (1),
cattle (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna Wood-
lands (AT1316), Kalahari Xeric Savanna (AT1309), Kalahari Acacia-
Baikiaea Woodlands (AT0709), Zambezian and Mopane Woodlands
(AT0725), marginal in two other ecoregions.
Bioregions South Africa: Primarily, Indian Ocean Coastal Belt Biome
2 mm
(CB), Central Bushveld (SVcb), Lowveld (SVl), Mopane (SVmp), also
Eastern Kalahari Bushveld (SVk) (Savanna Biome) with scattered re-
cords in five other bioregions.
Assessment rationale: EOO = 4 290 470 km2; extremely widespread,
assuming records to the N represent a single species; trophic records
suggest a carrion specialist whereas the wide scatter of records suggest
habitat generalisation, but support is severely limited; unlikely to face
any threats over most of range; assessed as Least Concern (LC) on basis
of available data.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations and stronger
support for carrion associations; protected in the Kruger National Park
(South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 455

Onthophagus binodis
(Thunberg, 1818)

= Onthophagus columella Fahraeus, 1857

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Copris binodis: ‘majorem horum partem in capite bonae

spei collegi’ [amongst material mostly collected in the Cape of Good
Hope, South Africa]; O. columella: ‘Boschiesmans Rand’ [Bushmans
Ridge = uncertain location].
Taxonomy: Accepted Group 4 species; sexually dimorphic (prothoracic
disc), character varies in prominence with body size. ♂
Distribution: Cool, moist, E Highveld descending to coastal regions of
the Eastern and Western Cape in the S: South Africa, Lesotho; intro-
duced into Australia.
Locality data (mean ± SD, range): Altitude: 1 092 ± 621, 0–2 286 m;
annual rainfall: 669 ± 167, 293–1 101 mm; annual temperature (max.
+ min. /2): 15.9 ± 1.7, 10.6–22.5°C (N=160).
Habitat: No adequate quantitative assessment; at Carolina: little influ-
enced by habitat disturbance, natural Themeda grassland (337), fallow
crop fields (119), improved Kikuyu pasture (276); at Boschberg: much
more abundant in cool Highveld Grassland at 1 676–1 687 m (2 294)
than in warmer lower-altitude grassland at 1 264–1 303 m (348);
DBRU collection records suggest bias to finer-grained soils in grass-
land: sand (9), sandy loam (28), sandy clay loam (45) in grassland/
pasture (61), open shrub/woodland (4), also fallow crop fields (5).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (104), horse (1).
Temporal activity: Diurnal flight activity in spring, summer and autumn
(Sept. to May) reflecting range in winter, bimodal and summer rainfall
regions of South Africa.
2 mm
Bioregions South Africa: Mesic Highveld Grassland (Gm), Sub-
Escarpment Grassland (Gs) (Grassland Biome); S Indian Ocean Coast-
al Belt Biome (CB); mostly transformed grassland habitats in moister
parts of Albany Thicket Biome (AT) plus Eastern Fynbos-Renosterveld
(FO 6), East Coast Renosterveld (FO 8), Southwest Fynbos (FO 2)
bioregions (Fynbos Biome).
Assessment rationale: EOO = 274 485 km2; widespread and abundant
in both natural and transformed farm grasslands; occurrence primarily
in transformed pasture habitats in Fynbos, Renosterveld and Albany
Thicket may indicate range expansion in response to clearance of natu-
ral shrubland; assessed as Least Concern (LC).
Conservation measures: Not currently threatened, but assessment of
conservation status would be improved by further quantitative data on
ecological associations, including tests of grassland versus shrubland as-
sociation in the S; protected in Suikerbosrand Nature Reserve, Golden
Gate National Park and lower-altitude parts of Ukhahlamba Drakens-
berg Park (World Heritage Site).

♀
2 mm

ONTHOPHAGINI
456 SURICATA 6 (2020)

Onthophagus bovinus
Péringuey, 1892

= Onthophagus lobigena d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: O. bovinus: Not stated, but possibly Ovamboland, Na-

mibia; O. lobigena: ‘Natal’ [KwaZulu-Natal, South Africa].
Taxonomy: Accepted Group 7 species; synonymy of O. lobigena should
be re-examined as type locality of ‘Natal’ lies outside known range of
O. bovinus; sexually dimorphic (head, prothoracic disc), characters vary
in prominence with body size.
♂ Distribution: Arid Karoo to dry savanna along a wide band of terrain
encompassing the E, S and W borders of the deep Kalahari sands: Na-
mibia, South Africa, Botswana, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 093 ± 282, 243–1 530 m;
annual rainfall: 416 ± 162, 124–674 mm; annual temperature (max. +
min. /2): 19.8 ± 2.2, 15.5–24.3°C (N=32).
Habitat: No quantitative data; limited DBRU collection records on sand
(4), sandy clay loam (3) in grassland (3), open shrub/woodland (4).
Food types: No quantitative data; limited DBRU collection records on
dung of cattle (3), wildebeest (1), elephant (1), human/cattle mix (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna Wood-
lands (AT1316), Kalahari Xeric Savanna (AT1309), Angolan Mopane
Woodlands (AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709),
Southern African Bushveld (AT0717), Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Bushmanland (NKb), Upper Karoo (NKu)
(Nama Karoo Biome); Kalahari Duneveld (SVkd), Eastern Kalahari
Bushveld (SVk), Central Bushveld (SVcb), Mopane (SVmp) (Savanna
2 mm
Biome); Dry Highveld Grassland (Gh) (Grassland Biome).
Assessment rationale: EOO = 642 681 km2; recorded across a wide area
on a range of dung types in relatively open vegetation on both coarse
and fine-grained soil types; arid part of range little transformed; cur-
rently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
much improved by taxonomic re-examination and quantitative data on
ecological associations; protected in Etosha National Park (Namibia),
Kgalagadi Transfrontier Park (South Africa/Botswana).

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 457

Onthophagus cameloides
d’Orbigny, 1900

= Onthophagus urus Harold, 1867c

Onthophagus quadricallosus
d’Orbigny, 1902

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: O. cameloides: nom nov. for O. urus [pre-occupied name] ♂

‘Cap der guten Hoffnung’; O. quadricallosus: ‘Cap de Bonne-Espérance’
[both Cape of Good Hope, South Africa].
Taxonomy: Accepted Group 30 species, but needs to be re-examined as a pos-
sible senior name for O. quadricallosus, which d’Orbigny described from
a single larger specimen (8.5 mm) that is close to O. cameloides described
from smaller specimens (4.00–5.25 mm); significance of climatic differ-
ences in seasonal activity also requires investigation; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Dry to arid areas in the winter, bimodal and late summer
rainfall regions of South Africa; absence of records from central S
Nama Karoo possibly a collection artefact; claimed records from Java
undoubtedly an error.
Locality data (mean ± SD, range): Altitude: 506 ± 495, 1–1598 m; an-
nual rainfall: 237 ± 98, 63–445 mm; annual temperature (max. + min.
/2): 16.8 ± 1.3, 14.0–20.0°C (N=97).
Habitat: No adequate quantitative assessment; in Western Cape: absent
from cool Cape Peninsula, but on W coast near Darling Hills, more
abundant in pastures (233, 123, 241) than in natural shrubland (sand:
31, sandy loam: 34), rare on coastal deep sands (9, 1, 4); DBRU collec-
tion records on sand (22), sandy loam (10), sandy clay loam (8), clay
2 mm
(1) in grassland/pasture (13), scrub (13), open shrub/woodland (2);
also fallow crop fields (9).
Food types: No quantitative assessment; DBRU collection records from
dung of buffalo (1), cattle (40), sheep (1), horse (6), donkey (1).
Temporal activity: Diurnal flight activity; seasonal activity varies with cli-
mate; in W Coast winter rainfall region: population shows spring to ear-
ly summer activity (July to Dec.); in spring: young, gravid, sclerotised
adults are dominant, but population then ages; in early dry summer:
callow, teneral adults with no follicular development are dominant; in
Northern Cape late summer rainfall region: also recorded Mar. to May.
Bioregions South Africa: Various drier Fynbos and Renosterveld biore-
gions (Fynbos Biome); various bioregions of the Succulent Karoo Bi-
ome; Upper Karoo (NKu), Bushmanland (NKb) (Nama Karoo Biome).
Assessment rationale: EOO = 241 800 km2 (inludes central S Nama
Karoo); widespread association with open vegetation and tolerance of
habitat transformation on various soils types together with ready attrac-
tion to dung of farm livestock suggest limited threats, particularly as
transformation is limited in the N and NE of the range used primarily
for livestock grazing; assessed as Least Concern (LC).
Conservation measures: None recommended although assessment of
conservation status would be improved by quantitative ecological data
and taxonomic revision; protected in Bontebok, West Coast and Nam-
aqua national parks. ♀ 2 mm

ONTHOPHAGINI
458 SURICATA 6 (2020)

Onthophagus cinctipennis
Quedenfeldt, 1884

= Onthophagus patricius Péringuey, 1901

Global: LC

J A S O N D J F M A M J

Type localities: O. cinctipennis: ‘Malange’ [Angola]; O. patricius: ‘Natal

(Estcourt)’ [KwaZulu-Natal, South Africa].
Taxonomy: Accepted Group 2 species; minor sexual dimorphism (head).
Distribution: Disjunct pattern on moist SE coastline and moist SE and
south central African uplands/highlands: N South Africa, E Zimbabwe,
Central Mozambique, W Angola, S Democratic Republic of the Congo
♂ (DRC).
Locality data (mean ± SD, range): Altitude: 1 240 ± 439, 7–2 274 m;
annual rainfall: 761 ± 210, 310–1 659 mm; annual temperature (max.
+ min. /2): 17.6 ± 2.1, 10.6–22.2°C (N=35).
Habitat: In Gauteng bushveld: extreme bias to deep sand (211) compared
to sandy clay loam (0), in grassland (140), open woodland (53), shaded
thickets (18); limited DBRU collection records support open vege-
tation association, grassland/pasture (5), shrubland (1), but not soil
association, sand (1), sandy clay loam (5).
Food types: No quantitative assessment; limited DBRU collection records
all on cattle dung (5).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season, also early dry season (Oct. to June); in Gauteng bushveld:
activity commencing in Oct., peaking in Dec., continuing in late rainy
season (Jan. to Apr.), tailing off into the dry season; in Gauteng bush-
veld: active in the cooler early morning in mid-summer (08:00–10:00,
Nov. to Feb.) moving to a midday flight activity window in cooler late
summer (10:00–14:00, March).
Ecoregions Zimbabwe: Eastern Zimbabwean Forest Grassland Mosaic
(AT1006), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Primarily straddling edges of bioregions at N
2 mm
and NE edge of Highveld: Central Bushveld (SVcb) (Savanna Biome);
Dry Highland Grassland (Gh), Mesic Highland Grassland (Gm),
Sub-Escarpment Grassland (Gs) (Grassland Biome); also Indian Ocean
Coastal Belt Biome (CB).
Assessment rationale: EOO = 217 320 km2 (underestimated); possible
restriction of AOO by soil specialisation requires further investigation
owing to conflicting results; known highland occurrence in South Af-
rica coincides with vegetation units subject to habitat transformation:
25–63% (Gh), 10–50% (Gs); however, widespread in open vegetation
types across several moist highland blocks in southern Africa; therefore,
currently assessed as Least Concern (LC)
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on soil and dung type association as
well as possible effects of habitat transformation; limited protection in
nature reserves, including Suikerbosrand (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 459

Onthophagus cineraceus
d’Orbigny, 1902

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Colonie du Cap.... : Cape Town’ [Western Cape, South

Africa].
Taxonomy: Currently an accepted Group 9 species, but type needs to be
compared with that of O. hyaena (Fabricius, 1801); O. cineraceus could
possibly be a junior synonym; sexually dimorphic (head, prothoracic
disc), characters vary in prominence with body size. ♂
Distribution: Currently known only from SW Western Cape in vicinity
of Cape Town, South Africa, possibly in mountainous terrain; citations
from KwaZulu-Natal in South Africa, Zimbabwe and Mozambique are
errors.
Locality data (mean ± SD, range): Altitude: 571 ± 144, 469–673 m;
annual rainfall: 473 ± 11, 465–480 mm; annual temperature (max. +
min. /2): 14.4 ± 0.2, 14.3–14.6°C (N=2).
Habitat: No quantitative assessment; unknown.
Food types: No quantitative assessment; one individual on jackal dung.
Temporal activity: Not known, but probably diurnal flight activity; re-
corded in the dry season of the winter rainfall region (Dec., Jan.).
Bioregions South Africa: Locality records inaccurate; possibly vegetation
units comprising Peninsula Sandstone Fynbos (FFs 9) and Kogelberg
Sandstone Fynbos (FFs 11) (Fynbos Biome) containing patches of
Southern Afrotemperate Forest (FOz 1).
Assessment rationale: EOO = 180 km2 (gross estimate); currently known
from two localities, one old and inexact (Cape Town), the other more
recent along a mountain hiking trail that passes through fynbos and
segments of old forest (near Stellenbosch); considering the absence of
ecological data, must be assessed as Data Deficient (DD); considering
the rarity in collections and apparent small range in a highly trans-
formed region, may deserve an IUCN threat category if surveys of the
Western and Eastern Cape fail to locate the species elsewhere.
Conservation measures: Assessment of conservation status will only be
possible once the EOO, AOO and ecological associations have been 2 mm

established; to this end, a quantitative survey of mountain fynbos and

forest patches is required during early summer, commencing in the
vicinity of Cape Town; protected in Jonkershoek Nature Reserve.

ONTHOPHAGINI
460 SURICATA 6 (2020)

Onthophagus convexus
d’Orbigny, 1908

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Rhodésia: Salisbury’ [Harare, Zimbabwe].

Taxonomy: Accepted Group 22 species described from a single individual;
at SW edge of range, Karoo individuals shiny, but otherwise similar to
NE populations; however, W populations should be compared with
Onthophagus aspericeps d’Orbigny, 1908, described from a single Group
22 individual recorded at ‘Okahandya’ [Okahandja, Namibia]; descrip-
♂ tion on previous page (148) to that of O. convexus (149); minor sexual
dimorphism (head).
Distribution: Centred on sandy soils across most of southern Africa: Na-
mibia, Botswana, Zimbabwe, South Africa.
Locality data (mean ± SD, range): Altitude: 1 015 ± 207, 278–1 410 m;
annual rainfall: 279 ± 133, 88–835 mm; annual temperature (max. +
min. /2): 19.3 ± 1.6, 14.7–23.7°C (N=183).
Habitat: In Gauteng bushveld: extreme association with deep sand (195)
compared to sandy clay loam (0) with a bias to grassland (148) com-
pared to open woodland (47) or shaded thickets (0) (cited as Ontho­
phagus sp. nr pullus (a)); limited DBRU collection records on sand (4),
sandy loam (2) in grassland (2), open shrub/woodland (4).
Food types: In Botswana: abundant on all dung types with a bias to omni-
vore dung (pig: 5 912, elephant: 2 877, cattle: 1 751, sheep: 1 931), rel-
atively uncommon on carrion (30) (cited as Onthophagus sp. nr pullus
(a)); limited DBRU collection records on dung of cattle (2), elephant
(1), donkey (1), human/cattle mix (1).
Temporal activity: Diurnal flight activity during the summer rainy sea-
son (Oct. to Mar.); on Gauteng bushveld: primarily active early to
mid-summer (Oct. to Jan.) becoming rare late summer (Feb., Mar.).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric savanna
(AT1309), Namibian Savanna Woodlands (AT1316), Angolan Mo-
pane Woodlands (AT0702), Kalahari Acacia-Baikiaea Woodlands
(AT0809), Zambezian and Mopane Woodlands (AT0725), Southern
African Bushveld (AT0717), Southern Miombo Woodlands (AT0719).
1 mm
Bioregions South Africa: Sandy soils in Kalahari Duneveld (SVkd),
Eastern Kalahari Bushveld (SVk), Central Bushveld (SVcb), Mopane
(SVmp), Lowveld (SVl) (Savanna Biome); Bushmanland (NKb), NW
Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 988 530 km2; AOO would be smaller, but,
nevertheless, widespread across sandy regions; much of range lies in
little-transformed areas used primarily for grazing of domestic livestock
to which the species is readily attracted; despite some taxonomic un-
certainty, assessed as Least Concern (LC) as no populations apparently
face threats.
Conservation measures: Resolution of Group 22 taxonomic uncertain-
ties is required to improve assessment of conservation status; protected
in Kgalagadi Transfrontier Park and Kruger National Park (Botswana,
South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 461

Onthophagus corniculiger
d’Orbigny, 1913

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Afrique orientale portugaise: Zambézie’ [Zambezia Prov-

ince, central Mozambique].
Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus), but
described from a single specimen; should be compared to type material
of O. decedens Péringuey, 1904, (not located) described from ‘Southern
Rhodesia (Sebakwe)’ [Zimbabwe]; like most Group 12 species O. cor­
niculiger is sexually dimorphic (head, prothoracic disc); as characters ♂
show great morphological variability according to body size, a review of
the group taxonomy would be useful.
Distribution: Most known records from reserves in moist savanna on the
S Mozambique Coastal Plain (Mozambique, South Africa), but also
from bushveld in NW South Africa; citation from Zambia requires
validation.
Locality data (mean ± SD, range): Altitude: 450 ± 402, 55–1 448 m;
annual rainfall: 629 ± 120, 435–900 mm; annual temperature (max. +
min. /2): 21.7 ± 1.7, 18.2–23.6°C (N=14).
Habitat: No quantitative assessment; sampled in small numbers on both
sandy granite-derived and finer gabbro-derived soils in very open
woodland at South African Wildlife College (Kruger National Park).
Food types: No quantitative assessment; DBRU collection records all
from elephant dung (4).
Temporal activity: Diel flight periodicity unknown, but probably in dark-
ness like some other Group 12 species; recorded during the summer
rainy season (Nov. to Mar.).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
2 mm
Assessment rationale: EOO = 145 035 km2 (gross estimate); AOO would
be much smaller owing to specialisation to dung of large monogas-
tric herbivores whose range is severely contracted within SE Africa;
elephants remain protected across circa 20% of the putative range for
O. corniculiger; unclear if AOO would also be influenced by habitat
associations as these are essentially unknown; currently assessed as Data
Deficient (DD).
Conservation measures: Conservation of this species would be dependent
on protection of elephants; to adequately assess conservation status, the
current EOO and AOO need to be more precisely determined by a
survey of elephant dung in game reserves of SE Africa; quantitative data
on habitat associations is also required; protected in Kruger National
Park, uMkhuze and Hluhluwe–iMfolozi game reserves (South Africa).

2 mm

ONTHOPHAGINI
462 SURICATA 6 (2020)

Onthophagus cretus
Péringuey, 1901

See Taxonomy section.

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Natal (Durban, Umkomas River)’ [Durban, KwaZulu-

Natal, South Africa]; Also ‘Ovampoland (Humbe)’ [Ovamboland,
Namibia – considered an error].
Taxonomy: Group 23 species currently synonymised with Onthophagus
setosus Fahraeus, 1857, described from ‘prope fluvium Limpopo’ [near
♂ Limpopo River]; this type locality lies outside of the range of the species
treated here as O. cretus, which may, therefore, deserve revalidation as a
‘bona fide’ species; however, further study required; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Primarily SE coastal and SE lower escarpment forest patch-
es in South Africa; validated as same species (red squares), tentative
(black square).
Locality data (mean ± SD, range): Altitude: 586 ± 459, 0–1 431 m; an-
nual rainfall: 876 ± 111, 482–1 015 mm; annual temperature (max. +
min. /2): 18.0 ± 1.8, 14.2–20.5°C (N=21).
Habitat: No quantitative assessment; 78.6% (22) of available records (28)
cited from forest; no soil type records.
Food types: No quantitative assessment; no records.
Temporal activity: Diel flight periodicity unknown; active primarily
during the summer rainy season (Aug. to Apr.).
Bioregions South Africa: Locality records primarily from patches of
Southern Mistbelt Forest (FOz 3) and Scarp Forest (FOz 5) within
the Indian Ocean Coastal Belt Biome (CB) and E edge of Grassland
Biome (G).
2 mm Assessment rationale: EOO = 44 150 km2; AOO would be much smaller
due to forest patch specialisation; often comparatively abundant; with-
in reserves, FOz 3 and FOz 5 are assessed as least threatened although
uncontrolled exploitation is a problem; O. cretus is currently assessed
as Least Concern (LC) owing to widespread occurrence in many forest
reserves.
Conservation measures: Taxonomic issues of species validity need to
be resolved. Assessment of conservation status would be improved by
quantitative data on ecological associations, particularly food and soil
type; protected in various forest reserves, including Dwesa, Ntsubane
and Silaka (Eastern Cape).

2 mm ♀

ONTHOPHAGINI
SURICATA 6 (2020) 463

Onthophagus cribripennis
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Afrique austral: Zoulouland’ [Zululand, South Africa].

Taxonomy: Accepted Group 17 species, but populations recorded primar-
ily in forest along SE coast require re-assessment (black squares); male
aedeagus of these populations identical to typical O. cribripennis, but
ridge on vertex of female somewhat arcuate rather than straight; sexu-
ally dimorphic (head, prothoracic disc), characters vary in prominence
with body size. ♂
Distribution: Primarily warm, moist woodland savanna and forest plus
moist coastal and upland grasslands of SE Africa: South Africa, Mo-
zambique, Zimbabwe; also cited from Malawi.
Locality data (mean ± SD, range): Altitude: 1 122 ± 521, 0–2 286 m;
annual rainfall: 800 ± 162, 305–1 659 mm; annual temperature (max.
+ min. /2): 17.5 ± 2.2, 10.6–22.7°C (N=146).
Habitat: At 13–1 400 m on Wolkberg: more abundant in forest (21.1)
than grassland (2.7); at Carolina in Dec.: more abundant in improved
Kikuyu pasture (25.8) and regenerating vegetation of fallow crop fields
(12.7) than in natural Themeda grassland (4.9); DBRU collection re-
cords suggest a bias to finer-grained soils (sand: 6, sandy loam: 17,
sandy clay loam: 28, clay: 4) although most recorded in pasture/grass-
land (43) or forest (8), also shrub/woodland (5).
Food types: At 13–1 400 m on Wolkberg: more abundant on pig dung
(16.2) than cattle dung (7.6); DBRU collection records primarily from
dung of cattle (50), also baboon (1), human (1), human/cattle or buf-
falo mix (8), rhinoceros (1), zebra (1).
Temporal activity: Diurnal flight activity during much of the year (Aug.
to May). 2 mm

Ecoregions Zimbabwe: Primarily Southern Miombo Woodlands

(AT0719), Eastern Zimbabwe Grassland Forest Mosaic (AT1006).
Bioregions South Africa: Straddling edges of Central Bushveld (SVcb)
and Lowveld (SVl) in Soutpansberg and Waterberg (Savanna Biome);
N and E edges of Mesic Highveld Grassland (Gm), N Sub-Escarpment
Grassland (Gs) (Grassland Biome); Indian Ocean Coastal Belt Biome
(CB); warmer S coast forest patches.
Assessment rationale: EOO = 206 820 km2; AOO may be influenced by
soil type bias, but quantitative support required; comparatively wide-
spread and abundant in both reserves and farm rangeland; abundance
apparently greater in improved pasture than in natural grassland; there-
fore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative ecological data to determine soil asso-
ciations and resolve conflicts in vegetation and dung associations; pro-
tected in Ithala and Hluhluwe–iMfolozi game reserves (South Africa).

♀ 1 mm

ONTHOPHAGINI
464 SURICATA 6 (2020)

Onthophagus croesulus
Bates, 1888

= Onthophagus obtusicollis Péringuey, 1888

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: O. croesulus: ‘Natal’ [KwaZulu-Natal, South Africa];

O. obtusicollis: ‘Potchefstroom and Pretoria, Transvaal’ [South Africa:
North-West Province and Gauteng].
Taxonomy: Accepted Group 18 species; limited sexual dimorphism (head,
prothoracic disc) varying in prominence with body size.
♂ Distribution: Patchy records from moist savanna and grasslands in N and
E South Africa; patchiness probably a collection artefact due to dietary
specialisation.
Locality data (mean ± SD, range): Altitude: 1 113 ± 323, 305–1 553 m;
annual rainfall: 680 ± 132, 525–1 022 mm; annual temperature (max.
+ min. /2): 18.4 ± 2.1, 15.4–22.5°C (N=13).
Habitat: No quantitative assessment; sampled in open woodland on kop-
pies in Telperion Nature Reserve (2); one DBRU collection record from
sandy clay loam in grassland.
Food types: No quantitative assessment; sampled to pig dung bait in Tel-
perion Nature Reserve (2); one DBRU collection record from a dead
millipede.
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl)
(Savanna Biome); N and E edges, Mesic Highveld Grassland (Gm)
(Grassland Biome) adjoining the savanna bioregions; also E edge, Sub-
Escarpment Grassland (Gs) adjoining Sub-Escarpment Savanna (SVs).
Assessment rationale: EOO = 128 590 km2; no quantitative data, but
probably a carrion specialist like other members of Group 18 Ontho­
phagus; recorded over a wide range including in grassland where carrion
would be readily available; probably not threatened, however, ecologi-
2 mm
cally poorly known, so assessed as Data Deficient (DD).
Conservation measures: Before an assessment may be made of conserva-
tion status, quantitative data on ecological associations are required; it
is also necessary to conduct a survey to determine the full EOO and
AOO; protected in Nascott and Telperion nature reserves.

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 465

Onthophagus cupricollis
Péringuey, 1888

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Lydenburg, Transvaal’ [Mashishing, Mpumalanga, South

Africa].
Taxonomy: Accepted Group 18 species; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size.
Distribution: South Africa: moist upland savanna on Waterberg (Limpo-
po and Mpumalanga provinces) and NE escarpment. ♂
Locality data (mean ± SD, range): Altitude: 1 228 ± 296, 603–1 627 m;
annual rainfall: 762 ± 127, 563–1 022 mm; annual temperature (max.
+ min. /2): 17.6 ± 2.0, 14.7–20.7°C (N=11).
Habitat: Not known.
Food types: No quantitative assessment; trapped to meat bait.
Temporal activity: Attracted to light, so presumably flies in darkness;
recorded primarily in the summer rainy season (Oct. to Mar.).
Bioregions South Africa: Moist uplands of Central Bushveld (SVcb) and
Lowveld (SVl) (Savanna Biome) plus NE extremity of Mesic Highveld
Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 26 645 km2; poorly known, but attraction
to meat bait suggests carrion associations like other close relatives in
Group 18; AOO may be restricted to moist savanna regions of central
N South Africa, but range probably somewhat underestimated; cur-
rently assessed as Data Deficient (DD).
Conservation measures: Before an assessment of conservation status
may be made, a quantitative survey is required to determine the full
EOO, AOO and ecological associations; the survey should be centred
on moist upland savanna along the Waterberg and NE escarpment;
protected in Mabalingwe Game Reserve and Nelspruit Nature Reserve.
2 mm

♀
2 mm

ONTHOPHAGINI
466 SURICATA 6 (2020)

Onthophagus cyaneoniger
d’Orbigny, 1902

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA (but see Distribution)

J A S O N D J F M A M J

Type localities: ‘Transvaal: Hamman’s Kraal près de Pretoria..... Bechoua-

na: Vryburg.....; État d’Orange: Bloemfontein....; Natal’ [South Africa:
Hammanskraal, Vryburg, Bloemfontein, KwaZulu-Natal].
Taxonomy: Accepted Group 23 species; sexually dimorphic (head), char-
acters vary in prominence with body size.
♂ Distribution: Centred on the grasslands of the South African Highveld
with peripheral records in the Northern Cape, highlands of Upper
Karoo and transformed Eastern Cape coastal habitat; disjunct pattern
may reflect soil specialisation, a collection artefact related to small body
size, or both; citations from Botswana, Zimbabwe, Democratic Repub-
lic of the Congo (DRC) require validation as they are likely errors.
Locality data (mean ± SD, range): Altitude: 1 411 ± 240, 290–1 860 m;
annual rainfall: 590 ± 168, 308–1 022 mm; annual temperature (max.
+ min. /2): 16.1 ± 1.5, 13.2–19.5°C (N=42).
Habitat: No adequate quantitative assessment, but recorded in abundance
in grassland on sand (24.3/sample, Telperion NR) and sandy loam
(65.0/sample, Abe Bailey NR), but uncommon on sandy clay loam
(0.5/sample, Suikerbosrand NR); DBRU collection records on sand
(1), sandy loam (6), sandy clay loam (1) in grassland/pasture (6), also
fallow crop field (1).
Food types: No quantitative assessment; sampled to pig dung and to com-
bined sheep/cattle dung bait; DBRU collection records exclusively on
cattle dung (13).
Temporal activity: Diel flight periodicity unknown, but probably diurnal;
1 mm
active in the summer rainy season (Oct. to May).
Bioregions South Africa: Dry Highveld Grassland (Gh), Mesic Highveld
Grassland (Gm) (Grassland Biome), E Upper Nama Karoo (NKu)
(Nama Karoo Biome); also in Eastern Kalahari Bushveld (SVk) (Sa-
vanna Biome).
Assessment rationale: EOO = 181 630 km2; AOO may be smaller due to
possible bias to sandier soils although this requires rigorous quantitative
support as the data cited above might be influenced by climate; range
is subject to extensive transformation, but recorded in cattle dung on
farms over a wide range; also abundant in some Highveld nature re-
serves; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, including effects
of pasture improvement or habitat modification; protected in Telperi-
on, Suikerbosrand, Abe Bailey nature reserves (Gauteng).

1 mm

ONTHOPHAGINI
SURICATA 6 (2020) 467

Onthophagus deterrens
Péringuey, 1901

Onthophagus variolosus
d’Orbigny, 1902

No synonyms
Global: DD
Endemic: RSA (but see Taxonomy section)

J A S O N D J F M A M J

Type localities: O. deterrens: ‘Cape Colony (Stellenbosch, in carrion; ♂

Paarl), Natal (Durban), Transvaal (Lydenburg), Southern Rhodesia
(Salisbury)’ [South Africa; Zimbabwe]; O. variolosus: ‘Colonie du Cap:
Cape Town’ [Cape Town, Western Cape, South Africa].
Taxonomy: Both accepted Group 19 species, but status needs validation
or revision after comparison between types; as aedeagi are identical,
O. variolosus may be a junior synonym and blue colour variety of metal-
lic green, O. deterrens; cited type localities for O. deterrens (SW Cape to
Zimbabwe) suggest inclusion of two or three species; in two, the vertex
of females bears a spine (O. deterrens, O. variolosus), in one, it bears a
ridge (Onthophagus sp. nr deterrens).
Distribution: O. variolosus (black squares): S Cape and NW Lesotho bor-
der, South Africa; O. deterrens (red squares): NE Highveld and E coast,
South Africa; O. sp. nr deterrens (turquoise squares): N savanna, South
Africa to Zimbabwe.
Locality data (mean ± SD, range): O. variolosus: altitude: 820 ± 630,
26–1 830 m; annual rainfall: 519 ± 150, 270–879 mm; annual tem- 2 mm
perature (max. + min. /2): 15.7 ± 1.6, 13.3–17.8°C (N=16). O. de­
terrens: altitude: 1 275 ± 463, 28–2 274 m; annual rainfall: 793 ±
156, 338–1 271 mm; annual temperature (max. + min. /2): 16.6 ±
2.1, 10.6–20.5°C (N=55). O. sp. nr deterrens: altitude: 1 119 ± 386, O. deterrens
58–1 746 m; annual rainfall: 719 ± 115, 533–1 013 mm; annual tem-
perature (max. + min. /2): 18.4 ± 1.8, 14.0–22.6°C (N=42).
Habitat: No quantitative assessment; limited DBRU records for species
complex from finer-grained soils: sandy loam (1), sandy clay loam (2)
in pasture/grassland (2).
Food types: No quantitative assessment; limited DBRU records for species
complex from carnivore dung (dog; 2) or dead animals (toad, millipede
and tenebrionid beetle; 3).
Temporal activity: Diel flight periodicity unknown; species complex ac-
tive throughout summer rainy season (Sept. to Apr.).
Bioregions South Africa: O. variolosus: S Mesic Highveld Grassland
(Gm) (Grassland Biome), mainly Renosterveld vegetation units (Fyn-
bos Biome); O. deterrens: primarily Sub-Escarpment Grassland (Gs) in
SE plus N and NE edge of Mesic Highveld Grassland (Gm) (Grassland
Biome), also Indian Ocean Coastal Bet Biome (CB); O. sp. nr deterrens:
primarily Central Bushveld (SVcb) and moister S and W of Lowveld
(SVl) (Savanna Biome).
Assessment rationale: EOO = 58 525 km2, 293 240 km2, 111 545 km2,
respectively, O. variolosus, O. deterrens, O. sp. nr deterrens; owing to 2 mm
localities cited in original description any one of the three could be
the ‘real’ O. deterrens; thus, impossible to assess conservation status ♀
although wide range and association with carnivore dung or carrion
O. deterrens

ONTHOPHAGINI
468 SURICATA 6 (2020)

Onthophagus deterrens & O. variolosus (continued)

♂ suggests no member of the species complex is threatened; currently

assessed as Data Deficient (DD).
Conservation measures: Uncertainties in species status and nomencla-
ture need to be resolved before conservation status may be assessed;
quantitative data on ecological associations are also required; protected
in various nature reserves: Coleford, Abe Bailey, Suikerbosrand (O. de­
terrens); Roodeplaat, Rustenburg (O. sp. nr deterrens).

2 mm

O. variolosus

ONTHOPHAGINI
SURICATA 6 (2020) 469

Onthophagus discretus
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Transvaal (Rustenburg)’ [Rustenburg, North-West Prov-

ince, South Africa].
Taxonomy: Accepted Group 7 species; minor sexual dimorphism (head).
Distribution: Disjunct, scattered pattern in SE Africa owing to speciali-
sation to moist, shaded, upland vegetation: South Africa, Zimbabwe;
also cited from Malawi.
Locality data (mean ± SD, range): Altitude: 1 166 ± 158, 973–1 553 m;
annual rainfall: 648 ± 51, 563–745 mm; annual temperature (max. +
min. /2): 19.0 ± 1.5, 15.4–21.9°C (N=13).
Habitat: In Gauteng bushveld: recorded on both deep sand (13) and sandy
clay loam (36), but with extreme bias to shaded thickets (49) compared
to open woodland (0) and grassland (0); similar at Roo­deplaat Nature
Reserve on sandy clay loam: shaded thicket (12), open woodland (1),
grassland (0); a single DBRU collection record from shaded Brachyste­
gia forest.
Food types: No quantitative assessment; a single DBRU collection record
from human/cattle dung mix.
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Mar.); in Gauteng bushveld: peak activity early to mid-
rainy season (Oct. to Dec.).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Primarily Central Bushveld (SVcb) (Savanna
Biome).
Assessment rationale: EOO = 42 255 km2 (underestimated); AOO much
smaller owing to association with shaded thicket and dense woodland;
would be susceptible to clearance of dense woody vegetation although
also recorded in eucalypt plantation; transformation in vegetation units
of known range in South Africa, 3–49%; currently assessed as Least
Concern (LC) pending further research.
Conservation measures: Assessment of conservation status would be im-
proved by a survey of the surviving dense woody vegetation in upland
2 mm
SE Africa to better determine the EOO, AOO as well as dung type
associations on both deep sand and finer-grained soils; protected in
Roodeplaat and Nascott nature reserves (South Africa), Lake Mutirik-
we Game Reserve (Zimbabwe).

ONTHOPHAGINI
470 SURICATA 6 (2020)

Onthophagus ebenicolor
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Afrique orientale allemande...., Zambèze’ [Tanzania; ?N

Mozambique].
Taxonomy: Accepted Group 19 species, but distribution data requires
validation owing to a number of putative close relatives of similar ap-
pearance; sexually dimorphic (head), character varies in prominence
with body size.
♂ Distribution: Dry to moist savanna from southern to E Africa: South Af-
rica, Namibia, Botswana, Zimbabwe, Mozambique, Zambia, Tanzania;
also reported from Kenya.
Locality data (mean ± SD, range): Altitude: 716 ± 445, 1–1 850 m; an-
nual rainfall: 619 ± 205, 162–1 177 mm; annual temperature (max. +
min. /2): 20.8 ± 2.0, 13.4–25.2°C (N=69).
Habitat: No quantitative assessment; limited DBRU collection records on
sand (1), sandy clay loam (2), clay (2) in pasture (1), shrub/woodland
(4) or forest (1).
Food types: No quantitative assessment; limited DBRU collection records
on gut content of impala (2), human/buffalo or cattle dung mix (2);
also dung of buffalo (1) or cattle (1); also see Assessment rationale.
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to May); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna
Woodlands (AT1316), Angolan Mopane Woodlands (AT0702), Ka-
lahari Acacia-Baikiaea Woodlands (AT0709), Zambezian and Mopane
Woodlands (AT0725), Southern African Bushveld (AT0717).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl),
Mopane (SVmp) (Savanna Biome); Indian Ocean Coastal Belt Biome
(CB); single records in two other ecoregions.
2 mm
Assessment rationale: EOO = 1 691 195 km2; AOO unclear; ecolog-
ically poorly known owing to lack of quantitative data, but Group
19 is thought to be characterised primarily by carrion associations;
habitat associations possibly biased to finer-grained soils and woody
vegetation; however, probably widespread; therefore, assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status needs to be
improved by quantitative ecological data to confirm bias in habitat
associations and bias in food associations with omnivore dung and/
or carrion specialisation; protected in Kruger National Park (South
Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 471

Onthophagus ebenus
Péringuey, 1888 (but see Taxonomy)

= Onthophagus natalicus d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: O. ebenus: ‘From the banks of the Vaal River’ [inexact,
Vaal River, N South Africa]; O. natalicus: ‘Natal...: Durban....; Trans-
vaal: Makapan près de Pietersburg’ [South Africa: Durban and Maka-
pan, KwaZulu-Natal and Limpopo provinces].
Taxonomy: Accepted Group 24 species; sexually dimorphic (head protho-
racic disc), characters vary in prominence with body size; subsequent to
completion of the manuscript for this book, this Group 24 species was ♂
placed in a new genus, Tiaronthophagus Roggero, Moretto, Palestrini
& Barbero, 2019, comprising 26 species from three different groups.
Distribution: Moist coastal and upland open savanna in SE Africa: South
Africa, N Botswana, Zimbabwe, Mozambique; also reported from Ma-
lawi, SE Tanzania.
Locality data (mean ± SD, range): Altitude: 953 ± 522, 0–1 824 m; an-
nual rainfall: 704 ± 147, 205–1 254 mm; annual temperature (max. +
min. /2): 19.3 ± 2.2, 14.2–25.2°C (N=123).
Habitat: No clear soil specialisation in Gauteng bushveld: deep sand (5),
sandy clay loam (11), or in uMkhuze Game Reserve: sand (0.1), sand
over clay (0.3), sandy clay loam (0.8), clay (0.3); at Roodeplaat Nature
Reserve: bias to open vegetation, grass (32), open woodland (20), shad-
ed thicket (2); limited DBRU collection records on sand (1), sandy
loam (3), sandy clay loam (3), clay (1).
Food types: In Gauteng bushveld: extreme bias to omnivore dung (pig,
12) and carrion (14) compared to dung of ruminant herbivore (cattle,
0) and monogastric herbivore (horse, 0); bias supported by limited
DBRU collection records from human/cattle or buffalo dung mix (5),
cattle dung (3).
Temporal activity: Flight activity in darkness; recorded during most
months of the year (Aug. to May) although primarily active during the 2 mm
summer rainy season (Oct. to Mar.); attracted to light.
Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725), Southern African Bushveld (AT0717), Southern Miombo
Woodlands (AT0719).
Bioregions South Africa: Primarily Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome) and Indian Ocean Coastal Belt Biome (CB);
also N margins of Grassland Biome (3 bioregions).
Assessment rationale: EOO = 847 490 km2; widespread omnivore dung
and carrion specialist showing a bias to open vegetation; possibly toler-
ant of urbanisation as it has been recorded on dog dung in Pretoria city
suburbs; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by stronger data to support soil generalisation or a slight bias to
finer-grained soils; protected in Kruger National Park, Leeuwfontein
and Abe Bailey nature reserves (South Africa).

♀
2 mm

ONTHOPHAGINI
472 SURICATA 6 (2020)

Onthophagus fimetarius
Roth, 1851

= Onthophagus piceus Fahraeus, 1857

= Onthophagus lugens Fahraeus, 1857, sensu Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: O. fimetarius: ‘Abyssinien’ [inexact, Ethiopia]; O. piceus:

‘prope fluvium Limpopo’ [near Limpopo River, southern Africa];
O. lugens sensu Péringuey: ‘Cape Colony (Kimberley), Natal (Newcas-
tle, Frere, Durban), Transvaal (Potchefstroom, Rustenburg), Southern
Rhodesia (Salisbury, Upper Hanyani River, Victoria Falls)’ [South Af-
♂ rica, Zimbabwe].
Taxonomy: Accepted Group 23 species, but revision required as species of
similar appearance have been variously synonymised or revalidated by
past researchers; synonymy of southern African O. piceus with Ethiopi-
an O. fimetarius requires further validation; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size.
Distribution: Cited from savanna throughout the Afrotropical region:
single species composition validated by extraction of aedeagi for South
Africa, Namibia, Angola, Zimbabwe, Mozambique, Malawi, Tanzania,
Kenya (black squares on map); also cited from Lesotho, Eswatini, Bot­
swana, Democratic Republic of the Congo (DRC), Uganda, Sudan,
Somalia, Ethiopia, Eritrea and 15 W African countries.
Locality data (mean ± SD, range): Altitude: 1 069 ± 453, 0–2 286 m;
annual rainfall: 683 ± 198, 120–1 415 mm; annual temperature (max.
+ min. /2): 18.8 ± 2.4, 10.6–24.8°C (N=199).
Habitat: On Gauteng bushveld: strong bias to deep sand (1 385) com-
pared to sandy clay loam (221), but a vegetation generalist (grassland:
491, open woodland: 563, shaded thickets: 532); at Carolina in Jan.:
much more abundant in natural Themeda grassland (71) compared to
regenerating vegetation of fallow crop fields (10) or improved Kikuyu
grass pasture (8); DBRU collection records from sand (19), sandy loam
2 mm (28), sandy clay loam (19), clay (4) in pasture/grassland (45), shrub­
land/open woodland (32), shaded thicket (1),
Food types: On Gauteng bushveld: primarily attracted to omnivore dung
(pig: 73), but also to herbivore dung (cattle: 29, horse: 15), rarely to
carrion (2); DBRU collection records on dung of cattle (70), wildebeest
(2), elephant (1), human (1), human/buffalo or cattle mix (6).
Temporal activity: Flight activity during darkness in the summer rainy sea-
son (Aug., Oct. to May); on Gauteng bushveld: primarily active in early
summer (Oct. to Dec.) tailing off from mid-summer into early winter
(Jan. to May); also on Gauteng bushveld: flies primarily early evening
(18:00–20:00) in early (Nov.) and late summer (Feb., Mar.), but also at
dawn in warmer mid-summer (Jan., 04:00–06:00), attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: N Namibian Savanna
Woodlands (AT1316), Kalahari Xeric Savanna (AT1309), Angolan
Mopane Woodlands (AT0702), Zambezian and Mopane Woodlands
(AT0725), Zambezian Baikiaea Woodlands (AT0726), Southern Af-
rican Bushvelld (AT0717), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); N Grassland Biome (Dry Highveld
Grassland (Gh), Mesic Highveld Grassland (Gm), Sub-Escarpment
Grassland (Gs)); Indian Ocean Coastal Belt Biome (CB).

ONTHOPHAGINI
SURICATA 6 (2020) 473

Assessment rationale: EOO = 5 546 895 km2 (gross estimate based on

range from S to NE Africa); extremely wide range with generalist
vegetation associations; possible soil bias and intolerance of habitat
disturbance requires further validation, assessed as Least Concern (LC).
Conservation measures: Taxonomic revision is required to validate
nomenclature used for a species that is ecologically well known in N ♀
South Africa; protected in many reserves including Kruger National
Park (South Africa), Serengeti National Park (Tanzania).

2 mm

ONTHOPHAGINI
474 SURICATA 6 (2020)

Onthophagus flavolimbatus
Klug, 1855

= Onthophagus bicolor Raffray sensu d’Orbigny, 1900

= Onthophagus semiflavus Boheman, 1857, sensu d’Orbigny, 1902
= Onthophagus breviculus d’Orbigny, 1905
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. flavolimbatus: ‘Sena’ [central Mozambique]; O. bicolor

sensu d’Orbigny: ‘Afrique allemande orientale: Bagamoyo’ [Tanzania];
O. semiflavus sensu d’Orbigny: ‘Transvaal’ [South Africa]; O. breviculus:
‘Transvaal: Hamman’s Kraal’ [Hammanskraal, South Africa].
♂ Taxonomy: Accepted Group 11 species; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size.
Distribution: Mostly, dry, lower-altitude savanna on the E seaboard from
S to E Africa: South Africa, N Botswana, Zimbabwe, Mozambique,
Kenya, Tanzania.
Locality data (mean ± SD, range): Altitude: 620 ± 419, 5–1 670 m; an-
nual rainfall: 627 ± 180, 304–1 275 mm; annual temperature (max. +
min. /2): 21.3 ± 1.9, 14.5–25.7°C (N=72).
Habitat: In Gauteng bushveld: bias to coarse-grained soils in open vegeta-
tion, sand (462), sandy clay loam (64), grassland (267), open woodland
(254), shaded thicket (5); in uMkhuze Game Reserve: similar bias,
sand (3.1), sand on clay (6.1), sandy clay loam (0.1), clay (1.3); DBRU
collection records less supportive, sand (8), sandy loam (10), sandy clay
loam (7), clay (2) in pasture (2), shrubland (7), open woodland (17).
Food types: Relative dung generalist in Gauteng bushveld, but lower on
monogastric herbivore dung: cattle (30), horse (19), pig (29), carrion
(1); similar at Phalaborwa: pig (1 001), cattle (1 061), elephant (351);
DBRU collection records from dung of cattle (20), buffalo (1), wilde-
1 mm beest (1), elephant (1), horse (1), donkey (2), human/cattle mix (2).
Temporal activity: Diurnal flight activity; in Gauteng bushveld: peaking
between 10:00–14:00 (Nov. to Feb.), tailing off to 16:00; active in the
summer rainy season (Oct. to Apr.); in Gauteng bushveld: peaking Oct.
to Feb., tailing off early Mar.; at Phalaborwa: more abundant in warm
sunny conditions after heavy rainfall (1 391) than in hot, sunny, dry
conditions (794), or warm cloudy conditions after light rainfall (55).
Ecoregions Botswana, Zimbabwe: Kalahari Acacia-Baikiaea Woodands
(AT0709), Southern African Bushveld (AT0717), Zambezian and Mo-
pane Woodlands (AT0725).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl), Mopane (SVmp) (Savanna Biome);
also N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 860 785 km2 (gross estimate); AOO
possibly smaller due to sandy soil bias; however, widespread in both
grassland and wooded habitats where it shows relatively generalist dung
associations; assessed as Least Concern (LC).
Conservation measures: None required; protected in various reserves in-
cluding Kruger National Park, uMkhuze Game Reserve (South Africa),
Gorongosa National Park (Mozambique).

1 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 475

Onthophagus fritschi
d’Orbigny, 1902

No synonyms
Global: LC
Endemic: RSA (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Afrique australe’ [southern Africa].

Taxonomy: Accepted Goup 31 species (subgenus Furconthophagus); sexu-
ally dimorphic (head), characters vary greatly in prominence with body
size.
Distribution: Mostly dry upland Karoo and grassland of central South
Africa and lower-altitude, dry areas inland from the S coast. ♂
Locality data (mean ± SD, range): Altitude: 1 333 ± 292, 210–2 199 m;
annual rainfall: 413 ± 142, 221–741 mm; annual temperature (max. +
min. /2): 15.4 ± 1.3, 11.2–18.6°C (N=80).
Habitat: No adequate quantitative data; in Northern Cape: almost ex-
clusively in Upper Karoo (frequency: 45.6%; 31 out of 68 samples),
soil data inconclusive: sand (10.2 per sample, N=18), sandy loam (5.6,
N=11) but 173.8 per sample in a moist, outlier clay vlei at arid W
margins of range; DBRU collection data from sand (8), sandy loam
(15), sandy clay loam (10), clay (3) in grassland/pasture (17), scrub/
shrubland (9).
Food types: No quantitative data; DBRU collection records primarily
from dung of cattle (43), also sheep (1), donkey (1), zebra (1), human/
cattle mix (1).
Temporal activity: Diel flight periodicity unknown, but presumably diur-
nal activity in summer rainy season (Oct. to May).
Bioregions South Africa: Centred on S Dry Highveld Grassland (Gh)
and SW margins of three other bioregions (Grassland Biome); also
Upper Karoo (NKu), Lower Karoo (NKl) (Nama Karoo Biome) and 1 mm
Albany Thicket Biome (AT).
Assessment rationale: EOO = 174 610 km2; available data suggest a soil
generalist in open grassland and open shrubland vegetation, but quan-
titative support required; widely recorded from dung of farm livestock
across a range that is little transformed to the W and only 10–12%
transformed to the NE; currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations and more precise
measurement of EOO as regards E edge of range, which approaches
the range of the very closely related O. lamnifer d’Orbigny, 1902; not
currently known from any protected area.

♀ 1 mm

ONTHOPHAGINI
476 SURICATA 6 (2020)

Onthophagus fugitivus
Péringuey, 1901

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal (Klerksdorp)’ [North-West Province, South Af-

rica].
Taxonomy: Accepted Group 31 species (subgenus Furconthophagus); sex-
ually dimorphic (head), characters vary in prominence with body size.
Distribution: Uplands and adjoining lowlands, N South Africa; patchy
♂ occurrence possibly related to soil specialisation; record from Botswana
requires validation.
Locality data (mean ± SD, range): Altitude: 1 042 ± 372, 286–1 553 m;
annual rainfall: 607 ± 84, 422–801 mm; annual temperature (max. +
min. /2): 19.2 ± 1.9, 15.4–22.5°C (N=18).
Habitat: No quantitative assessment; limited DBRU collection records
from sandy clay loam (1) or clay (1) in shrubland (1) or woodland (1);
long series from Nylsvlei (Ditsong Museum), all but one collected near
a river, which may indicate alluvial soils rather than sand.
Food types: No quantitative assessment; limited DBRU collection records
from cattle dung (2).
Temporal activity: Diel periodicity not known; active in the summer
rainy season (Oct. to Apr.).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); N Dry Highveld Grassland (Gh), N Mesic Highveld
Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 118 900 km2; AOO probably much
smaller, but poorly known ecologically; AOO may be influenced by
specialised soil association, but quantitative support required; probably
2 mm
no bias in association with open or partially shaded vegetation, but
quantitative support is lacking; although probably not threatened due
to extensive EOO, currently assessed as Data Deficient (DD).
Conservation measures: Quantitative data on ecological associations are
required before an assessment may be made of conservation status;
protected in Roodeplaat and Mogol nature reserves, also on Satara
Floodplain in Kruger National Park.

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 477

Onthophagus giraffa
(Hausmann, 1807)

= Copris pectoralis Thunberg, 1818

= Copris platycera Wiedemann, 1823
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Copris giraffa: ‘Vorgebirges der guten Hoffnung’ [Cape

of Good Hope, South Africa]; C. pectoralis: ‘majorem horum partem
in capite bonae spei collegi’ [amongst material mostly collected in the
Cape of Good Hope, South Africa]; C. platycera: ‘Prom. bon. sp.’ [Cape
of Good Hope].
♂
Taxonomy: Accepted Group 17 species; sexually dimorphic (head), char-
acter varies in prominence with body size.
Distribution: Dry coastal lowlands and coastal mountains from SW
Western Cape to Eastern Cape, South Africa, encompassing the moist-
er SW of the winter rainfall region and the bimodal (spring/autumn)
rainfall region; W/E disjunction would be a collection artefact.
Locality data (mean ± SD, range): Altitude: 480 ± 508, 0–1 993 m;
annual rainfall: 408 ± 78, 246–564 mm; annual temperature (max. +
min. /2): 15.8 ± 1.9, 9.4–18.0°C (N=52).
Habitat: Available quantitative data are difficult to interpret; averaged data
suggest generalist habits; however, local data from natural West Coast
shrubland at Pampoenvlei suggest bias to deep sand (74) compared to
sandy loam (8); on deep sands of the Cape Peninsula, abundance is
lower in dense natural Restio-dominated shrubland (519) than in Ki-
kuyu grass pasture (1 041) created by clearance of natural shrubs; simi-
lar results are shown on deep sands of the West Coast at Geelbek (sparse
grass = 669, open shrubland = 199); DBRU collection records from sand
(8), sandy loam (5), sandy clay loam (4) in disturbed pasture (11), nat- 2 mm
ural shrubland (3), open woodland (1), forest (1), fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (18) and single records from buffalo, elephant, horse,
human/cattle dung mix.
Temporal activity: Diurnal flight activity during most of the year (Aug.
to May); on Cape Peninsula and West Coast: peaks in spring (Aug. to
Nov.) and autumn (Mar.) tailing off into dry mid-summer and cold
mid-winter; peaks in both sclerotised and teneral adults in spring (Aug.
to Nov.) and of teneral adults in late summer (Feb. to Mar.).
Bioregions South Africa: Most elements of Fynbos, Fynbos-Renosterveld
and Renosterveld Bioregions (FO 1–3, 5–8), also West Strandveld (F
11) (Fynbos Biome).
Assessment rationale: EOO = 59 495 km2; widespread along SW and
S coastline of South Africa, apparently with greater abundance in
transformed pasture habitats than natural shrubland; although trans-
formation for arable farming is extensive, particularly in the centre
of its range, it is assessed as Least Concern (LC) since it remains well
represented on various soil types in both natural and disturbed habitat
used for grazing.
Conservation measures: Assessment of conservation status would ben-
efit from more clearly defined data on soil, vegetation and dung type
associations, particularly precise responses to widespread habitat trans-
formation; protected in West Coast and Bontebok national parks, also
Cape of Good Hope Nature Reserve. ♀ 2 mm

ONTHOPHAGINI
478 SURICATA 6 (2020)

Onthophagus giuseppecarpanetoi
Tagliaferri & Moretto, 2012

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Mozambique, [Maputo], Matutuine distr., Lake Piti (east-

ern shore), 26°36’40.9’’S 32°53’19.7’’E’.
Taxonomy: Accepted Group 7 species; sexually dimorphic (head, front
tibiae); characters vary a little with body size.
Distribution: Open vegetation on deep coastal sands of the Maputaland
Centre of Endemism: SE Mozambique and NE coastal KwaZulu-
♂ Natal, South Africa.
Locality data (mean ± SD, range): Altitude: 36 ± 47, 0–222 m; annual
rainfall: 754 ± 78, 634–971 mm; annual temperature (max. + min. /2):
22.3 ± 0.3, 21.6–22.6°C (N=27).
Habitat: No quantitative assessment of soil association; on deep sand
in Maputo Special Reserve: strong bias to grassland on fossil lagoons
(30.0), few in sand forest patches (large, 1.1; medium, 1.9; small, 2.7)
or cooler coastal dune forest (0.03); severely limited DBRU collection
records in grassland (2), shaded thicket (1).
Food types: No quantitative assessment; known from elephant dung.
Temporal activity: Flight activity during darkness in the summer rainy
season (Oct. to Jan.).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB) and
adjoining SE Lowveld (SVl) Savanna Biome.
Assessment rationale: EOO = 8 945 km2; probably a deep sand specialist
primarily in open vegetation, but no formal data to support soil asso-
2 mm ciation; known EOO relatively small, but most records from protected
areas; therefore, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation assessment would be
improved by data to support soil specialisation and determine dung
type associations; protected in Tembe Elephant Park and uMkhuze
Game Reserve (South Africa), Maputo Special Reserve (Mozambique).

2 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 479

Onthophagus gonopygus
d’Orbigny, 1902

= Onthophagus phalopsides Frey, 1955

Global: LC

J A S O N D J F M A M J

Type localities: O. gonopygus: ‘Côte du Congo.....Benguella’ [?Congo

coast; Benguela, Angola]; O. phalopsides: ‘Karibib, S. W. Afrika’ [Karib-
ib, Namibia].
Taxonomy: Accepted Group 30 species; recent synonymy of O. phalopsides
with O. gonopygus based on material from Benguela; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Arid edge of W escarpment in Namibia; presumably also ♂
arid coastal regions of SW Angola near type locality of Benguela; one
record from Tswalu Kalahari Reserve (Northern Cape, South Africa)
requires validation; citations from tropical Democratic Republic of the
Congo (DRC), Gabon also require validation.
Locality data (mean ± SD, range): Altitude: 881 ± 443, 58–1 884 m;
annual rainfall: 185 ± 104, 14–375 mm; annual temperature (max. +
min. /2): 17.0 ± 1.4, 13.3–18.8°C (N=17).
Habitat: No quantitative assessment; DBRU collection records from sand
(7), sandy loam (1) in grassland (2), scrub/shrubland (5), open wood-
land (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (5), horse (2).
Temporal activity: Diel flight periodicity unknown; active in late summer
rainy season (Feb. to Apr.).
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316), Namib
Desert (AT1315).
Assessment rationale: EOO = 102 535 km2; wide range along the little-
transformed, arid, SW coastal region that is used primarily for grazing
of domestic livestock and conservation; AOO would be smaller if asso- 2 mm

ciation with sandy soils is supported by quantitative data; nevertheless,

assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by a survey to ascertain the full EOO and quantitative data on
ecological associations; protected in parts of Namib-Naukluft National
Park (Namibia).

♀
2 mm

ONTHOPHAGINI
480 SURICATA 6 (2020)

Onthophagus granulifer
Harold, 1886

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Oranje-Freistaat’ [Free State, South Africa].

Taxonomy: Accepted Group 24 species; sexually dimorphic (head), char-
acters vary in prominence with body size.
Distribution: Deep sands of S Kalahari: South Africa, Botswana, Namib-
ia, Angola; tentative identification of species recorded at three E outlier
localities; validation required for citations from Zimbabwe, Democrat-
♂ ic Republic of the Congo (DRC) owing to presence of close relatives.
Locality data (mean ± SD, range): Altitude: 1 060 ± 115, 799–1 345 m;
annual rainfall: 299 ± 117, 171–813 mm; annual temperature (max. +
min. /2): 19.7 ± 1.1, 16.8–23.1°C (N=127).
Habitat: No adequate quantitative assessment; in Northern Cape: high
frequency in SW Kalahari (92.8%, 90/97 samples) and high abundance
on deep sand (33.5/sample) or sandy loam (21.5); DBRU collection
records exclusively on deep sand (15) in grassland (5), scrub/shrubland
(5), open woodland (4).
Food types: In Botswana: well represented on various dung types, but with
clear bias: pig (4 052), sheep (2 179), elephant (1 363), cattle (550),
poorly attracted to carrion: chicken livers (34) (cited as Onthophagus
sp. k); DBRU collection records on dung of cattle (10), wildebeest (1),
elephant (2), donkey (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Oct. to May).
Ecoregions Namibia, Botswana: N Namibian Savanna Woodlands
(AT1316), Kalahari Xeric Savanna (AT1309), Angolan Mopane
Woodlands (AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709),
Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea
2 mm Woodlands (AT0726).
Bioregions South Africa: Kalahari Duneveld (SVkd), Eastern Kalahari
Bushveld (SVk) (Savanna Biome); sand patches in N Bushmanland
(NKb), N Upper Karoo (NKu) (Nama Karoo Biome); sand outliers
in Dry Highveld Grassland (Gh) (Grassland Biome), Central Bushveld
(SVcb) (Savanna Biome).
Assessment rationale: EOO = 1 080 205 km2 (possibly underestimated);
AOO would be limited by soil specialisation although deep sand is
widespread across the EOO; SW part of large range is centred on a
little-transformed region used primarily for conservation and grazing
of farm livestock to which it is attracted in abundance; assessed as Least
Concern (LC).
Conservation measures: Asessment of conservation status would be im-
proved by better quantitative data on habitat associations; protected in
Kgalagadi Transfrontier Park (South Africa/Botswana).

2 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 481

Onthophagus graphicus
Wallengren, 1881

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘ad Christianam inventum’ [interpreted as Christiana,

South Africa].
Taxonomy: Accepted Group 15 species; another group member, Ontho­
phagus maculatus (Fabricius, 1801), is now assigned to the genus
Paracatonomus Paulian, 1932; however, pending validation of changed
generic affiliations of O. graphicus, we continue to treat it as an Ontho­
phagus species.
Distribution: Widespread distribution in dry savanna from S to E Afri-
ca; patchiness possibly a collection artefact due to food specialisation:
South Africa, Zimbabwe, Botswana, Namibia, Kenya.
Locality data (mean ± SD, range): Altitude: 935 ± 373, 58–1 201 m;
annual rainfall: 486 ± 157, 279–717 mm; annual temperature (max. +
min. /2): 20.0 ± 1.7, 17.8–22.6°C (N=9).
Habitat: Not known.
Food types: No quantitative assessment; two records from meat bait; re-
corded to dead millipede bait in Kenya (Krell 2004).
Temporal activity: Diel flight periodicity unknown; recorded in the sum-
mer rainy season (Nov. to Feb.).
Ecoregions Namibia, Zimbabwe: N Namibian Savanna Woodlands
(AT1316), Southern African Bushveld (AT0717), Zambezian and Mo-
pane Woodlands (AT0725).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 088 955 km2 (gross estimate); scattered
records may be a collection artefact related to putative carrion special-
isation, but further supporting data are required; probably deserves
Least Concern (LC) status on basis of widespread range, but currently
assessed as Data Deficient (DD) since poorly known taxonomically and 2 mm

ecologically.
Conservation measures: A quantitative survey of ecological associations
is required before an assessment may be made of conservation status;
protected within Stevensford (Botswana) and Ndumo (South Africa)
game reserves.

ONTHOPHAGINI
482 SURICATA 6 (2020)

Onthophagus herus
Péringuey, 1901

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Southern Rhodesia (Salisbury)’ [Harare, Zimbabwe].

Taxonomy: Accepted Group 16 species; sexually dimorphic (head, fore
tibia, prothoracic disc), characters vary in prominence with body size.
Distribution: Dry to moist shaded vegetation on the Mozambique Coast-
al Plain and upland moist savanna in SE Africa; NE tip South Africa,
Zimbabwe, Mozambique; also cited from Malawi, Tanzania, Demo-
♂ cratic Republic of the Congo (DRC).
Locality data (mean ± SD, range): Altitude: 439 ± 489, 76–1 410 m;
annual rainfall: 722 ± 237, 383–1 007 mm; annual temperature (max.
+ min. /2): 23.3 ± 2.6, 18.7–25.9°C (N=8).
Habitat: No quantitative assessment; several Ditsong Museum records
from dense woodland or forest.
Food types: No quantitative assessment; single male type found in carri-
on; single DBRU collection record from baboon dung.
Temporal activity: Diel flight periodicity unknown, but presumably diur-
nal during the summer rainy season (Nov. to Mar.).
Ecoregions Zimbabwe: Dense riverine vegetation at edges of Zambezian
and Mopane Woodlands (AT0725) and Southern Miombo Woodland
(AT0719).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 1 168 730 km2 (gross estimate); AOO
would be very much smaller owing to probable vegetation (shade) and
food type specialisation (omnivore dung, carrion), but this needs quan-
titative support; if correct, clearance of dense woody vegetation would
be detrimental to population density; however, S extremity of range in
NE South Africa entirely protected; currently assessed as Least Concern
2 mm (LC) on basis of wide distribution.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Kruger National Park (South Africa).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 483

Onthophagus hyaena
(Fabricius, 1801)

= Onthophagus impictus Fahraeus, 1857

= Onthophagus scabrosus Péringuey, 1901
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus hyaena: ‘Cap. Bon. Spei.’ [Cape of Good

Hope, South Africa]; O. impictus: ‘in terra Natalensi’ [KwaZulu-
Natal, South Africa]; O. scabrosus: ‘Natal (Newcastle, Frere), Transvaal
(Potchefstroom, Lydenburg)’ [South Africa: KwaZulu-Natal; Potchef-
stroom, North-West Province; Mashishing, Mpumalanga]. ♂
Taxonomy: Accepted Group 9 species, but requires validation; type should
be compared with that of putative closely related, O. cineraceus d’Or-
bigny, 1902 (type locality: Cape Town); synonymy of O. impictus and
O. scabrosus are probable errors as type localities fall outside of ranges
of O. cineraceus and O. hyaena; sexually dimorphic (head, prothoracic
disc), characters vary in prominence with body size.
Distribution: Lowland, S coastal belt, Eastern Cape and highlands along
W Lesotho border primarily in E Free State, South Africa; disjunction
may be a collection artefact, but probably real across unsuitable biore-
gions.
Locality data (mean ± SD, range): Altitude: 1 250 ± 869, 21–2 199 m;
annual rainfall: 614 ± 116, 421–814 mm; annual temperature (max. +
min. /2): 15.2 ± 1.7, 11.2–17.6°C (N=38).
Habitat: No quantitative assessment; DBRU collection records on sand
(3), sandy loam (14), sandy clay loam (11) in pasture/grassland (25),
fallow crop fields (2); also recorded in fynbos shrubland and forest.
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (30), horse (2), human/cattle mix (2).
Temporal activity: Diel flight periodicity unknown, but probably diur-
nal, primarily in the summer rainy season (Oct.–May) or autumn in 1 mm
bimodal rainfall region of S coast (Feb.–May).
Bioregions South Africa: S Mesic Highveld Grassland (Gm), SE tip Dry
Highveld Grassland (Gh), S tip Drakensberg Grassland (Gd) (Grass-
land Biome); Eastern Fynbos-Renosterveld (FO 6) (Fynbos Biome).
Assessment rationale: EOO = 38 760 km2; vegetation associations
unclear; primarily recorded in pasture grassland, but also series from
Tsitsikamma Forest; however, considering fairly large range and fre-
quent occurrence in farmland pastures, including fallow crop fields,
currently assessed as LC.
Conservation measures: Taxonomic issues need to be resolved by com-
parison of types of O. hyaena and O. cineraceus plus possible revision of
the status and relationships of O. impictus and O. scabrosus; assessment
of conservation status of O. hyaena would be improved by quantitative
data on ecological associations; protected in Tsitsikamma forest.

♀ 1 mm

ONTHOPHAGINI
484 SURICATA 6 (2020)

Onthophagus immundus
Boheman, 1858

= Onthophagus vitulus Péringuey, 1901

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: O. immundus: ‘Promontorium Bonae Spei’ [Cape of

Good Hope, South Africa]; O. vitulus: ‘Cape Colony (Cape Town,
Stellenbosch, Worcester, Paarl, Malmesbury)’ [Western Cape, South
Africa].
Taxonomy: Accepted Group 31 species (subgenus: Furconthophagus); sex-
♂ ually dimorphic (head), characters vary somewhat in prominence with
body size.
Distribution: Centred on Renosterveld regions, SW Western Cape to W
Eastern Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 521 ± 367, 62–962 m; an-
nual rainfall: 340 ± 86, 246–539 mm; annual temperature (max. +
min. /2): 16.0 ± 1.4, 13.1–18.2°C (N=13).
Habitat: In Western Cape: not recorded on cooler Cape Peninsula (0),
rare on warmer W coastal sands (1), primarily recorded on sandy loam
in three pastures cleared of natural shrubland in Darling Hills (70, 74,
23); DBRU collection records from sand (2), sandy loam (7), sandy
clay loam (1) in pasture/grassland (7), or crop fields (4).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (11).
Temporal activity: Diel flight periodicity unknown; recorded in all
months of the year except mid-winter (Aug. to May); but, in Western
Cape: primarily recorded in spring (sclerotised: Aug. to Oct., especially
Aug.; tenerals: especially Oct.), rare in autumn (May).
Bioregions South Africa: Centred on West Coast Renosterveld (FO 7),
1 mm
East Coast Renosterveld (FO 8), Western Fynbos-Renosterveld (FO 5)
(Fynbos Biome).
Assessment rationale: EOO = 26 615 km2; restricted to SW South Afri-
ca; occupies a fair-sized EOO although AOO would, presumably, be
smaller according to occurrence of Renosterveld; although endemic to
the winter and W bimodal rainfall regions, all known vegetation records
are from pasture or crop fields cleared of natural shrubland; because of
large EOO and apparent tolerance of habitat disturbance, currently
assessed as Least Concern (LC), although monitoring of population
density would be advisable, particularly in the most highly disturbed
W part of its range.
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations, par-
ticularly formal comparisons of population density in natural versus
replacement pasture vegetation since most Renosterveld has been
transformed; protected in the Eastern Cape in the conservation area
incorporating Swartberg Pass.

♀
1 mm

ONTHOPHAGINI
SURICATA 6 (2020) 485

Onthophagus interstitialis
Fahraeus, 1857

= Onthophagus glaber Boheman, 1857

Global: LC
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type localities: O. interstitialis: ‘in terra Natalensi’ [KwaZulu-Natal,

South Africa]; O. glaber: ‘Caffraria interiore’ [probably interior of
South Africa].
Taxonomy: Accepted Group 7 species probably belonging to the new sub-
genus, Bicornonthophagus Tagliaferri & Moretto, 2012; sexually dimor-
phic (head, prothoracic disc), characters vary somewhat in prominence ♂
with body size.
Distribution: Widespread across drier W Highveld to moist NE Highveld
and NE lowlands, South Africa; record from Zimbabwe requires valida-
tion; currently considered primarily endemic to South Africa.
Locality data (mean ± SD, range): Altitude: 1 042 ± 562, 43–1 914 m;
annual rainfall: 653 ± 119, 435–947 mm; annual temperature (max. +
min. /2): 17.9 ± 3.0, 12.9–23.6°C (N=95).
Habitat: In Gauteng bushveld: extreme bias to sandy clay loam (56)
compared to deep sand (2) and to grassland (47) compared to open
woodland (4) and shaded thickets (7); in lowlands of uMkhuze Game
Reserve: sand (2.3), duplex soil (sand over clay) (55.8), sandy clay loam
(238.8), clay (18.0) with 64% of abundance in grassland; supported by
DBRU collection records primarily from NE lowlands: clay (9), sandy
clay loam (23), sandy loam (18), sand (8) in grassland/pasture (28),
open shrub/woodland (11), forest/thicket (3); as well as fallow crop
fields (4).
Food types: In Ithala Game Reserve: extreme bias to dung of pig (119) 2 mm
compared to horse (1) and cattle (2); however, DBRU collection re-
cords biased to more frequently recorded dung of cattle (54), wilde-
beest (1), elephant (8), rhinoceros (2), zebra (3), horse (1), warthog (1),
human/cattle mix (3).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Oct. to May); in Gauteng bushveld primarily active
Oct. to Feb., tailing off thereafter; attracted to light.
Bioregions South Africa: Moister E Dry Highveld Grassland (Gh), warm-
er N Sub-Escarpment Grassland (Gs) (Grassland Biome); Lowveld
(SVl), Mopane (SVmp) (Savanna Biome); scattered records in warmer
parts of Mesic Highveld Grassland (Gm).
Assessment rationale: EOO = 251 810 km2; AOO presumably smaller
due to specialisation on finer-grained soils; high frequency of records
from cattle dung across farmland despite measured bias to omnivore
dung, which might, however, lower population density; assessed as
Least Concern (LC) on basis of widespread occurrence.
Conservation measures: Assessment of conservation would be improved
by tests of population density in improved pasture versus natural high-
veld grasslands; lowland populations protected in Kruger National
Park, uMkhuze and Hluhluwe–iMfolozi game reserves.

2 mm

ONTHOPHAGINI
486 SURICATA 6 (2020)

Onthophagus juvencus
Klug, 1835

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Isle de Prince’ [Isle of Príncipe, São Tomé e Príncipe, Gulf
of Guinea].
Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus),
but, given the type locality and extremely wide cited distribution, S,
E, and W African representatives should, perhaps, be re-examined for
validity as a single species; minor sexual dimorphism (head).
Distribution: Widespread from S to E to W Africa: South Africa, Mozam-
bique, Botswana, Namibia, Kenya, Príncipe Island, Côte d’Ivoire; also
cited from Ethiopia, Eritrea, Uganda, Rwanda, Democratic Republic
of the Congo (DRC), Gabon, Chad, Niger, Benin, Guinea (Conakry),
Guine (Bissau), Gambia, Senegal; in southern Africa: open vegetation
on deep sands of the E coast and Okavango Delta.
Locality data (mean ± SD, range): Altitude: 67 ± 185, 0–919 m; annual
rainfall: 865 ± 173, 646–1 275 mm; annual temperature (max. + min.
/2): 22.5 ± 0.9, 21.6–24.8°C (N=25).
Habitat: On deep sand in Maputo Special Reserve: almost exclusively
in grassland of fossil lagoons (7.43) compared to sand forest patches
(large, 0; medium, 0; small, 0.02); DBRU collection records from sand
(7), sandy clay loam (1) in grassland (13), shrubland (1), open wood-
land (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (5), human (1), human/buffalo mix (1).
Temporal activity: Flight activity during darkness (18:00–22:00); record-
ed both during the summer rainy season (Oct. to Apr.) and mild dry
season at the coast (May, July, Aug.).
Ecoregions Botswana, Mozambique: Zambezian and Mopane Wood-
lands (AT0725) in Okavango; Maputaland Coastal Forest Mosaic
2 mm
(AT0119), also Southern Zanzibar-Inhambane Coastal Forest Mosaic
(AT0128), Zambezian Coastal Flooded Savanna (AT0906) (not shown
on map).
Bioregions South Africa: Primarily Indian Ocean Coastal Belt Biome
(CB).
Assessment rationale: EOO = 2 971 515 km2 (underestimated, gross es-
timate); cited from localities across a very wide range, but in southern
Africa primarily range restricted to moist grasslands on deep E coastal
sands; also recorded as a rarity in Okavango Swamps and an untraced
locality in Namibia (‘Okurokandovi sandveld’); assessed as Least Con-
cern (LC) owing to wide range and frequent occurrence in open vege-
tation and protected areas in South Africa.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on soil and food type associations in S, E
and W Africa; in southern Africa, protected in iSimangaliso Wetland
Park (World Heritage Site) (South Africa), Maputo Special Reserve
(Mozambique).

ONTHOPHAGINI
SURICATA 6 (2020) 487

Onthophagus kochi
Frey, 1957

No synonyms
Global: DD
Endemic: Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Tulli Distr., Betschuanaland/Südafrika’ (inexact locality:

Tuli Block, NE Botswana).
Taxonomy: Accepted Group 19 species; sexually dimorphic (head), char-
acters vary in prominence with body size; identity of species treated,
here, as O. kochi, requires validation in comparison with O. otjivaron­
gus Balthasar, 1967, described from Abachaus (N Namibia); differs ♂
from close relative in Zambezian and Mopane Woodlands (AT0725) of
the Kruger National Park (South Africa).
Distribution: Dry savanna: N Namibia, N Botswana.
Locality data (mean ± SD, range): Altitude: 949 ± 222, 683–1 246 m;
annual rainfall: 442 ± 105, 356–660 mm; annual temperature (max. +
min. /2): 20.4 ± 1.9, 17.8–22.7°C (N=7).
Habitat: No quantitative assessment; no records.
Food types: No quantitative assessment; reference material recorded from
baits of human dung and meat; a few from banana.
Temporal activity: Diel flight periodicity unknown, but probably in dark-
ness like other Group 19 species; recorded primarily in the summer
rainy season (Oct. to Mar.).
Ecoregions Namibia, Botswana: Namibian Savanna Woodlands
(AT1316), Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian
and Mopane Woodlands (AT0725).
Assessment rationale: EOO = 224 650 km2; recorded from scattered
localities across a wide area of dry savanna, but poorly known ecolog-
ically although Group 19 species are considered to comprise carrion
specialists; currently assessed as Data Deficient (DD), but probably
Least Concern (LC).
2 mm
Conservation measures: Before conservation status may be adequately
assessed, this species requires validation as O. kochi; quantitative data
on ecological associations are also required; not currently known from
any protected region.

♀ 2 mm

ONTHOPHAGINI
488 SURICATA 6 (2020)

Onthophagus lacustris
Harold, 1877b

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Nyassa’ [Lake Malawi region].

Taxonomy: Accepted Group 7 species; aedeagi of NE Tanzanian and
South African specimens identical, but minor differences in punctation.
Distribution: Moister, densely vegetated parts of tropical and subtropical
E seaboard; in the S: Mozambique Coastal Plain and moist edges of
major river valleys (Limpopo, Save, Zambezi); Tanzania, Mozambique,
Malawi, Zimbabwe, South Africa; also reported from Kenya; record
from Democratic Republic of the Congo (DRC) requires validation
(probable error).
Locality data (mean ± SD, range): Altitude: 245 ± 404, 0–1 464 m; an-
nual rainfall: 846 ± 215, 383–1 659 mm; annual temperature (max. +
min. /2): 22.3 ± 1.5, 17.2–25.2°C (N=38).
Habitat: In uMkhuze Game Reserve: recorded only on deep sand (3.3)
and in shaded habitat, not on sandy clay loam or clay (0.0); on deep
sand in Maputo Special Reserve: recorded only in sand forest (large
patches: 16.4, medium: 19.6, small: 7.2) and cooler coastal dune for-
est (6.6), not in grassland (0); supported by limited DBRU collection
records from sand (2) in forest (8) or tree plantation (2); recorded on
finer-grained soil types elsewhere, e.g. Tanzania (East Usambaras).
Food types: No quantitative assessment; limited DBRU collection records
from human/cattle or buffalo dung mix (5); also dung of baboon (1),
cattle (2).
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Oct. to Mar.); attracted to light.
Ecoregions Zimbabwe, Mozambique: Southern Miombo Woodland
(AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: N part of Indian Ocean Coastal Belt Biome
2 mm
(CB); densely vegetated areas in Mopane (SVl) (Savanna Biome).
Assessment rationale: EOO = 874 260 km2 (gross estimate, validated
range); AOO would be smaller owing to shade specialisation in forest
or dense woodland; soil association data may be characteristic only of
coastal NE South Africa; possible bias to omnivore dung, but not well
supported; clearance of dense shady vegetation would be detrimental
but assessed as Least Concern (LC) owing to widespread range and
protection in various reserves.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and vegetation associations at
different points across the EOO; protected in uMkhuze Game Reserve,
Kruger National Park (South Africa), Lake Mutirikwe Game Reserve
(Zimbabwe), Gorongosa National Park (Mozambique).

ONTHOPHAGINI
SURICATA 6 (2020) 489

Onthophagus lamelliger
Gerstaecker, 1871

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Zanzibar’ [Tanzania].

Taxonomy: Accepted Group 31 species (subgenus Furconthophagus); sexu-
ally dimorphic (head, prothoracic disc), characters vary in prominence
with body size.
Distribution: Centred on finer-grained soils in moist, lower-altitude sa-
vanna from southern to southern central Africa; EOO in tropics needs
to be validated by comparison with close relatives, O. exisiciceps d’Or- ♂
bigny (E to W central Africa) and O. ochropygus d’Orbigny (W central
Africa); South Africa, Botswana, Zimbabwe, Mozambique, Malawi,
Tanzania; records from Angola, Zambia, Kenya require validation.
Locality data (mean ± SD, range): Altitude: 591 ± 353, 55–1 573 m;
annual rainfall: 593 ± 170, 304–1 177 mm; annual temperature (max.
+ min. /2): 21.7 ± 1.7, 15.1–25.2°C (N=107).
Habitat: In uMkhuze Game Reserve: conflicting results, deep sand
(186.8), sand on clay (1 180.4), sandy clay loam (354.6), clay (1 155.6)
with a 68% bias to shade; DBRU collection records weakly biased to
finer-grained soils, sand (12), sandy loam (13), sandy clay loam (10),
clay (2), and strongly biased to more shaded vegetation, grassland/pas-
ture (3), shrubland (6), open woodland/woodland (23), forest/thickets
(6); fallow crop field (1).
Food types: At Phalaborwa: strong bias to dung of omnivores (pig,
21 850) compared to monogastric (elephant, 6 335) or ruminant her-
bivores (cattle, 4 125); DBRU collection records from various dung
types, buffalo (1), cattle (24), wildebeest (2), impala (1), donkey (2),
horse (2), zebra (2), elephant (5), rhinoceros (2), warthog (1), human
(2), human/cattle mix (3).
Temporal activity: Flight activity in darkness, primarily during the sum- 1 mm
mer rainy season (Oct. to Apr.); at Phalaborwa: much more abundant
in warm conditions after light rainfall (18 612) than under dry condi-
tions (8 544) or after heavy rainfall (5 146).
Ecoregions Botswana, Zimbabwe: Primarily Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian and Mopane Woodlands (AT0725),
Southern African Bushveld (AT0717).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); marginal in two other bioregions.
Assessment rationale: EOO = 1 440 975 km2; abundant and widespread;
more or less equally abundant in conserved and disturbed areas of a
mining concession at Phalaborwa; perceived bias to finer-grained soils
not well supported by available data; clearance of woodland would,
presumably, influence population density owing to bias to partially
shaded vegetation; assessed as Least Concern (LC) owing to extensive
EOO and great local abundance on various dung types in both reserves
and exploited areas.
Conservation measures: Assessment of conservation status would be im-
proved by further data on soil associations; in South Africa protected
in various reserves, especially, uMkhuze Game Reserve and Kruger
National Park.
2 mm
♀

ONTHOPHAGINI
490 SURICATA 6 (2020)

Onthophagus lamnifer
d’Orbigny, 1902

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal’ [N South Africa].

Taxonomy: Accepted Group 31 species (subgenus Furconthophagus); sexu-
ally dimorphic (head); characters vary greatly in prominence with body
size.
Distribution: Apparently centred on moist upland grassland: South Af-
♂ rica; black squares (tentative localities) within range of close relative,
O. fritschi d’Orbigny, 1902.
Locality data (mean ± SD, range): Altitude: 1 429 ± 344, 21–1 922 m;
annual rainfall: 691 ± 105, 289–804 mm; annual temperature (max. +
min. /2): 15.9 ± 1.3, 12.7–18.6°C (N=38).
Habitat: No adequate quantitative data; on sandy loam in Abe Bailey
Nature Reserve (26oS; 1 496–1 541 m): strong bias to grassland (107.1
per sample) compared to open woodland (7.9); on sandy loam at
more southerly farm Boschberg (28°S): almost exclusively recorded in
mid-altitude grassland (1 264–1 303 m; 31) compared to Highveld
Grassland (1 676–1 687 m; 1); DBRU collection records from sand
(1), sandy loam (8), sandy clay loam (8), clay (2) with all validated
records from grassland/pasture (16) bar for one record from a fallow
crop field.
Food types: No quantitative data; DBRU collection records entirely from
cattle dung (22).
Temporal activity: Diel flight periodicity unknown, but presumably diur-
nal during the summer rainy season (Oct. to May).
Bioregions South Africa: Validated range centred on E Dry Highveld
Grassland (Gh), Mesic Highveld Grassland (Gm), NW Sub-Escarpment
2 mm Grassland (Gs) (Grassland Biome).
Assessment rationale: EOO = 96 245 km2 (validated range, red squares);
limited data suggest a soil generalist with a bias to open grassland that
is readily attracted to the dung of farm livestock although further quan-
titative data is required in support; also one record from a fallow crop
field, which would suggest some tolerance of transformation across its
validated range (25–69%, lower-altitude Gm); therefore, assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; also
data on altitudinal centring and a more exact measurement of EOO as
regards W edge of range, which abuts onto that of very closely related
O. fritschi d’Orbigny, 1902; protected in Abe Bailey Nature Reserve.

2 mm ♀

ONTHOPHAGINI
SURICATA 6 (2020) 491

Onthophagus leroyi
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Sud de l’Afrique orientale anglaise: Escarpment....Mom-

basa....Nord de l’Afrique orientale allemande: Kilima Ndjaro’ [S Kenya:
Escarpment, Mombasa; N Tanzania: Kilimanjaro].
Taxonomy: Accepted Group 23 species, but validation required for occur-
rence in the S although aedeagi of S and E African individuals appear
the same or very similar; sexually dimorphic (head, prothoracic disc),
characters vary in prominence with body size. ♂
Distribution: Wide climatic range in savanna with putative occurrence
from lowlands in the S to lowlands and highlands in the N: NE South
Africa, N Tanzania, S Kenya.
Locality data (mean ± SD, range): Altitude: 382 ± 296, 55–1 205 m;
annual rainfall: 612 ± 152, 305–900 mm; annual temperature (max. +
min. /2): 21.8 ± 1.3, 17.6–24.4°C (N=27).
Habitat: No quantitative assessment; DBRU collection records on sand
(1), sandy loam (3), sandy clay loam (1) with bias to woody vegetation:
shrubland (1), woodland (2), shaded forest/thicket (3).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (1), wildebeest (1), impala (1), elephant (2).
Temporal activity: Diel flight periodicity unknown, but probably in dark-
ness like its close relative, O. fimetarius Roth, 1851; active during the
summer rainy season (Oct. to Mar.).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 228 595 km2 (putative estimate assumes
disjunct populations); assuming S and E African populations are the
same species, widespread in savanna, but ecologically poorly known;
2 mm
possible bias to woody vegetation could indicate potential threats from
clearance in the S; however, often locally abundant with most available
South African records from reserves; assessed as Least Concern (LC).
Conservation measures: Confirmation of single species status is required
for E and S African populations; assessment of conservation status
would be improved by quantitative ecological data, particularly veg-
etation associations and effects of woodland clearance on population
density in the S; in South Africa, protected in Kruger National Park.

♀ 2 mm

ONTHOPHAGINI
492 SURICATA 6 (2020)

Onthophagus leucopygus
Harold, 1867

= Onthophagus opimus Péringuey, 1901

Global: LC (see IUCN Red List – LC as O. impressicollis Boheman,
1860)

J A S O N D J F M A M J

Type localities: O. leucopygus: ‘Inneres Südafrika’ [interior, South Africa];

O. opimus: ‘Cape Colony’ (Griqualand West) [South Africa].
Taxonomy: Accepted Group 24 species with varieties that are entirely
metallic green or with elytra that are pale coloured; sexually dimorphic
(head), characters vary in prominence with body size.
♂ Distribution: Centred on deep Kalahari sands and its outliers: validated
for South Africa, Botswana, Namibia; green variety (red squares), pale
elytra variety (black squares), both varieties (grey squares).
Locality data (mean ± SD, range): Both varieties: Altitude: 1 224 ± 236,
817–1 998 m; annual rainfall: 381 ± 115, 254–660 mm; annual tem-
perature (max. + min. /2): 18.8 ± 1.5, 15.9–22.7°C (N=35).
Habitat: No adequate quantitative assessment; in Northern Cape: record-
ed in very low numbers only on deep sands of the SW Kalahari (13.3%
of samples, 13 of 98); DBRU collection records from sand (7), sandy
loam (3), sandy clay loam (3) in grassland/pasture (5), shrubland (2),
open woodland (4).
Food types: No adequate quantitative assessment; on deep sands in Bot­
swana: recorded only in the arid SW: carrion (1), dung of pig (4), ele-
phant (3), cattle (2), sheep (0); DBRU collection records from dung of
cattle (11), donkey (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Apr.).
2 mm
Ecoregions Namibia, Botswana: Kalahari Xeric Savanna (AT1309), Ka-
lahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), also, sand
outlier in Central Bushveld (SVcb) (Savanna Biome); also in N Dry
Entirely green variety.
Highveld Grassland (Gh) (Grassland Biome).
Assessment rationale: EOO = 587 600 km2; no clear soil or vegetation as-
sociations despite Kalahari centre of distribution; widespread although
recorded in low population density; currently assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data to assess ecological associations; protected
in Witsand Nature Reserve (South Africa).

2 mm
♂
Pale elytra variety.

ONTHOPHAGINI
SURICATA 6 (2020) 493

Onthophagus lugubris
Fahraeus, 1857

= Onthophagus lutulentus Harold, 1867a

Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: O. lugubris: ‘in terra Natalensi’ [KwaZulu-Natal, South

Africa]; O. lutulentus: invalid alternative name, postdates O. lugubris.
Taxonomy: Accepted Group 16 species; sexually dimorphic (head), char-
acters vary in prominence with body size.
Distribution: From cool, moist, upland to high altitude grassland in N
of range to lower-altitude upland and coastal grassland in S of range ♂
within South Africa.
Locality data (mean ± SD, range): Altitude: 1 429 ± 512, 157–2 194 m;
annual rainfall: 719 ± 113, 382–960 mm; annual temperature (max. +
min. /2): 14.9 ± 1.7, 11.0–18.3°C (N=47).
Habitat: No quantitative assessment; DBRU collection records primarily
from finer-grained soils: sand (1), sandy loam (9), sandy clay loam (13)
in pasture/grassland (20); also fallow crop fields (2).
Food types: No quantitative assessment; DBRU collection records pri-
marily from dung of cattle (26), also horse (1), zebra (1), human/cattle
mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
Bioregions South Africa: Mesic Highveld Grassland (Gm), Sub-
Escarpment Grassland (Gs), Drakensberg Grassland (Gd) (Grassland
Biome); also scattered localities (possibly modified to pasture) in Alba-
ny Thicket (AT) and E Fynbos Biomes.
Assessment rationale: EOO = 180 110 km2; recorded in low numbers
at many localities in both reserves and farm pastures; difficult to assess
threats due to absence of quantitative data on habitat associations, but
occurrence across a wide range including fallow crop fields suggests it
may currently be assessed as Least Concern LC). 2 mm

Conservation measures: Assessment of conservation status would be

improved by quantitative data on ecological associations; protected in
Royal Natal National Park, Ithala Game Reserve, Suikerbosrand Na-
ture Reserve.

♀
2 mm

ONTHOPHAGINI
494 SURICATA 6 (2020)

Onthophagus minutus
(Hausmann, 1807)

= Onthophagus opacus Fahraeus, 1857

= Onthophagus rusticus Boheman, 1858
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Copris minuta: ‘Vorgebirges der guten Hoffnung’

[Cape of Good Hope, South Africa]; O. opacus: ‘in terra Natalensi’
[KwaZulu-Natal, South Africa]; O. rusticus: ‘Promontorium Bonae
Spei’ [Cape of Good Hope, South Africa].
♂ Taxonomy: Accepted Group 9 species; sexually dimorphic (head), prom-
inence of characters varies with body size; synonymy of O. opacus
requires validation as the type locality lies far outside of the known
species range of O. minutus.
Distribution: Centred on dry, coastal and inland valley, deep sands of
winter and bimodal rainfall regions, Western and Eastern Cape, South
Africa; W/E disjunction possibly a mix of collection artefact and un-
suitable soil (finer-grained) or climate (higher rainfall).
Locality data (mean ± SD, range): Altitude: 118 ± 146, 0–868 m; annual
rainfall: 303 ± 99, 132–455 mm; annual temperature (max. + min. /2):
16.6 ± 0.9, 15.2–18.4°C (N=39).
Habitat: On West Coast in SW Western Cape: abundance high on coastal
sands in moist natural shrubland (3 548–5 361), substantial in dry nat-
ural shrubland (635) and sparse grassland cleared of shrubs (586–894),
low in moist Kikuyu pasture on sandy loam (32–37); on sand of Cape
Peninsula: high abundance in Kikuyu pasture with high cattle density
(883), low in natural shrubland with low density of herbivores (75–92);
DBRU collection records primarily from deep sand (11), also sandy
loam (1), sandy clay loam (1) primarily from fallow crop fields (7), also
1 mm scrub/shrubland/open woodland (3), grassland cleared of shrubs (2).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (12), buffalo (1), horse (1).
Temporal activity: Diurnal flight activity throughout the year except
June; in Western Cape; activity peak in moist warm spring to early
summer (Aug. to Dec.) becoming very low in dry summer and low
in cool autumn (Jan. to May), absent in June becoming active end of
July; sclerotised individuals throughout year (especially Aug. to Oct.),
tenerals (Sept. to Dec.; mainly Nov.).
Bioregions South Africa: In the W: centred on Western Strandveld (F 11)
in vegetation units (FS 1–3, FS 5, FS 7); also Southwest Fynbos (FO 2),
Sand Fynbos vegetation units (FFd 2–5, FFd 7) (Fynbos Biome); in the
E: recorded in Albany Thicket Biome (AT)
Assessment rationale: EOO = 44 485 km2; wide EOO, but AOO pre-
sumably smaller due to sandy soil bias; abundance variable according
to habitat type and density of dung; apparently somewhat tolerant of
habitat transformation, which is high on coastal sands of Western Cape
(FS: 25–70%; FFd: 27–80%); assessed as Least Concern (LC) owing to
wide range and tolerance of transformation.
Conservation measures: Assessment of conservation status would be
improved by standardised data on ecological associations; protected in
West Coast National Park and Cape of Good Hope Nature Reserve.

1 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 495

Onthophagus monodon
Fahraeus, 1857

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘in terra Natalensi’ [KwaZulu-Natal, South Africa].

Taxonomy: Accepted Group 9 species, but validation required by com-
parison with type(s); original description insufficiently clear to fully
validate the species cited, here, as O. monodon, which is undoubtedly
different to a single specimen in the National Collection of Insects la-
belled ‘Onthophagus monodon’ and ‘Natal’; colour, sheen and rugosity of ♂
interstriae (‘Elytra.....nigro-violacea, nitidissima, rugulosa’) in original
description, better matches the species treated here as O. monodon; sex-
ually dimorphic (head), characters vary in prominence with body size.
Distribution: Cool Highveld Grassland: South Africa; record from Zim-
babwe requires validation (probable error).
Locality data (mean ± SD, range): Altitude: 1 416 ± 165, 1 167–1 672 m;
annual rainfall: 753 ± 174, 514–1 157 mm; annual temperature (max.
+ min. /2): 16.3 ± 1.0, 14.5–17.4°C (N=16).
Habitat: No adequate quantitative assessment; single DBRU collection re-
cord from pasture on sandy clay loam; on sandy clay loam at Carolina:
weak bias to natural Themeda grassland (7) compared to regenerating
crop fields (4) and improved Kikuyu grass pasture (1).
Food types: No quantitative assessment; single DBRU collection record
on dung of cattle; also sampled to cattle dung.
Temporal activity: Diel flight periodicity unknown; active during the
summer rainy season (Nov. to Apr.).
Bioregions South Africa: Dry Highveld Grassland (Gh), Mesic Highveld
Grassland (Gm), margins of Sub-Escarpment Grassland (Gs) (Grass-
land Biome).
Assessment rationale: EOO = 100 835 km2; sampled at scattered localities 1 mm
over a wide area of the Highveld: Mpumalanga, Gauteng, KwaZulu-
Natal, North-West Province, but taxonomically and ecologically poorly
known; population density may be influenced by disturbance of nat-
ural grassland; assessed as Least Concern (LC) on basis of large EOO,
but essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by taxonomic validation and quantitative data on ecological
associations; protected in Abe Bailey Nature Reserve.

♀
1 mm

ONTHOPHAGINI
496 SURICATA 6 (2020)

Onthophagus naso
Fahraeus, 1857

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘juxta fluvium Gariep’ [near Orange River, South Africa –
probably inexact].
Taxonomy: Accepted Group 19 species; minor sexual dimorphism (head).
Distribution: Moist coastal and lower E escarpment grasslands in the
summer rainfall region of SE South Africa.
Locality data (mean ± SD, range): Altitude: 578 ± 517, 0–1 406 m;
annual rainfall: 833 ± 99, 664–988 mm; annual temperature (max. +
min. /2): 18.5 ± 2.1, 13.5–21.5°C (N=18).
Habitat: No adequate quantitative data; in Ithala Game Reserve: single
specimens recorded on finer-grained soils in higher (1 360 m) and
lower middle veld (910 m) grassland; at farm Boschberg, KwaZulu-
Natal: only in middle veld grassland (9 at 1 283 m) not on Highveld (0
at 1 683 m); severely limited DBRU collection records on sandy clay
loam (1), clay (1) in grassland (2).
Food types: No quantitative data; sampled to pig dung at farm Boschberg,
KwaZulu-Natal; many Ditsong Museum specimens trapped to meat
bait.
Temporal activity: Diel activity unknown, but active in summer rainy
season (Oct. to Mar.).
Bioregions South Africa: Mainly Sub-Escarpment Grassland (Gs),
(Grassland Biome); grassland in Sub-Escarpment Savanna (SVs), SE
Lowveld (SVl) (Savanna Biome) and Indian Ocean Coastal Belt Biome
(CB).
Assessment rationale: EOO = 70 340 km2; would have a smaller AOO if
specialisation to omnivore dung /carrion on finer-grained soils in grass-
land is supported; not well represented in most collections; probably a
collection artefact due to food specialisation; however, sizeable range
2 mm that would not be susceptible to woodland clearance if a grassland spe-
cialist; currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
grassland patches in Hluhluwe Game Reserve.

ONTHOPHAGINI
SURICATA 6 (2020) 497

Onthophagus obtusicornis
Fahraeus, 1857

= Onthophagus imitativus Péringuey, 1901

Global: LC (see IUCN Red List – LC)
J A S O N D J F M A M J

Type localities: O. obtusicornis: ‘terra Natalensi’ [KwaZulu-Natal, South

Africa]; O. imitativus: ‘Transvaal (Rustenburg)’ [North-West Province,
South Africa].
Taxonomy: Accepted Group 16 species; sexually dimorphic (head), char-
acters vary in prominence with body size.
Distribution: Known disjunct distribution pattern in three main upland
and coastal savanna centres in southern Africa; possibly relict outlier ♂
populations in Eastern Cape Province (black squares): South Africa,
Botswana, Namibia, Zimbabwe, Mozambique; citation from Zambia
requires validation.
Locality data (mean ± SD, range): Altitude: 1 014 ± 537, 0–1 998 m;
annual rainfall: 634 ± 170, 128–1 100 mm; annual temperature (max.
+ min. /2): 19.0 ± 2.2, 14.5–23.7°C (N=179).
Habitat: Soil associations variable; in Gauteng bushveld: strong bias to
sandy clay loam (3 622) compared to deep sand (263), in uMkhuze
Game Reserve: bias to deep sand (0.9) and sand over clay (3.2) com-
pared to sandy clay loam (0) and clay (0.1); in Gauteng bushveld: bias
to open vegetation, grassland (2 204), open woodland (1 283), shaded
thickets (398); on deep sands in Maputo Elephant Reserve: extreme bias
to grassland (7.8) compared to forest (0.3); DBRU collection records:
sand (18), sandy loam (20), sandy clay loam (10) in grassland/pasture
(26), open shrub/woodland (21), thickets/forest (6), also crop fields (2).
Food types: Somewhat of a food generalist; in Gauteng bushveld: biased
to carrion (33) and pig dung (33), also dung of horse (12), cattle (9),
rotted grass clippings (1), overripe banana (5); similar bias at Phalabor-
wa: dung of pig (119), elephant (14) and cattle (20); DBRU collection
records on dung of cattle (27), elephant (2), rhinoceros (1), zebra (2),
horse (1), donkey (3), human (5), human/cattle mix (4), hyaena (1). 2 mm

Temporal activity: Diurnal flight activity recorded throughout the year,

but in Gauteng bushveld: peak activity in the summer rainy season
continuing into early dry season (Oct. to May); very low activity in
later dry season unless there is unseasonal rainfall (increased activity
recorded after June rain); at Phalaborwa: most active on a warm sunny
day after heavy rainfall (92), less so on a warm cloudy day after light
rain (36) or on a hot dry sunny day (25).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Namibian
Savanna Woodlands (AT1316), Kalahari Xeric Savanna (AT1309),
Southern Miombo Woodlands (AT0719), Maputaland Coastal Forest
Mosaic (AT0119).
Bioregions South Africa: Main range: Lowveld (SVl), Central Bushveld
(SVcb), Eastern Kalahari Bushveld (SVk) (Savanna Biome); N Indi-
an Ocean Coastal Belt Biome (CB); Dry Highveld Grassland (Gh), N
Sub-Escarpment Grassland (Gs) (Grassland Biome).
Assessment rationale: EOO = 418 675 km2; widespread in both farmland
and reserves with a bias to open vegetation and some tolerance of habi-
tat transformation; wide range of food types despite a bias to omnivore
dung; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data to resolve apparent conflict in cur-
rent results; occurs in various reserves including Kruger National Park
and Hluhluwe–iMfolozi Game Reserve. ♀
2 mm

ONTHOPHAGINI
498 SURICATA 6 (2020)

Onthophagus obtutus
Péringuey, 1901

No synonyms
Global: LC
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type localities: O. obtutus: ‘Cape Colony (Burghersdorp), Transvaal

(Potchefstroom)’ [South Africa: Burgersdorp, Eastern Cape; Potchef­
stroom, North-West Province].
Taxonomy: Accepted Group 17 species; sexually dimorphic (head and
prothoracic disc), characters vary in prominence with body size.
Distribution: Cool, dry to moist, highland grassland range from NE to
SW along central Highveld: South Africa; also cited from Lesotho.
Locality data (mean ± SD, range): Altitude: 1 579 ± 226, 688–2 009 m;
annual rainfall: 677 ± 107, 289–924 mm; annual temperature (max. +
min. /2): 15.0 ± 1.2, 12.3–19.4°C (N=33).
Habitat: No quantitative assessment; at farm Boschberg, KwaZulu-Natal:
only on hills of Highveld (22 at 1 677 m) not in nearby middle veld
(0 at 1 283 m), less in burnt (5) than natural grassland (17); DBRU
collection records suggest bias to finer-grained soils, sand (3), sandy
loam (4), sandy clay loam (14), clay (1) primarily in grassland/pasture
(12), shrubland (1), also fallow crop fields (3).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (28).
Temporal activity: Diel flight periodicity unknown, but probably diurnal
during the summer rainy season (Oct. to May).
Bioregions South Africa: Centred on Mesic Highveld Grassland (Gm);
marginal in edge of three other grassland bioregions (all Grassland
Biome).
2 mm Assessment rationale: EOO = 89 365 km2; AOO may be smaller if a
bias to finer-grained soils is supported; range centred on a bioregion
subject to 6–69% transformation in vegetation units that are 23–67%
transformed, mostly above 50%; currently assessed as Least Concern
(LC) owing to widespread range and occurrence at many localities.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
effects of habitat transformation since few known records are from
protected areas; protected in Suikerbosrand Nature Reserve (Gauteng).

ONTHOPHAGINI
SURICATA 6 (2020) 499

Onthophagus pallidipennis
Fahraeus, 1857

Onthophagus suturalis
Péringuey, 1888

Onthophagus politissimus
d’Orbigny, 1908

Global: LC

J A S O N D J F M A M J

Type localities: O. pallidipennis: ‘juxta fluvium Gariep’ [near Orange

River, South Africa]; O. suturalis: ‘Limpopo River, Pretoria and Rusten­
burg, Transvaal’ [South Africa: Limpopo, Gauteng and North-West
provinces]; O. politissimus: ‘Rhodésia: Salisbury’ [Harare, Zimbabwe].
Taxonomy: Accepted Group 11 species, but revision required concerning
status of species of similar appearance to O. pallidipennis that are prob-
ably unrecognised synonyms described from within its range; these spe-
cies are currently regarded as valid, i.e. O. suturalis Péringuey, O. poli­
tissimus d’Orbigny, (also see notes on species account for O. peringueyi
Shipp, 1895c).
Distribution: Dry to moist sandy savanna of central and SE southern
Africa: E central Namibia, Botswana, South Africa, Zimbabwe, S Mo-
zambique; Kenya record undoubtedly an error.
Locality data (mean ± SD, range): Altitude: 947 ± 371, 55–1 958 m;
annual rainfall: 502 ± 192, 194–1 075 mm; annual temperature (max.
+ min. /2): 20.1 ± 1.6, 14.3–23.4°C (N=97).
Habitat: In Gauteng bushveld: extreme bias to deep sand (379) compared
to sandy clay loam (4) and very strong bias to open woodland (302)
compared to grassland (60) or shaded thickets (21); in uMkhuze Game
Reserve: strong bias to deep sand (87.4) compared to sand over clay
(25.5), sandy clay loam (18.0), clay (10.5); largely supported by DBRU
collection records on sand (18), sandy loam (11), sandy clay loam (2) in
grassland/pasture (9), scrub/shrubland (7), open woodland (13).
Food types: At Phalaborwa: equally abundant on dung of omnivore (pig:
173) and ruminant herbivore (cattle: 186), less so on monogastric her-
bivore (elephant: 22); DBRU collection records on dung of cattle (21),
buffalo (1), wildebeest (1), impala (1), donkey (2), horse (2), zebra (1).
Temporal activity: Diurnal flight activity in the summer rainy season 2 mm
(Oct. to Apr.); in Gauteng bushveld: primarily active in early rainy
season (Oct. to Dec.) declining in Jan. becoming uncommon Feb. to
Apr.; at Phalaborwa: primarily active on a warm day after heavy rainfall
(295), less active on a hot, dry day (66), least active on a warm cloudy
day (19).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea Woodlands (AT0726), Southern Miombo
Woodlands (AT0719).
Bioregions South Africa: Lowveld (SVl), Mopane (SVmp), Central Bushveld (SVcb), Eastern Kalahari Bushveld (SVk), moist-
er W Kalahari Duneveld (SVkd) (Savanna Biome).
Assessment rationale: EOO = 1 056 735 km2; widespread EOO, but AOO would be smaller due to soil type bias; also, pop-
ulation density possibly influenced by clearance of woodland, which is extensive in parts of range; however, assessed as Least
Concern (LC) owing to large EOO and protection in various reserves.
Conservation measures: Assessment of conservation status would be improved by resolution of apparent confusion in taxo-
nomic relationships; protected in Kruger National Park, uMkhuze Game Reserve (South Africa).

ONTHOPHAGINI
500 SURICATA 6 (2020)

Onthophagus parumnotatus
Fahraeus, 1857

= Onthophagus parumnotatus Fahraeus, 1857, var. conjunctus d’Orbigny,

1905
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. parumnotatus: ‘prope fluvium Gariep’ [near Orange

River, South Africa]; var. conjunctus: ‘Afrique oriental anglaise: Boura’
[Kenya: Bura].
Taxonomy: Accepted Group 16 species; sexually dimorphic (head), char-
acter varies in prominence with body size.
♂ Distribution: Disjunct pattern; moist SE coast and N uplands of South
Africa, moist upland blocks in SE and S central Africa, moist East
African highlands and wet coast (Zanzibar): South Africa, Eswatini,
Zimbabwe, Malawi, Angola, Kenya; also reported from Mozambique,
Democratic Republic of the Congo (DRC), Rwanda, Tanzania.
Locality data (mean ± SD, range): Altitude: 1 213 ± 420, 46–2 286 m;
annual rainfall: 812 ± 145, 549–1 415 mm; annual temperature (max.
+ min. /2): 17.5 ± 2.2, 10.6–22.0°C (N=103).
Habitat: No quantitative assessment; DBRU collection records biased to
finer-grained soils: sand (5), sandy loam (11), sandy clay loam (11),
clay (4) in open vegetation: pasture/grassland (23), open shrub/wood-
land (2), woodland (2).
Food types: No adequate quantitative assessment; in mid-altitude grass-
land and open woodland in Ithala Game Reserve: primarily sampled
to pig dung (26) compared to cattle (6) and horse dung (1); DBRU
collection records primarily from cattle dung (29), also dung of buffalo
(1), horse (2), zebra (2), elephant (1), human (1), and human/cattle
mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
2 mm rainy season (Sept. to Apr.).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719), East-
ern Zimbabwean Montane Forest Grassland Mosaic (AT1006).
Bioregions South Africa: Primarily moist, warm edges of Mesic Highveld
Grassland (Gm) (Grassland Biome) with Central Bushveld (SVcb) or
Lowveld (SVl) (Savanna Biome); also N Sub-Escarpment Grassland
(Gs) (Grassland Biome) and Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 2 391 165 km2 (misleading); AOO very
much smaller on finer-grained soils of moist, warm temperate to trop-
ical uplands and wet coastal situations, hence, the disjunction between
moist upland ranges in South Africa and Zimbabwe across the hot, dry
Limpopo Valley; type locality presumably inaccurate as it is cited from
a cooler area; not abundant, but not uncommon; primarily in open
vegetation across an extensive EOO; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
various nature reserves including Abe Bailey, Suikerbosrand and Ver-
non Crookes (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 501

Onthophagus pauxillus
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Transvaal: Hamman’s Kraal près de Pretoria...; Bechoua-

na : Vryburg’ [South Africa: Hammanskraal, Vryburg].
Taxonomy: Accepted Group 10 species.
Distribution: Centred on moist, lower-altitude savanna in SE Africa:
South Africa, Zimbabwe; also reported from Botswana, Mozambique.
Locality data (mean ± SD, range): Altitude: 949 ± 452, 8–1 670 m;
annual rainfall: 657 ± 99, 306–897 mm; annual temperature (max. +
min. /2): 19.3 ± 2.2, 14.5–23.7°C (N=67).
Habitat: In Gauteng bushveld: no significant soil bias although many
more on sandy clay loam (2 624) than deep sand (798) with bias to
open woodland (2 061) compared to grassland (1 164) and shaded
thickets (197) (cited as Onthophagus sp. nr pullus (b)); in open wood-
land at Leeuwfontein Nature Reserve: occurrence biased to sand
(145.3) compared to sandy loam (103.0) and stony sandy loam (95.7);
DBRU collection records from various soil types: sand (2), sandy loam
(5), sandy clay loam (1), clay (1) in woody vegetation: shrubland (4),
open woodland (3).
Food types: At Phalaborwa: similar numbers on dung of cattle (215) and
pig (163) but few on coarse-fibred elephant dung (22); supported by
DBRU collection records: cattle dung (5), human/cattle dung mix (4).
Temporal activity: Diurnal flight activity primarily during the summer
rainy season (Oct. to May); in Gauteng bushveld: abundant from Oct.
to Mar. with mid-dry season activity in response to unseasonal rainfall
(June–July 1984); at Phalaborwa: primarily active on a warm sunny
day after heavy rainfall (302) rather than on a warm cloudy day after
light rainfall (64) or a hot, sunny day some time after the previous 1 mm
substantial rainfall (34).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Primarily Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome); margins of six other bioregions.
Assessment rationale: EOO = 316 435 km2 (underestimated); small body
size probably responsible for under-collection and under-estimation of
EOO; often abundant with relatively generalist ecological associations;
therefore, likely resilient to less-extreme forms of habitat transforma-
tion; assessed as Least Concern (LC).
Conservation measures: None recommended; protected in various nature
reserves including, Leeuwfontein, Tswaing, Telperion (South Africa).

ONTHOPHAGINI
502 SURICATA 6 (2020)

Onthophagus peringueyi
Shipp, 1895d

= Onthophagus biplagiatus Péringuey, 1888

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: O. peringueyi nom nov. for O. biplagiatus [pre-occupied

name]: ‘Beaufort West, Cape Colony’ [Western Cape, South Africa].
Taxonomy: Group 11 species; treated here as a valid taxon although not
formally revalidated; currently listed in error as a synonym of Onthoph­
agus suturalis Péringuey, 1888, whose type localities of ‘Limpopo River,
Pretoria and Rustenburg’, lie outside of the range of O. peringueyi;
O. suturalis is, currently, listed as a valid species, but would be an un-
recognised synonym of O. pallidipennis Fahraeus, 1857.
Distribution: Cooler S of the arid SW of Africa: South Africa; citations
from Botswana are an error.
Locality data (mean ± SD, range): Altitude: 1 157 ± 243, 97–1 632 m;
annual rainfall: 247 ± 74, 118–538 mm; annual temperature (max. +
min. /2): 17.0 ± 1.8, 13.4–20.2°C (N=136).
Habitat: No adequate quantitative assessment; in scrub and grassland of
Northern Cape: higher frequency in samples from cooler uplands to
S of Orange River (5.0/sample; 90.1% frequency = 109/121) and in
Upper Karoo (5.6; 95.6% = 65/68) than in hot, arid Bushmanland
(0.1; 28.8% = 19/66) with no clear soil association: sand (5.5/sample),
sandy loam (5.1), sandy clay loam (3.2); DBRU collection records on
sand (2), sandy loam (4), sandy clay loam (5), clay (1) in grassland (1),
scrub (2), shrubland (3).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (9), buffalo (2), sheep (2), horse (1).
Temporal activity: Diurnal flight activity during the late summer rainy
season (Jan. to May).
Bioregions South Africa: Bushmanland (NKb), Upper Karoo (NKu),
Lower Karoo (NKl) (Nama Karoo Biome); Albany Thicket Biome
(AT); Rainshadow Valley Karoo (SKv) (Succulent Karoo Biome); mar-
2 mm ginal in two other bioregions to NE.
Assessment rationale: EOO = 219 865 km2; sampled with high frequen-
cy across an arid, little-transformed range used primarily for grazing
of domestic livestock to whose dung it is readily attracted; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by resolution of taxonomic errors and better quantitative data
on ecological associations; protected in Addo Elephant National Park.

ONTHOPHAGINI
SURICATA 6 (2020) 503

Onthophagus pilosus
Fahraeus, 1857

= Onthophagus costipennis Fahraeus, 1857

= Onthophagus pellax Péringuey, 1901
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: O. pilosus: ‘ad Boschiesmanns Rand’ [uncertain origin

meaning: ?to Bushmans Ridge]; O. costipennis: ‘juxta fluvium Gariep’
[near Orange River, South Africa]; O. pellax: ‘Transvaal (Klerksdorp)’
[Klerksdorp, North-West Province, South Africa].
Taxonomy: Accepted Group 16 species, but synonymy of O. pellax and O. pi­ ♂
losus requires validation; aedeagi identical, but clypeal margin of putative
O. pellax (black squares) more sinuous than that of O. pilosus (red squares);
sexually dimorphic (head), character varies in prominence with body size.
Distribution: Apparently restricted to South Arica; O. pilosus: moist NE
Highveld and E coastal occurrence; O. pellax: bushveld occurrence de-
creasing in population density to the SW.
Locality data (mean ± SD, range): O. pilosus: altitude: 1 353 ± 519,
0–2 653 m; annual rainfall: 757 ± 159, 246–1 271 mm; annual tem-
perature (max. + min. /2): 16.2 ± 2.3, 7.9–21.3°C (N=51). O. pel­
lax: altitude: 1 151 ± 150, 903–1 485 m; annual rainfall: 591 ± 75,
338–664 mm; annual temperature (max. + min. /2): 19.1 ± 1.0, 17.2–
21.3°C (N=24).
Habitat: O. pellax: in Gauteng bushveld, extreme bias to sand (16) com-
pared to sandy clay loam (0) with strong bias to open vegetation, grass-
land (13), open woodland (3), shaded thickets (0); in open woodland at
Leeuwfontein Nature Reserve: sand (40), sandy loam (8), stony sandy
loam (29); O. pilosus: on sandy loam in Abe Bailey Nature Reserve: bias
to open vegetation, grassland (10.6), open woodland (3.1); on sandy
clay loam at Carolina: natural Themeda grassland (20), regenerating veg-
etation on former crop fields (27), improved Kikuyu grass pasture (2). 1 mm
Food types: No quantitative assessment; attracted to cattle and pig dung,
but no standardised comparison.
Temporal activity: Diel flight pattern unknown; active primarily in the
summer rainy season (Oct. to Apr.) with July activity recorded in
warmer coastal regions (Hluhluwe Game Reserve).
Bioregions South Africa: O. pilosus: Centred on Mesic Highveld Grass-
land (Gm), Sub-Escarpment Grassland (Gs) (Grassland Biome) and
Indian Ocean Coastal Belt Biome (CB), marginal occurence in five
other bioregions; O. pellax: Central Bushveld (SVcb), Eastern Kalaha-
ri Bushveld (SVk) (Savanna Biome), also N Dry Highveld Grassland
(Gh) (Grassland Biome).
Assessment rationale: EOO = 156 440 km2 (O. pilosus), EOO =
40 975 km2 (O. pellax); possible differences in distribution pattern and
ecological associations may support two cryptic species, but further
data required, particularly to determine if differences in soil association
exist; large ranges, bias to open vegetation, and some resilience to dis-
turbance suggest an overall assessment of Least Concern (LC) is justi-
fied even without taxonomic revision.
Conservation measures: Accurate assessment of conservation status re-
quires that questions of taxonomic status should be resolved; assess-
ment would be improved by further quantitative data on ecological
associations; O. pellax protected in Tswaing and Leeuwfontein nature
reserves, O. pilosus in Abe Bailey and Lekgalameetse nature reserves. ♀
1 mm

ONTHOPHAGINI
504 SURICATA 6 (2020)

Onthophagus plebejus
Klug, 1855

= Onthophagus stercorarius Fahraeus, 1857

Global: LC

J A S O N D J F M A M J

Type localities: O. plebejus: ‘Sena’, lectotype: ‘Sinna’ [= Sena, central

Mozambique]; O. stercorarius: ‘prope fluvium Gariep’ [inexact, near
Orange River, South Africa].
Taxonomy: Accepted Group 5 species (subgenus Afrostrandius); minor
sexual dimorphism (head, prothoracic disc), characters vary in promi-
nence with body size.
♂ Distribution: Widespread in savanna of S and S central Africa, but now
range restricted to game reserves, presumably due to range contraction
of monogastric herbivores from whose dung most field collections have
been made: NE South Africa, N Namibia, N Botswana, Zimbabwe,
S Angola, Zambia, Mozambique, SE Tanzania; citation from Kenya
requires validation.
Locality data (mean ± SD, range): Altitude: 654 ± 403, 55–1 207 m;
annual rainfall: 619 ± 117, 379–871 mm; annual temperature (max. +
min. /2): 22.6 ± 1.1, 21.1–25.7°C (N=32).
Habitat: No quantitative data; DBRU collection records from sand (8),
sandy loam (7), sandy clay loam (4) in grassland (2), shrubland (1),
open woodland (12).
Food types: In Botswana: strong bias to omnivore dung (pig, 71) as op-
posed to herbivore dung (elephant, 25; cattle, 0; sheep, 13) and carrion
(chicken livers, 2); however, DBRU collection records almost exclusive-
ly from dung of monogastric herbivores: elephant (12), rhinoceros (3),
horse (1); ruminant herbivore: cattle (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.), also Aug. at warmer coastline; attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
2 mm
lands (AT0702), Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Mopane (SVmp), Lowveld (SVl) (Savanna
Biome).
Assessment rationale: EOO = 1 949 910 km2; despite a measured bias
to omnivore dung, all DBRU records from game parks (23) or areas
containing elephants (5); overall centring on game reserves is reflected
by the mapped fragmented range; however, currently assessed as Least
Concern (LC) on the basis of small, scattered AOO protected in vari-
ous game reserves within a very large EOO.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations, particularly if there
is a bias to sandy soils and to open woodland whose clearance might
have a negative impact on population density; further food association
data is also required to determine if conservation of this species is tied
to that of monogastric herbivores, particularly elephants, rather than
to that of omnivores; protected in various national parks and game re-
serves including: Kruger, uMkhuze (South Africa), Hwange (Zimbab­
we), Chobe (Botswana), Etosha (Namibia), Bicuar (Angola).

ONTHOPHAGINI
SURICATA 6 (2020) 505

Onthophagus probus
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: ‘Cape Colony (Hope Town), Mossamedes’ [Hopetown,

Northern Cape, South Africa; Namibe, Angola].
Taxonomy: Accepted Group 20 species.
Distribution: Arid, late summer rainfall region, SW Africa: central N
South Africa, SW Botswana, W Namibia, SW Angola.
Locality data (mean ± SD, range): Altitude: 1 010 ± 195, 47–1 884 m;
annual rainfall: 205 ± 63, 22–356 mm; annual temperature (max. +
min. /2): 18.8 ± 1.3, 13.3–21.4°C (N=218).
Habitat: No adequate quantitative assessment; in the Northern Cape:
abundance was lower in arid Bushmanland (9.9) than the moister SW
Kalahari (34.6) where it was equally abundant on deep sand (38.3) and
sandy loam (36.5); however, limited DBRU collection records were pri-
marily from sand (13) rather than finer-grained soils (sandy clay loam:
1) and entirely in open vegetation: grassland (6), scrub/shrubland (6),
open woodland (2).
Food types: In SW Botswana: well represented on various dung types
with a slight bias to omnivore dung (pig: 544) rather than monogas-
tric herbivore dung (228) or ruminant herbivore pads (cattle: 199) or
pellets (sheep: 339), rare on carrion (chicken livers: 2); limited DBRU
collection records from dung of cattle (7), wildebeest (1), horse (2).
Temporal activity: Diurnal flight activity recorded throughout most of
the year; abundance known to be high during the late summer rainy
season (Feb. to Apr.).
Ecoregions Namibia, Botswana: Namibian Savanna Woodlands
(AT1316), SW Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Bushmanland (NKb), NW Upper Karoo
(NKu) (Nama Karoo Biome); Kalahari Duneveld (SVkd), W Eastern
Kalahari Bushveld (SVk) (Savanna Biome).
Assessment rationale: EOO = 830 525 km2; widespread across a little-
transformed region used primarily for the grazing of farm livestock 1 mm

where it is attracted to a wide range of dung types; unclear if AOO is

influenced by soil type; but wide expanses of various soil types occur
within its EOO; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would benefit
from improved quantitative data on habitat associations; protected in
the Kgalagadi Transfrontier Park (South Africa/Botswana).

ONTHOPHAGINI
506 SURICATA 6 (2020)

Onthophagus producticollis
d’Orbigny, 1908

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Natal: Estcourt’ [Estcourt, KwaZulu-Natal, South Africa].

Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus);
sexually dimorphic (head, prothoracic disc), characters vary in promi-
nence with body size.
Distribution: Primarily finer-grained soils in moist upland savanna re-
gions, South Africa: E–W range along N edge of Highveld, Waterberg
♂ and Soutpansberg; also lower SE escarpment, KwaZulu-Natal, South
Africa, plus E central Zimbabwe; disjunctions may be a mix of collect-
ing artefacts and regional centring of distributions.
Locality data (mean ± SD, range): Altitude: 1 283 ± 300, 273–1 736 m;
annual rainfall: 719 ± 79, 587–900 mm; annual temperature (max. +
min. /2): 17.5 ± 1.3, 16.2–17.3°C (N=18).
Habitat: No adequate quantitative assessment; in Gauteng bushveld: not
present on deep sand (0); recorded only on sandy clay loam (13) in
grassland (2), open woodland (5), shaded thicket (6); limited DBRU
collection records on sand (1), sandy clay loam (2), clay (1) in pasture/
grassland (4).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (3).
Temporal activity: Putative diurnal flight activity although other Group
12 species fly in darkness; active during the summer rainy season (Nov.
to Apr.).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719).
2 mm Bioregions South Africa: Central Bushveld (SVcb at N edge of Highveld,
Waterberg, Soutpansberg), edge of Sub-Escarpment Grassland (Gs)
and Sub-Escarpment Savanna (SVs), also S Lowveld (SVl) (Savanna
Biome).
Assessment rationale: EOO = 85 445 km2; AOO may be more restrict-
ed due to soil, vegetation and climate type specialisations, but further
quantitative support required; uncommonly recorded, therefore, diffi-
cult to determine conservation status; assessed as Data Deficient (DD),
but probably Least Concern (LC) on basis of large range.
Conservation measures: Adequate assessment of conservation status
requires improved quantitative data on ecological associations to deter-
mine how much smaller AOO would be compared to EOO; protected
in Rustenburg Nature Reserve and Hluhluwe–iMfolozi Game Reserve
(South Africa).

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 507

Onthophagus pugionatus
Fahraeus, 1857

= Onthophagus pugionatus Fahraeus, 1857, var. latefulvus d’Orbigny, 1902

= Onthophagus rufaticollis d’Orbigny, 1904
= Onthophagus pugionatus Fahraeus var. rufus d’Orbigny, 1905
= Onthophagus pugionatus Fahraeus, 1857 var. quadraticornis d’Orbigny,
1905
= Onthophagus caucanus Balthasar, 1939b
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: O. pugionatus: ‘prope fluvium Gariep et in terra Natalensi’

[South Africa: near Orange River and KwaZulu-Natal]; var. latefulvus:
♂
‘Afrique orientale allemande: Mhonda,..Zanzibar,..Haut-Zambèze,..
Nyassa,..lac Ngami’ [Tanzania, S central Africa, Botswana]; O. rufa­
ticollis: ‘Congo: Haute Maringa’ [Democratic Republic of the Congo
(DRC)]; var. rufus: ‘Afrique orientale allemande: Tanga’ [Tanzania]; var.
quadraticornis: ‘Kibwezi’ [Kenya]; O. caucanus: ‘Columbien, Cauca’
[Colombia: Cauca; Neotropical type locality undoubtedly an error].
Taxonomy: Accepted Group 24 species; sexually dimorphic (head, protho-
racic disc), characters vary markedly in prominence with body size;
also some colour pattern variation; some taxonomic re-examination,
perhaps, justified.
Distribution: Widespread in moist, shaded savanna thicket and forest
from S to NE Africa; also cool highland grasslands at S edge of range:
South Africa, Zimbabwe, Malawi, Mozambique, Tanzania; synonyms
from Botswana, Democratic Republic of the Congo (DRC), Kenya;
also cited from Gabon, Angola, Namibia, Rwanda, Uganda, Ethiopia,
Somalia; many records N of 15°S require validation.
Locality data (mean ± SD, range): Altitude: 916 ± 541, 0–1 824 m; an-
nual rainfall: 714 ± 199, 213–1 394 mm; annual temperature (max. +
min. /2): 19.3 ± 2.6, 14.0–25.2°C (N=129).
Habitat: In Gauteng bushveld: slight bias to deep sand (27) compared 2 mm
to sandy clay loam (9) with an extreme bias to shaded thicket (34)
rare in open woodland (1) and grassland (1) (sampled to cattle dung);
on sandy clay loam at Roodeplaat Nature Reserve (1 254–1 288 m):
thickets (113), open woodland (35), grassland (19) (sampled to com-
posite pig/cattle dung); on sandy loam at Abe Bailey Nature Reserve
(1 496–1 541 m): open woodland (28), grassland (4); supported by
DBRU collection records from sand (7), sandy loam (5), sandy clay
loam (4), clay (1) in thicket/forest (21), shrub/woodland (4), grassland/
pasture (5).
Food types: No quantitative data; DBRU collection records biased to
omnivore dung: human/cattle or buffalo mix (15), human (4), baboon
(1) compared to herbivore dung: cattle (7), buffalo (1), wildebeest (1),
zebra (1), horse (1), warthog (1).
Temporal activity: Flight activity in darkness (18:00–20:00) primarily
during the summer rainy season (Oct. to May), also Aug. at warmer
coast.
Ecoregions Zimbabwe: Shaded situations in Zambezian and Mopane
Woodlands (AT0725), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Primarily moister SE Central Bushveld (SVcb),
Lowveld (SVl) (Savanna Biome), Indian Ocean Coastal Belt Biome
(CB); also E and N margins of four bioregions in Grassland Biome
plus rare records in dry bioregions along Orange River, Northern Cape. ♀
2 mm

ONTHOPHAGINI
508 SURICATA 6 (2020)

Onthophagus pugionatus (continued)

Assessment rationale: EOO = 4 391 285 km2 (gross estimate based on

limited available data); AOO would be much smaller, largely accord-
ing to vegetation offering shade; with such a bias, population density
would be excessively reduced by clearance of dense woody vegetation
over most of its range; high tranformation across main part of range
in South Africa (0–58% (SVl); 2–49% (SVcb)); however, extremely
widespread so assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations with a
design that takes apparent dung and vegetation bias into consideration;
protected in various national parks, game and nature reserves with
dense vegetation offering shade including Kruger, Hluhluwe–iMfolozi,
Roodeplaat (South Africa), Lake Mutirikwe, Victoria Falls (Zimbabwe).

ONTHOPHAGINI
SURICATA 6 (2020) 509

Onthophagus pullus
Roth, 1851

= Onthophagus brevicornis Fahraeus, 1857

= Onthophagus infuscatus Raffray, 1877
= Onthophagus schimbanus Harold, 1886
Global: LC

J A S O N D J F M A M J

Type localities: O. pullus: ‘Tigré in N. Abyssinien’ [Tigray, N Ethiopia];

O. brevicornis: ‘prope fluvium Limpopo’ [near Limpopo River, south-
ern Africa]; O. schimbanus: nom nov. for O. infuscatus (pre-occupied
name): ‘vom Berge Schimba’ [Shimba Mts, SE Kenya].
Taxonomy: Accepted Group 10 species, but further validation of syn- ♂
onyms described from Ethiopia and South Africa would be useful; sex-
ually dimorphic (head), character varies in prominence with body size.
Distribution: Woody savanna from S to NE to W Africa: South Africa,
Namibia, N Botswana, Zimbabwe, Mozambique, Kenya, Ethiopia;
also cited from Malawi, Democratic Republic of the Congo (DRC),
Tanzania, Somalia, Eritrea; West African citations (Gabon, Gambia,
Senegal) and occurrences (Ghana, Côte d’Ivoire) should be validated.
Locality data (mean ± SD, range): Altitude: 852 ± 478, 0–1 815 m; an-
nual rainfall: 636 ± 224, 306–1 254 mm; annual temperature (max. +
min. /2): 20.1 ± 2.4, 15.4–24.8°C (N=46).
Habitat: In Gauteng bushveld: extreme bias to deep sand (443) compared
to sandy clay loam (2) with strong bias to shade or partial shade: shaded
thickets (318), open woodland (105), grassland (32) (cited as Onthoph­
agus sp. a); supported by DBRU collection records on sand (2) in open
woodland (1), woodland (1), forest (2).
Food types: No quantitative assessment; DBRU collection records from
dung of human (1), human/cattle or buffalo mix (3).
Temporal activity: Diurnal flight activity, primarily in the summer rainy
season (Oct. to May); in Gauteng bushveld: abundant Oct. to Mar., 1 mm
numbers falling off steeply Apr. and May.
Ecoregions Namibia, Zimbabwe: N Namibian Savanna Woodlands
(AT1316), Kalahari Xeric Savanna (AT1309), Zambezian and Mopane
Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); N Indian Ocean Coastal Belt Biome
(CB); N margins Grassland Biome (Mesic Highveld Grassland (Gm),
Dry Highveld Grassland (Gh)).
Assessment rationale: EOO = 2 692 290 km2 (gross estimate of range
restricted to type origins in S, E and NE Africa); widespread, but possi-
bly poorly understood taxonomically; AOO in South Africa, probably
smaller due to soil and vegetation specialisation; clearance of woody
vegetation offering shade on sandy soils probably detrimental to popu-
lation density although does occur in low numbers in open situations;
therefore, assessed as Least Concern (LC).
Conservation measures: Taxonomic revision would be useful to deter-
mine if O. pullus comprises a single widespread species or a species
complex; quantitative data on food type association required to support
apparent bias to omnivore dung; protected in Kruger National Park
(South Africa).

♀ 1 mm

ONTHOPHAGINI
510 SURICATA 6 (2020)

Onthophagus quadrimaculatus
Raffray, 1877

Onthophagus quadrinotatus
d’Orbigny, 1905

No synonyms
Global: DD

J A S O N D J F M A M J

Type localities: O. quadrimaculatus: ‘Zanguebar. Montagnes de Schim-

ba’ [Shimba Mts, SE Kenya]; O. quadrinotatus: ‘Kibwézi’ [Kibwezi,
Kenya].
♂ Taxonomy: Both accepted Group 2 species that need to be validated as
distinct taxa or revised as synonyms; minor colour pattern variation in
coastal individuals considered intraspecific; minor sexual dimorphism
(head), character varies in prominence with body size.
Distribution: Recorded from savanna of South Africa, Botswana, Namib-
ia, Zimbabwe, Mozambique; O. quadrimaculatus: cited from Kenya,
Tanzania, Malawi, Zimbabwe, Mozambique; O. quadrinotatus: cited
only from Kenya.
Locality data (mean ± SD, range): Altitude: 827 ± 449, 0–1 430 m; an-
nual rainfall: 618 ± 195, 306–1 003 mm; annual temperature (max. +
min. /2): 20.3 ± 1.8, 17.4–24.4°C (N=34).
Habitat: In Gauteng bushveld: strong bias to finer-grained soils (deep
sand: 4; sandy clay loam: 14) and shaded vegetation (grassland: 0; open
woodland: 0; shaded thicket: 18); severely limited DBRU collection
records: sandy loam (3) on grass verges (2), open woodland (1).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (1), dog (2), dead tenebrionid beetle (1); long series
from Nylsvlei sampled to meat, faeces and banana bait.
Temporal activity: Diel flight perodicity unknown; active in the summer
rainy season (Oct. to Apr.).
Ecoregions Namibia, Botswana: N Namibia Savanna Woodlands
(AT1316); Angolan Mopane Woodlands (AT0702), Kalahari Acacia-
1 mm Baikiaea Woodlands (AT0709), Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl) (Savanna Biome); N Indian Ocean
Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 875 455 km2 (gross estimate); difficult
to assess conservation status given taxonomic uncertainties and limited
ecological data; may be an omnivore dung/carrion specialist, but sup-
porting quantitative data required; strong association with shade would
suggest possible threats from clearance of woody vegetation; assessed as
Data Deficient (DD) although putative widespread range would sug-
gest an assessment of Least Concern (LC) would be justified.
Conservation measures: Taxonomic study of these two species and re-
lated taxa is required before an adequate assessment may be made of
conservation status in conjunction with improved data on ecological
associations, particularly food; protected in Nylsvlei Nature Reserve
(South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 511

Onthophagus quadrinodosus
Fahraeus, 1857

= Onthophagus truncaticornis Boheman, 1860

= Onthophagus trucidatus Harold, 1870
Global: LC

J A S O N D J F M A M J

Type localities: O. quadrinodosus: ‘in terra Natalensi’ [KwaZulu-Natal,

South Africa]; O. truncaticornis Boheman: ‘juxta lacum N’Gami’ [near
Lake Ngami, Botswana]; O. trucidatus: nom nov. for O. truncaticornis
Boheman [pre-occupied name].
Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus),
sexually dimorphic (head, prothoracic disc), characters vary in promi- ♂
nence with body size.
Distribution: Primarily moist woody savanna on sandy soils in southern
Africa: South Africa, NE Namibia, Botswana, Zimbabwe, Angola;
also cited from Malawi, Democratic Republic of the Congo (DRC);
citations from Somalia, Ethiopia, seem unlikely and require validation.
Locality data (mean ± SD, range): Altitude: 955 ± 450, 0–1 640 m; an-
nual rainfall: 676 ± 168, 194–1 010 mm; annual temperature (max. +
min. /2): 19.5 ± 1.9, 15.4–23.3°C (N=52).
Habitat: In Gauteng bushveld: extreme bias to deep sand (542) compared
to sandy clay loam (4) and to woody vegetation, shaded thicket (295),
open woodland (235) compared to grassland (16); in open woodland at
Leeuwfontein Nature Reserve: sand (30), sandy loam (5), stony sandy
loam (0); DBRU collection records from sand (3), sandy loam (5) in
grassland (5), open shrub/woodland (3).
Food types: In Gauteng bushveld: strong bias to omnivore dung (pig, 86)
rather than herbivore dung (horse, 12; cattle, 14), rare on carrion (2);
DBRU collection records from dung of cattle (3), elephant (2).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to May); in Gauteng bushveld: little activity until Dec.
continuing to Mar., tailing off steeply to early May; in Gauteng
bushveld: only evening flight on cooler early and late summer nights
(18:00–20:00 in Nov., Feb., Mar.), both evening and dawn flight in
warmer mid-summer (18:00–20:00, 04:00–06:00 in Dec.). 2 mm

Ecoregions Namibia, Botswana, Zimbabwe: SE Kalahari Xeric Savanna

(AT1309), Zambezian and Mopane Woodlands (AT0725); Zambe-
zian Baikiaea Woodlands (AT0726); Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 2 016 665 km2 (gross underestimate);
AOO would be smaller owing to soil and vegetation specialisation;
bias to woody vegetation suggests that clearance of woodland would
adversely influence population density; transformation across South
African range: 0–58% (SVl), 2–49% (SVcb); however, assessed as Least
Concern (LC) owing to large EOO and occurrence in both farmland
and reserves.
Conservation measures: None recommended as long as woodland savan-
na is conserved within its range; protected within Kruger National Park
(South Africa).

♀
2 mm

ONTHOPHAGINI
512 SURICATA 6 (2020)

Onthophagus rasipennis
d’Orbigny, 1908

= Onthophagus pallidipennis Fahraeus, 1857, sensu d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: ‘Rhodésia: Bulawayo..., monts Matopos...; Transvaal:

Shilouvane près de Leydsdorp...; Makapan près de Pietersburg...;
Hamman’s Kraal près de Pretoria’ [Zimbabwe: Bulawayo and Matopos;
South Africa: Shiluvane, Makapan and Hammanskraal]; O. pallidipen­
nis sensu d’Orbigny, 1902: No locality stated.
Taxonomy: Accepted Group 11 species; difficult to separate from close
relative, Onthophagus pallidipennis Fahraeus, 1857, with whose body
size range it overlaps.
Distribution: Primarily finer-grained soils in moist, lowland woodland
savanna, SE Africa: South Africa, Eswatini, E Botswana, Zimbabwe,
Mozambique (red squares); W populations, characterised by dark area
in centre of elytra, may represent a variety or a further species: N Na-
mibia (black squares).
Locality data (mean ± SD, range): Altitude: 811 ± 457, 48–1 784 m;
annual rainfall: 655 ± 129, 306–1 145 mm; annual temperature (max.
+ min. /2): 20.0 ± 2.1, 14.6–24.4°C (N=73) (SE populations only).
Habitat: In Gauteng bushveld: bias to finer-grained soils, sandy clay loam
(15), deep sand (6) and partial shade, grassland (2), open woodland
(15), shaded thicket (4); partly supported by DBRU collection records
on sand (6), sandy loam (9), sandly clay loam (1), clay (1) in grassland/
pasture (2), shrubland (3), open woodland (11).
Food types: No quantitative assessment; DBRU collection records on a
wide range of dung types: impala (1), cattle (12), wildebeest (1), don-
2 mm
key (2), horse (2), zebra (1) elephant (2), human/cattle mixture (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Apr.).
Ecoregions Botswana, Zimbabwe: SE Kalahari Xeric Savanna (AT1309),
E Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern African
Bushveld (AT0717), Southern Miombo woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); marginal in two other bioregions.
Assessment rationale: EOO = 230 805 km2 (SE populations only); AOO
may be smaller owing to possible soil type bias; widely distributed in
savannas of SE Africa, but bias to partially shaded habitat suggests pop-
ulation density would be reduced by clearance of woodland; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations; protect-
ed in uMkhuze Game Reserve, Kruger National Park (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 513

Onthophagus scapularis
d’Orbigny, 1902

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Colonie du Cap: Port-Elizabeth’ [Eastern Cape, South

Africa].
Taxonomy: Accepted Group 18 species.
Distribution: Coastal lowlands, SE Eastern Cape, South Africa, at the E
limits of the bimodal rainfall region.
Locality data (mean ± SD, range): Altitude: 233 ± 232, 69–397 m; an- ♂
nual rainfall: 493 ± 54, 455–531 mm; annual temperature (max. +
min. /2): 17.6 ± 0.0, 17.6–17.6°C (N=2).
Habitat: Not known.
Food types: Not known.
Temporal activity: Not known.
Bioregions South Africa: Albany Thicket Biome (AT).
Assessment rationale: EOO = 1 725 km2 (presumably underestimated);
extremely poorly known, but probably carrion-associated like other
close relatives in Group 18; apparent rarity and small known range
possibly a collection artefact related to food specialisation; assessed as
Data Deficient (DD).
Conservation measures: Assessment of conservation status will only be
possible following a quantitative survey centred on SE coastal regions
of the Eastern Cape; this should determine the EOO, AOO and eco-
logical associations; not currently known from any protected area.

2 mm

ONTHOPHAGINI
514 SURICATA 6 (2020)

Onthophagus semiflavus
Boheman, 1860

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘in vicinitate fluvii svakop’ [in the vicinity of the Swakop
River, W central Namibia].
Taxonomy: Accepted Group 11 species.
Distribution: Centred on the N and centre of the arid, late summer rain-
fall region, SW Africa: N central South Africa, SW Botswana, primarily
W Namibia, SW Angola.
Locality data (mean ± SD, range): Altitude: 966 ± 229, 47–1 407 m;
annual rainfall: 210 ± 77, 14–428 mm; annual temperature (max. +
min. /2): 19.0 ± 1.5, 14.2–21.8°C (N=109).
Habitat: No quantitative assessment; in Northern Cape: greater frequency
at study sites in arid SW Kalahari (67.3%, 66 out of 98) than in Bush-
manland (32.8%, 22 out of 67) or Karoo to S of Orange River (8.1%,
10 out of 121); DBRU collection records entirely from sand (15) in
grassland/pasture (5), scrub/shrubland (8), open woodland (2).
Food types: In Botswana: attracted to various dung types with a bias
to omnivore dung: pig (2 527), elephant (1 000), cattle (813), sheep
(1 128), rare on carrion (1); DBRU collection records from dung of
cattle (5), wildebeest (1), horse (1), donkey (3).
Temporal activity: Diurnal flight activity during the late summer rainy
season (Dec. to May) with additional records during winter and spring,
probably from warmer regions.
Ecoregions Namibia, Botswana: Namib Desert (AT1315), Namibian
Savanna Woodlands (AT1316), SW Kalahari Xeric Savanna (AT1309);
marginal in Kaokoveld Desert (AT1310), N Succulent Karoo
(AT1322), Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Bushmanland (NKu) (Nama Karoo); Kalaha-
ri Duneveld (SVkd), W Eastern Kalahari Bushveld (SVk) (Savanna
Biome).
Assessment rationale: EOO = 226 630 km2; AOO may be smaller if
specialisation to sandy soils is shown, but currently no supporting
quantitative data; attracted to various dung types across a large arid
range where land usage is primarily conservation or grazing in little-
2 mm
transformed habitat; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations; protected in
Kgalagadi Transfrontier Park (Botswana and South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 515

Onthophagus suffusus
Klug, 1855

= Onthophagus crucifer Klug, 1855

Global: LC

J A S O N D J F M A M J

Type localities: O. suffusus: ‘Inhambane’ [Mozambique]; O. crucifer:

‘Sena’ [central Mozambique].
Taxonomy: Accepted Group 31 species (subgenus Furconthophagus); sex-
ually dimorphic (head); characters vary in prominence with body size.
Distribution: Widespread in dry to moist southern African savanna, but
now mostly localised to game reserves: South Africa, Mozambique,
Zimbabwe, Botswana, Namibia; also cited from Angola, Malawi. ♂
Locality data (mean ± SD, range): Altitude: 719 ± 403, 55–1 410 m;
annual rainfall: 586 ± 113, 362–835 mm; annual temperature (max. +
min. /2): 21.8 ± 1.4, 18.6–25.7°C (N=31).
Habitat: No adequate quantitative assessment; at South African Wildlife
College in Kruger National Park: no soil bias, finer-grained gabbro-
derived soils (101.4), coarse-grained granite-derived soils (92.1), but
fewer in nearby communal rangeland, gabbro-derived (12.3), granite-
derived (11.4); limited DBRU collection records from sand (4) and
sandy loam (3) only in open woodland (7).
Food types: On deep sand in the moister NE of Botswana: sampled to
various dung types, but with a distinct bias to omnivore dung (pig,
851) compared to monogastric (elephant, 396) or ruminant herbivore
dung (cattle, 153; sheep, 25), or carrion (213); similar bias at Phal-
aborwa: dung of pig (741), elephant (105), cattle (5); limited DBRU
collection records from dung of cattle (3), elephant (3), human/cattle
mix (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Sept. to Mar.); at Phalaborwa: more active on a warm
evening following light rainfall (437), fewer on evenings during hot,
dry weather (210) or during warm weather following heavy rainfall
2 mm
(206).
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709),
Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea
Woodlands (AT0726), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 962 255 km2 (underestimated); despite
a measured bias to omnivore dung, most available records out of 34
were close to (7) or within (24) game reserves, which accounts for the
scattered distribution pattern; thus, AOO would be much smaller than
EOO; soil data inconclusive and bias to woody vegetation unsupported
quantitatively; however, owing to wide range and abundance within a
number of reserves, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data to determine if there is a bias to sandy
soils and woody vegetation; also to determine if the distribution pattern
represents range contraction into game reserves driven by range con-
traction of indigenous monogastric herbivores despite conflicting food
association data; protected in various national parks including: Kruger
(South Africa), Chobe (Botswana), Hwange (Zimbabwe).
♀
2 mm

ONTHOPHAGINI
516 SURICATA 6 (2020)

Onthophagus tricorniger
Boheman, 1860

No synonyms
Global: DD
Endemic: South Africa, Namibia, ?Botswana

J A S O N D J F M A M J

Type localities: ‘prope lacum N’Gami et in regione fluvii Svakop’ [inex-

act: near Lake Ngami, Botswana, and in Swakop River region, central
Namibia].
Taxonomy: Accepted Group 12 species, but needs to be validated by
comparison with close relative or synonym, O. tenuigraniger d’Orbigny,
♂ 1913 (type locality (single specimen): ‘Rhodésia: rivière Mpuzi dans le
Manica’ [inexact: Mpudzi River, Zimbabwe]); distribution of individu-
als treated here as O. tricorniger better match the type localities.
Distribution: Dry SW African savanna of the late summer rainfall region:
South Africa, Namibia, Botswana; Botswana type locality (inexact)
requires validation.
Locality data (mean ± SD, range): Altitude: 1 430 ± 248, 1 140–1 712 m;
annual rainfall: 333 ± 85, 184–409 mm; annual temperature (max. +
min. /2): 19.2 ± 2.3, 16.7–22.2°C (N=6).
Habitat: No quantitative assessment; limited DBRU collection records:
biased to finer-grained soils, sand (1), sandy loam (2), sandy clay loam
(3) in low profile woody vegetation, scrub (2), shrubland (3), open
woodland (1).
Food types: No quantitative assessment; limited DBRU collection records
on dung of cattle (4), wildebeest (1), zebra (1).
Temporal activity: Diel flight periodicity unknown; active during the late
summer rainy season (Jan., Feb.).
Ecoregions Namibia: Namibian Savanna Woodlands (AT1316), NW
Kalahari Xeric Savanna (AT1309), Angolan Mopane Woodlands
(AT0702).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk) (Savanna
2 mm Biome).
Assessment rationale: EOO = 65 590 km2 (underestimated); recorded
from scattered localities across a large area in the late summer rainfall
region of SW Africa; probably not currently threatened, but poorly
known taxonomically and ecologically; therefore, assessed as Data De-
ficient (DD).
Conservation measures: Attention needs to be paid to taxonomic issues
with nomenclature; a survey to determine full EOO and quantitative
data on ecological associations are required before an assessment may
be made of conservation status; protected in Etosha National Park
(Namibia).

ONTHOPHAGINI
SURICATA 6 (2020) 517

Onthophagus trinodosus
Fahraeus, 1857

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘juxta fluvium Limpopo’ [near Limpopo River, southern

Africa].
Taxonomy: Accepted Group 16 species; minor sexual dimorphism (head
prothoracic disc), characters vary in prominence with body size.
Distribution: Bushveld at N and NW edge of South African Highveld;
currently considered endemic to South Africa; citation from Mozam- ♂
bique requires validation.
Locality data (mean ± SD, range): Altitude: 1 020 ± 272, 406–1 311 m;
annual rainfall: 635 ± 88, 491–871 mm; annual temperature (max. +
min. /2): 19.2 ± 1.7, 16.9–23.1°C (N=14).
Habitat: No adequate quantitative assessment; in Gauteng bushveld: re-
corded as a rarity only in grassland (2) and open woodland (2) on sandy
clay loam (Roodeplaat), no records on deep sand (Boekenhoutskloof );
but in Tswaing Nature Reserve: sampled on various soil types in
grassland (8), shrubland (15), open woodland (3), shaded thicket (1);
limited DBRU collection records on sandy loam (3) in grassland (1),
shrubland (1), open woodland (1).
Food types: No adequate quantitative assessment; over 15 months sam-
pling to cattle dung at Roodeplaat: recorded as a rarity (4); on a single
occasion in Tswaing Nature Reserve: sampled in greater numbers to
composite pig/cattle dung baits (27); limited DBRU collection records
on cattle dung (3).
Temporal activity: Diel flight periodicity unknown; active during the
summer rainy season (Oct. to Feb.), one record in June.
Bioregions South Africa: SE Central Bushveld (SVcb); single records in
Lowveld (SVl), Mopane (SVmp) (Savanna Biome). 2 mm

Assessment rationale: EOO = 32 470 km2; poorly known species; size-

able, but fairly restricted range; recorded only in very low numbers
bar for samples from Tswaing Nature Reserve; transformation 17–49%
across SVcb part of range, but available data inadequate for determin-
ing conservation status; assessed as Data Deficient (DD).
Conservation measures: Improved quantitative data on ecological associ-
ations are required before conservation status may be assessed; protect-
ed in Tswaing Nature Reserve.

♀
2 mm

ONTHOPHAGINI
518 SURICATA 6 (2020)

Onthophagus ursinus
d’Orbigny, 1902

No synonyms
Global: LC

J A S O N D J F M A M J

Type localities: ‘Mozambique: Delagoa......Natal’ [Maputo, Mozam-

bique; KwaZulu-Natal, South Africa].
Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus).
Distribution: Moist deep sands along the SE coast from Tongaat, South
Africa, at least to Beira, central Mozambique.
Locality data (mean ± SD, range): Altitude: 39 ± 76, 0–319 m; annual
rainfall: 861 ± 158, 654–1 275 mm; annual temperature (max. + min.
/2): 22.2 ± 1.0, 20.7–24.6°C (N=17).
Habitat: No quantitative assessment of soil association; across a gradient
of rehabilitating vegetation on deep sand at Richards Bay: absent from
highly shaded, dense, early succession woodland (9–12 yr) and natural
dune forest; present in less-shaded, early succession shrubland (6 yr:
1.7) and more open later succession woodland (15–24 yr: 0.1–1.2);
most abundant in least shaded, early succession grassland (0 yr: 7.2)
and open senescent late succession woodland with many fallen trees
(27 yr: 15.8); limited DBRU collection records entirely on sand (3) in
pasture (1), shrubland (1), open woodland (1).
Food types: No quantitative assessment; limited DBRU collection records
entirely on cattle dung (3).
Temporal activity: Diel flight periodicity unknown; active in the summer
rainy season (Oct. to Apr.) with records also from the dry season (July).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 38 715 km2; difficult to assess owing to
limited ecological data; probably a coastal deep sand specialist in moist
open vegetation although further quantitative support is required; be-
cause of apparent vegetation bias and fairly extensive range, currently
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Sileza Game Reserve and Tembe Elephant Park (South Africa).

2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 519

Onthophagus venustulus
Erichson, 1843

See Taxonomy section.

Global: LC

J A S O N D J F M A M J

Type locality: ‘Angola’ [also cited from Senegal and Egypt by Erichson].
Taxonomy: Status of O. venustulus dependent on whether or not the ?dis-
junct Kalahari populations of southern Africa are regarded as a valid
Group 31 species (subgenus Furconthophagus); or a synonym of O. var­
iegatus (Fabricius, 1798) (= current valid status), a species with a Saharo–
Sindian distribution pattern from Senegal to Pakistan/Afghanistan that
is also cited from coastal East Africa; sexually dimorphic (head), char-
♂
acters vary in prominence with body size; this variation may be why the
type description of males cites two horns on the vertex instead of the
three borne by major males; identity of this species requires validation.
Distribution: Southern African populations: dry savanna on S Kalahari
deep sands extending to the middle Limpopo Valley and as a rarity in
SE sand outliers: Angola, Namibia, Botswana, Zimbabwe, South Afri-
ca; citation from Lesotho probably an error.
Locality data (mean ± SD, range): Altitude: 1 084 ± 228, 328–1 815 m;
annual rainfall: 329 ± 119, 34–690 mm; annual temperature (max. +
min. /2): 19.6 ± 1.5, 15.9–23.7°C (N=184).
Habitat: No adequate quantitative assessment; in Northern Cape: fre-
quency and abundance high in SW Kalahari (89.8% of samples, 88
out of 98; 85.1/sample on sand, 80.4 on sandy loam), low in Bush-
manland (8.8%, 5/67; 0.04/sample) and uplands to S of Orange River
(7.4%, 9/121; 0.3/sample); DBRU collection records indicate bias to
coarse-grained soils: sand (19), sandy loam (5), sandy clay loam (2) in
grassland (5), scrub/shrubland (9), open woodland (11).
Food types: In Botswana: bias to dung of omnivores (pig, 434) and mo-
nogastric herbivores (elephant, 324) compared to ruminant herbivores 1 mm
(cattle, 84; sheep, 48) and carrion (chicken livers, 72); however, DBRU
collection records entirely from cattle dung (28).
Temporal activity: Flight activity during darkness, primarily in the sum-
mer rainy season (Sept. to Apr.).
Ecoregions Namibia, Botswana: Namibian Savanna Woodlands (AT1316),
Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands
(AT0709); marginal in Zambezian Baikiaea Woodlands (AT0726), An-
golan Mopane Woodlands (AT0702) Zambezian and Mopane Wood-
lands (AT0725) and one other ecoregion.
Bioregions South Africa: Primarily, Kalahari Duneveld (SVkd), Eastern Ka-
lahari Bushveld (SVk), sands of N Mopane (SVmp) in Limpopo Valley, rare
in sand outliers of Central Bushveld (SVcb) (Savanna Biome), also N ex-
tremes Bushmanland (NKb), Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 1 414 010 km2 (underestimated, Kalahari
populations only); AOO possibly smaller according to distribution of
sandy soils, but requires quantitative support; large part of range coin-
cides with little-transformed, arid areas used primarily for conservation
and grazing of farm livestock, assessed as Least Concern (LC).
Conservation measures: Taxonomic issues need to be resolved; assess-
ment of conservation status would be improved by formal quantitative
data on habitat associations; protected in Kgalagadi Transfrontier Park
(South Africa/Botswana), Central Kalahari Game Reserve, Chobe Na-
tional Park (Botswana). ♀ 1 mm

ONTHOPHAGINI
520 SURICATA 6 (2020)

Onthophagus verticalis
Fahraeus, 1857

= Onthophagus haroldi Péringuey, 1888

= Onthophagus hector Shipp, 1895c
= Onthophagus corniculatus Reiche, 1847 sensu Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: O. verticalis: ‘juxta fluvium Gariep’ [near Orange River,

South Africa]; O. hector: nom nov. for O. haroldi Péringuey (pre-occupied
name): ‘Potchefstroom, Transvaal, and Zambesi River’ [North-West
Province, South Africa; inexact, Zimbabwe]; O. corniculatus sensu
♂ Péringuey: recorded from ‘Natal (Estcourt), Transvaal (Potchefstroom,
Rustenburg), Southern Rhodesia (Victoria Falls, Middle Limpopo),
Ovampoland (Omrromba)’ [South Africa, Zimbabwe, Namibia].
Taxonomy: Accepted Group 12 species (subgenus: Trichonthophagus);
sexually dimorphic (head, prothoracic disc), characters vary in promi-
nence with body size.
Distribution: Widespread on sandy soils in dry to moist savanna of
southern Africa: South Africa, Namibia, Botswana, Zimbabwe, Mo-
zambique.
Locality data (mean ± SD, range): Altitude: 1 045 ± 415, 58–1 958 m;
annual rainfall: 520 ± 178, 160–1 220 mm; annual temperature (max.
+ min. /2): 20.2 ± 1.8, 14.3–23.4°C (N=65).
Habitat: In Gauteng bushveld: extreme bias to deep sand (60) compared to
sandy clay loam (0) with a bias to woody vegetation: grassland (6), open
woodland (22), shaded thicket (32); similar soil bias in uMkhuze Game
Reserve: deep sand (1.4), sand on clay (0.7), sandy clay loam (0.3), clay
(0); supported by DBRU collection records on sand (15), sandy loam
(9), clay (1) in grassland (7), scrub/shrubland (6), open woodland (11).
Food types: In Botswana: attracted to various dung types with a bias to mono-
gastric herbivore (elephant, 87) and omnivore dung (pig, 64) compared
2 mm
to ruminant herbivore dung (pellet-dropping sheep, 37; pad-dropping
cattle, 11), not recorded on carrion (0); DBRU collection records on
dung of cattle (15), buffalo (2), elephant (5), horse (1), donkey (2).
Temporal activity: Flight activity during darkness in the summer rainy
season (Oct. to Apr.); in Gauteng bushveld: mid-summer flight activity
peaks close to dawn (04:00–06:00, Nov. to Jan.), lower abundance for
evening flight activity (18:00–20:00).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Xeric Savanna
(AT1309), Kalahari Acacia-Baikiaea Woodlands (AT0709), Angolan
Mopane Woodlands (AT0702), Zambezian and Mopane Woodlands
(AT0725), Zambezian Baikiaea Woodlands (AT0726), Southern
Miombo Woodlands (AT0719).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 288 690 km2; AOO undoubtedly
somewhat smaller due to bias to sandy soils; measured bias to woody
vegetation suggests clearance of woodland would be detrimental to
population density in transformed areas in the NE of its range; trans-
formation: 0–58% (SVl), 2–49% (SVcb); however, assessed as Least
Concern (LC) on basis of wide EOO and protection in reserves.
Conservation measures: None recommended; protected in several na-
tional parks and game reserves including Kruger, Ndumo (South Afri-
2 mm
♀ ca), Chobe (Botswana).

ONTHOPHAGINI
SURICATA 6 (2020) 521

Onthophagus vigens
Péringuey, 1901

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal (Lydenburg)’ [Mashishing, Mpumalanga, South

Africa].
Taxonomy: Accepted Group 17 species, but should be compared to
Onthophagus simillimus d’Orbigny, 1913, described from Chirinda
Forest, Zimbabwe; sexually dimorphic (head), character varies in
prominence with body size. ♂
Distribution: South Africa: from cool, moist, high altitude, forest and
grassland in N of range (E edge of Highveld) to lower-altitude upland
grassland and coastal forest in S of range (central SE Eastern Cape).
Locality data (mean ± SD, range): Altitude: 1 367 ± 462, 112–2 274 m;
annual rainfall: 834 ± 168, 482–1 271 mm; annual temperature (max.
+ min. /2): 15.4 ± 2.3, 10.6–22.8°C (N=44).
Habitat: No adequate quantitative assessment; along a 500–2 800 m
altitudinal gradsect from the coast to the Sani Pass, KwaZulu-Natal:
recorded in grassland only at 1 500 m (0.8) and 1 900 m (1.0); DBRU
collection records entirely on finer-grained soils: sandy loam (3), sandy
clay loam (8), clay (1) in pasture/grassland (10) or forest (2).
Food types: No quantitative assessment; DBRU collection records pri-
marily from cattle dung (8), also human/cattle dung mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
Bioregions South Africa: Patchy records centred on E Mesic Highveld
Grassland (Gm), Sub-Escarpment Grassland (Gs) and Drakensberg
Grassland (Gd) (Grassland Biome) with records from encompassed
forest types, including Northern Afrotemperate (FOz 2), Southern 2 mm
(FOz 3) and Northern Mistbelt (FOz 4) and Scarp Forest (FOz 5); also
Southern Coastal Forest (FOz 6).
Assessment rationale: EOO = 61 695 km2; AOO probably smaller
according to occurrence of suitable cool moist grassland and forest
patches, but survey is required to improve the locality data; all known
collection records from finer-grained soils; because of protection of
some forest and grassland areas across its wide range, currently assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
relative bias to soil type or bias to forest or grassland, as these remain
unknown; protected in the uKhahlamba Drakensberg Park (World
Heritage Site).

2 mm
♀

ONTHOPHAGINI
522 SURICATA 6 (2020)

Onthophagus vinctus
Erichson, 1843

= Onthophagus obesus Fahraeus, 1857

Global: LC
J A S O N D J F M A M J

Type localities: O. vinctus: ‘Angola’; O. obesus: ‘terra Natalensi’ [KwaZulu-

Natal, South Africa].
Taxonomy: Accepted Group 7 species; O. vinctus complex recently subdi-
vided into three species, thus reducing known distribution from 34 to 25
African countries; limited sexual dimorphism (head, prothoracic disc).
Distribution: Widespread, mainly on sandy soils, in dry, lowland, wood-
♂ ed savanna and moist coastal regions from southern to E to W Africa:
recorded in 25 countries from Guinea (Bissau) to Somalia to Angola,
Namibia, Botswana, Zimbabwe, Mozambique and South Africa.
Locality data (mean ± SD, range): Altitude: 797 ± 487, 0–2 274 m; an-
nual rainfall: 663 ± 234, 125–1 659 mm; annual temperature (max. +
min. /2): 20.2 ± 2.2, 10.6–25.7°C (N=305).
Habitat: In Gauteng bushveld: extreme bias to deep sand (9 090) compared
to sandy clay loam (20) and a bias to shaded thickets (6 253) compared to
open woodland (2 229) and grassland (628); in uMkhuze Game Reserve:
bias to deep sand (7.6) compared to duplex sand over clay (1.5), sandy clay
loam (0) and clay (1.9); on deep sands in Botswana: primarily in the moister
wooded NE (3 608) not the arid SW (3); partial support from DBRU col-
lection records: sand (36), sandy loam (14), sandy clay loam (14) in pasture/
grassland (22), shrub/woodland (29) or thicket/forest/plantation (10).
Food types: In Gauteng bushveld: primarily attracted to pig dung (1 245),
much less abundant on horse (69) and cattle dung (58) or carrion (10); in
Botswana: mainly dung of pig (2 269), also elephant (671), sheep (595),
less on cattle (57) and carrion (16); DBRU collection records reflect com-
monly sampled dung types: cattle (44) and a wide range of others, human
2 mm (2), baboon (1), human/cattle mix (7), elephant (6), rhinoceros (2), hip-
popotamus (1), zebra (2), horse (1) donkey (2), buffalo (5), wildebeest
(1), waterbuck (1).
Temporal activity: In Gauteng bushveld: flight activity primarily at dusk
(87–99%) from 18:00–20:00 (Nov.–Mar.); recorded throughout the
year, but primarily active from end of the dry season till end of the
summer rainy season (Sept. to Apr.); attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Namibian Savanna Wood-
lands (AT1316), NW Kalahari Xeric Savanna (AT1309), Angolan
Mopane Woodlands (AT0702), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian and Mopane Woodlands (AT0725), Southern
African Bushveld (AT0717), Southern Miombo Woodlands (AT0719),
Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Primarily Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp), but also Eastern Kalahari Bushveld (SVk)
(Savanna Biome); Indian Ocean Coastal Belt Biome (CB); marginal
in Sub-Escarpment Grassland (Gs), N Dry Highveld Grassland (Gh)
(Grassland Biome).
Assessment rationale: EOO = 8 771 775 km2; AOO influenced by soil
and vegetation bias; clearance of woodland would be detrimental to
population density, but very wide EOO; in a study at Phalaborwa: re-
corded primarily in disturbed mine habitats; therefore, assessed as Least
Concern (LC).
Conservation measures: None required; widespread, abundant and tol-
2 mm
erant of disturbance; recorded in both farmland and protected areas
♀ including Kruger National Park (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 523

Onthophagus virescens
Harold, 1867c

= Onthophagus nitidulus Klug, 1855

Global: LC

J A S O N D J F M A M J

Type locality: O. virescens, nom nov. for O. nitidulus (pre-occupied name):

‘Sena’ [central Mozambique].
Taxonomy: Accepted Group 11 species; name requires validation, but
species characteristics match the redescription in d’Orbigny (1913);
sexually dimorphic (head, disc of prothorax), characters vary in prom-
inence with body size.
Distribution: Lowland savanna on sandy soils, particularly deep sand ♂
patches, on the S Mozambique Coastal Plain and up the Zambezi
Valley: NE South Africa, Mozambique, N Botswana; also cited from
Zimbabwe; north Botswana populations need to be validated by fur-
ther comparison with an unidentified species recorded in southwest
Zimbabwe and north Namibia.
Locality data (mean ± SD, range): Altitude: 572 ± 345, 59–1 000 m;
annual rainfall: 651 ± 160, 487–1 136 mm; annual temperature (max.
+ min. /2): 22.6 ± 1.3, 21.5–25.7°C (N=14).
Habitat: No quantitative assessment; nine out of 16 records on deep sand
in savanna; limited abundance recorded on sandy loam.
Food types: On deep sand in Botswana: strong bias to omnivore dung
(470) compared to herbivore dung (elephant: 106; cattle: 14; sheep:
145), rare on carrion (chicken livers: 2); limited DBRU collection re-
cords from dung of elephant (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Apr.).
Ecoregions Botswana: Zambezian and Mopane Woodlands (AT0725),
Zambezian Baikiaea Woodlands (AT0726)
Bioregions South Africa: Lowveld (SVl), Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 137 725 km2 (gross estimate); large EOO,
but AOO would probably be much smaller according to distribution
of sandy soils in the SE, although quantitative support is required; only
patchily recorded, but most localities lie in protected regions where it
may be abundant (e.g. Chobe); therefore, assessed as Least Concern
(LC). 2 mm
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations; protected in various
national parks: Kruger (South Africa), Chobe (Botswana), Gorongosa
(Mozambique).

ONTHOPHAGINI
524 SURICATA 6 (2020)

Onthophagus vylderi
d’Orbigny, 1913

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Sud-Ouest africain allemand: Damara’ [inexact: Dama­

raland, Namibia].
Taxonomy: Accepted Group 20 species; material cited, here, matches
original description.
Distribution: Dry to moist savanna on deep Kalahari sands: South Africa,
Botswana; also Namibia, but no recent records.
Locality data (mean ± SD, range): Altitude: 1 050 ± 173, 871–1 376 m;
annual rainfall: 588 ± 101, 352–659 mm; annual temperature (max. +
min. /2): 20.8 ± 1.5, 18.6–22.4°C (N=8).
Habitat: No quantitative assessment; only known from deep sands in
open woodland.
Food types: On deep sands in Botswana: sampled only to carrion (chicken
livers: 36), not recorded to dung (0: pig, elephant, cattle, sheep) (cited
as Onthophagus sp. 3); long series from deep sands at Nylsvlei sampled
to faeces, meat or banana; limited DBRU collection records from goat
carrion and dead millipede.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Feb.).
Ecoregions Botswana: Deep sands in Kalahari Xeric Savanna (AT1309),
Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea
Woodlands (AT0726).
Bioregions South Africa: Deep sand patches in Central Bushveld (SVcb)
(Savanna Biome).
Assessment rationale: EOO = 241 110 km2 (based on available Botswana
2 mm
and South Africa localities); probably a deep sand, carrion specialist, but
quantitative data on habitat associations are lacking; food specialisation
may be the reason for patchy records; however, widespread range across
an area where deep sands are extensive with several localities coinciding
with protected areas; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative ecological data, including further food asso-
ciation records; protected in Chobe National Park (Botswana), Nylsvlei
Nature Reserve (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 525

Genus Phalops Erichson, 1847b

Type species and designation: Copris ciconia Fabricius, 1801, by subsequent designation (Shipp 1895c).
Synonyms: = Coprioides Gistel, 1857: Type species: Onthophagus fimbriatus Klug, 1835, by mono-
typy.
= Ephillopus Reitter, 1892: Type species: Scarabaeus iphis Olivier, 1789, by original
designation.
Last review: Entire genus reviewed by Barbero et al. (2003); one species transferred from Ontho­
phagus Latreille, 1802 (Génier 2013).

The long-established genus, Phalops Erichson, 1847b, currently comprises 38 species with distributions in the savannas,
upland grasslands and arid areas of the Afrotropical (33), W Oriental (4) and extreme S Palaearctic regions (Arabia: 1).
The genus has been divided into six species groups (Barbero et al. 2003).

In South Africa, Botswana and Namibia, Phalops is represented by 13 species in five different species groups.
(1) iphis group: One dry savanna species with a range extending into the E tropics.
(2) boschas group: Two species with W savanna or E savanna ranges, the latter with a range extending into the E
tropics.
(3) prasinus group: One species with a dry W savanna range.
(4) divisus group: Three species; one with a dry E savanna grassland range extending into the E tropics; the other two
closely related and restricted to southern Africa; one with a dry W savanna and arid SW Karoo range; one with an
arid W savanna range.
(5) fimbriatus group: Five species; one with an E savanna range extending into the E tropics; the others all closely
related and restricted to southern Africa; one with a savanna, dry W highland grassland and Upper Karoo range;
one with an Upper Karoo and dry W savanna range; one with a dry W savanna range; one centred on deep Kalahari
sands and outliers.

One recently transferred species, centred on Highveld Grassland and E Upper Karoo of South Africa, has not been as-
signed to a group.

As most species show widespread distributions with high frequency of occurrence in open vegetation, all but one have been
assessed as Least Concern (LC). However, some savanna species show a bias to open woodland, mostly those species with
ranges extending into the E tropics. Thus, clearance of trees and shrubs would reduce their population density.

ONTHOPHAGINI
526 SURICATA 6 (2020)

Phalops ardea
(Klug, 1855)

= Onthophagus (Phalops) ardea var. chloritus d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus ardea: ‘Tette und Sena’, lectotype:

‘Senna...Sinna Tette’ [central Mozambique: Sena and Tete]; var. chlo­
ritus: ‘Afrique orientale allemande: Bagamoyo’ [Bagamoyo, Tanzania]
Taxonomy: Accepted species of the iphis group; sexually dimorphic (head,
prothoracic disc), characters vary in prominence with body size; irides-
cent colour variation from cupreous to green to blue.
♂ Distribution: Centred on hot, dry lowland savanna in southern Africa
with a recorded distribution extending northwards from dry S central
African savanna along the hot, moist E seaboard: NE South Africa,
E Botswana, Zimbabwe, N Namibia, SW Angola, Zambia, Malawi,
Mozambique, E Tanzania; reports from Kenya and Uganda require
validation; black square: surprising locality.
Locality data (mean ± SD, range): Altitude: 662 ± 374, 0–1 751 m; an-
nual rainfall: 618 ± 240, 304–1 748 mm; annual temperature (max. +
min. /2): 21.5 ± 1.8, 16.5–25.9°C (N=97).
Habitat: No adequate quantitative assessment; at Leeuwfontein Nature
Reserve: bias to finer-grained soils without stones: sand (7), sandy loam
(26), stony sandy loam (10); supported by DBRU collection records:
sand (6), sandy loam (22), sandy clay loam (6) with bias to open wood-
land vegetation: grassland/pasture (7), shrubland (1), open woodland
(24), forest (1).
Food types: At Phalaborwa: greatest attraction to omnivore dung (pig:
234), frequency less on ruminant and monogastric herbivore dung
(cattle: 41; elephant: 17); DBRU collection records reflect commonly
sampled dung types: cattle (26), elephant (9), donkey (4), horse (3),
also on single rhinoceros, zebra, hyaena and impala droppings.
2 mm
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Mar.); at Phalaborwa: frequency much greater in
hot, dry conditions (243) than in warm cloudy conditions following
light rainfall (13), or, in warm sunny conditions soon after heavy rain-
fall (35).
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Southern African Bushveld (AT0717), Zambezian
and Mopane Woodlands (AT0725).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), N
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 179 470 km2; AOO may be influenced
by soil and vegetation specialisation; across South African range, hab-
itat transformation varies from 0 to 40% (SVmp), 0–58% (SVcb),
2–49% (SVl); nevertheless, widespread in hot, dry, lowland savanna
both within and outside of reserves where it is commonly found in the
dung of domestic livestock despite a bias to omnivore dung; assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on habitat associations, particu-
larly, effects of woodland clearance; protected in Kruger National Park
(South Africa).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 527

Phalops boschas
(Klug, 1855)

= Onthophagus (Phalops) boschas var. chloronotus d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus boschas: ‘Sena’, lectotype: ‘Sinna’ [Sena,

central Mozambique]; var. chloronotus: ‘Nyassa’ [Mozambique].
Taxonomy: Accepted species of the boschas group; little colour variation;
mostly dark, sometimes with green sheen; limited sexual dimorphism
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Centred on hot, fairly dry, southern African savanna with
a recorded distribution extending northwards from S central African ♂
savanna along the hot, moist E seaboard: NE South Africa, NE Bot­
swana, Zimbabwe, NE Namibia, SE Angola, Zambia, Malawi, Mo-
zambique, S Democratic Republic of the Congo (DRC), E Tanzania;
report from Kenya requires validation.
Locality data (mean ± SD, range): Altitude: 724 ± 381, 0–1 503 m; an-
nual rainfall: 630 ± 136, 306–1 151 mm; annual temperature (max. +
min. /2): 21.1 ± 1.8, 17.1–25.7°C (N=130).
Habitat: No adequate quantitative assessment; at Leeuwfontein Nature
Reserve: biased to finer-grained soils without stones: sand (8), sandy
loam (13), stony sandy loam (7); supported by DBRU collection re-
cords: sand (12), sandy loam (27), sandy clay loam (8), clay (1) with
bias to woodland vegetation: grassland/pasture (10), shrubland (2),
open woodland (30).
Food types: At Phalaborwa: greatest attraction to omnivore dung (pig:
173), frequency lower on ruminant and monogastric herbivore dung
(cattle: 19; elephant: 9); DBRU collection records reflect commonly
sampled dung types: cattle (41), elephant (8), rhinoceros (4), also on
single baboon, donkey, warthog, wildebeest and waterbuck droppings.
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.); at Phalaborwa: frequency much greater in
2 mm
hot, dry conditions (176) than in warm cloudy conditions following
light rainfall (9), or in warm sunny conditions soon after heavy rainfall
(16).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: E Kalahari
Acacia-Baikiaea Woodlands (AT0709), Zambezian Baikiaea Wood-
lands (AT0726), Zambezian and Mopane Woodlands (AT0725),
Southern African Bushveld (AT0717), Southern Miombo Woodlands
(AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 2 069 530 km2; centred in hot lowland sa-
vanna both within and outside of reserves where it is found sporadically
in dung of domestic livestock, perhaps reflecting the measured bias to
omnivore dung; across South African range, habitat transformation
varies from 0 to 40% (SVmp), 0–58% (SVcb), 2–49% (SVl); however,
very widespread, so assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on habitat associations, particu-
larly, effects of woodland clearance; protected in Kruger National Park
(South Africa).

♀ 2 mm

ONTHOPHAGINI
528 SURICATA 6 (2020)

Phalops bubalus
(Harold, 1867c)

No synonyms
Global: LC (see IUCN Red List as Onthophagus – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Onthophagus bubalus: ‘Cap der guten Hoffnung’ [Cape

of Good Hope, South Africa].
Taxonomy: Accepted species; not assigned to a species group; no formal-
ly described synonyms although Harold cites O. bubalus Klug in litt.
and O. varipennis Dejean (catalogue) as synonyms; recently transferred
♂ from Onthophagus Group 13, but horn of males differs to those of other
Phalops; sexually dimorphic (head, prothoracic disc), characters vary in
prominence with body size.
Distribution: Centred on the South African Highveld and cooler coastal
parts of the Eastern Cape lying within the summer rainfall region; pos-
sibly more abundant in drier areas.
Locality data (mean ± SD, range): Altitude: 1 256 ± 417, 145–2 199 m;
annual rainfall: 527 ± 163, 301–843 mm; annual temperature (max. +
min. /2): 15.6 ± 1.4, 11.2–18.7°C (N=108).
Habitat: No adequate quantitative assessment; on the KwaZulu-Natal es-
carpment (29–30°S): limited numbers recorded in grassland on sandy
clay loam only at 1 500 m, not at lower (500 m, 1 000 m) nor higher
altitudes (1 900 m, 2 400 m, 2 800 m); at the SW edge of the range
in E Upper Nama Karoo: recorded at an average of three per sample at
36 study sites on nine farms; DBRU collection records on sand (12),
sandy loam (23), sandy clay loam (33), clay (5) in grassland/pasture
(35), scrub/shrubland (5), open woodland (2).
Food types: No quantitative assessment; DBRU collection records from
cattle (78) and buffalo (1) dung.
Temporal activity: Diel flight periodicity unknown; active primarily
during the summer rainy season (Oct. to May).
Bioregions South Africa: Centred on Dry Highveld Grassland (Gh),
2 mm Mesic Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs)
(Grassland Biome); Albany Thicket Biome (AT); Eastern Upper Karoo
(NKu 4) (Nama Karoo Biome).
Assessment rationale: EOO = 355 495 km2; primarily distributed in
farmland across the grasslands of the South African Highveld where
it readily colonises cattle dung; range extensively transformed by culti-
vation and pasture improvement (1–63%, Gh; 1–69%, Gm; 2–50%,
Gs; 2% NKu); however, widespread occurrence; therefore, assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly,
effects of habitat transformation such as pasture improvement in natu-
ral grassland; protected in Addo Elephant National Park.

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 529

Phalops dregei
(Harold, 1867c)

= Phalops rufosignatus van Lansberge, 1885a (pars)

= Phalops adspersipennis Boheman, 1857, sensu Péringuey, 1901 (pars)
= Phalops congenitus Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus dregei: ‘Cap der guten Hoffnung’, lec-

totype: ‘Pr. b. sp.’ [Cape of Good Hope, South Africa]; P. rufosignatus
(pars): ‘l’Afrique méridionale’ [southern Africa]; P. adspersipennis (pars):
‘Ovampoland (Omrramba), ? Southern Rhodesia’ [Namibia, Zimbabwe];
♂
P. congenitus: ‘Transvaal (Potchefstroom, Lydenburg), British Bechuana-
land (Palapye), Southern Rhodesia (Buluwayo)’ [South Africa, Botswana,
Zimbabwe].
Taxonomy: Accepted species in the fimbriatus group, but see also taxo-
nomic notes under P. rufosignatus van Lansberge; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size;
colour variation from iridescent cupreous in warmer NE lowland range
to black head and thorax with brown elytra in higher SW Highveld
range.
Distribution: Dry savanna in the NE to dry grassland or shrubland in
the SW; primarily found in South Africa with scattered records from
savannas of Zimbabwe, Botswana and Namibia.
Locality data (mean ± SD, range): Altitude: 1 102 ± 341, 106–1 860 m;
annual rainfall: 431 ± 171, 195–769 mm; annual temperature (max. +
min. /2): 17.9 ± 2.4, 13.2–23.2°C (N=178).
Habitat: No adequate quantitative assessment; in Gauteng bushveld: sand
(5), sandy clay loam (8) in grassland (8), open woodland (5), shaded
thickets (0); in open woodland at Leeuwfontein Nature Reserve: clear
bias to stony finer-grained soils: sand (68), sandy loam (109), stony 2 mm
sandy loam (321); DBRU collection records: sand (16), sandy loam (24),
sandy clay loam (11) in grassland/pasture (18), scrub/shrubland (12),
open woodland (10).
Food types: At Phalaborwa: greatest attraction to omnivore dung (pig:
140), frequency lower on ruminant and monogastric herbivore dung
(cattle: 43; elephant: 27); DBRU collection records reflect commonly
sampled dung types: cattle (43), zebra (2), horse (1).
Temporal activity: Diurnal flight activity, during the summer rainy sea-
son (Oct. to Apr.); at Phalaborwa: frequency much greater in hot, dry
conditions (141) than in warm cloudy conditions following light rain-
fall (20), or in warm sunny conditions soon after heavy rainfall (26).
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709),
Southern African Bushveld (AT0717), S Zambezian and Mopane
Woodlands (AT0725).
Bioregions South Africa: Central Lowveld (SVl), Central Bushveld
(SVcb), Eastern Kalahari Bushveld (SVk) (Savanna Biome); N and
S Dry Highveld Grassland (Gh) (Grassland Biome); N Upper Karoo
(NKu) (Nama Karoo Biome); Albany Thicket Biome (AT); margins of
five other bioregions.
Assessment rationale: EOO = 500 430 km2; associated with drier climate
in warm savanna, grassland and Karoo; widely recorded in dung of do-
mestic livestock in farmland despite measured bias to omnivore dung; ♀
2 mm

ONTHOPHAGINI
530 SURICATA 6 (2020)

Phalops dregei (continued)

SW part of range in Karoo little transformed; greater transformation to

NE in SVcb (0–58%) and SVl (2–49%); however, widespread and little
threatened over much of range; therefore, assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on soil and vegetation associations;
protected in various national parks and nature reserves including Kru-
ger, Telperion (South Africa), Etosha (Namibia).

ONTHOPHAGINI
SURICATA 6 (2020) 531

Phalops flavocinctus
(Klug, 1855)

= Onthophagus praeustus Fahraeus, 1857

= Onthophagus (Phalops) flavocinctus var. nigrostriatus d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus flavocinctus: ‘Sena’ [Sena, central Mo-

zambique], lectotype: ‘Port Natal Sinna’ [Unclear: Durban and Sena?];
O. praeustus: ‘prope fluvium Limpopo’ [near Limpopo River, southern
Africa]; var. nigrostriatus: ‘Transvaal’ [South Africa].
Taxonomy: Accepted species in the divisus group; sexually dimorphic
♂
(head, prothoracic disc), characters vary in prominence with body size;
dull, irridescent, colour variation from cupreous to green to blue.
Distribution: Widespread in open savanna on finer-grained soils along
the SE seaboard of Africa: South Africa, Botswana, Zimbabwe, Zam-
bia, Malawi, Mozambique, E Tanzania.
Locality data (mean ± SD, range): Altitude: 716 ± 481, 0–1 652 m; an-
nual rainfall: 677 ± 147, 304–1 191 mm; annual temperature (max. +
min. /2): 20.6 ± 2.3, 14.3–26.1°C (N=156).
Habitat: In Gauteng bushveld: extreme bias to sandy clay loam (20) com-
pared to sand (0) in grassland (16) rather than open woodland (4),
shaded thicket (0); in uMkhuze Game Reserve: similar bias to finer-
grained soils: sand (0.4), sand over clay (3.3), sandy clay loam (9.4),
clay (5.3) in unshaded (158) rather than shaded conditions (24); partly
supported by DBRU collection records: sand (7), sandy loam (21), san-
dy clay loam (21), clay (6) in pasture/grassland (15), shrubland (10),
open woodland (19), forest/thicket (3).
Food types: At Phalaborwa: greatest attraction to omnivore dung (pig:
60), frequency less on ruminant and monogastric herbivore dung (cat-
tle: 6; elephant: 7); DBRU collection records reflect commonly sam- 2 mm
pled dung types: cattle (41), buffalo (3), wildebeest (3), elephant (13),
rhinoceros (6), horse (2), also on single zebra, donkey, warthog, baboon
and impala droppings.
Temporal activity: Diurnal flight activity primarily during the summer
rainy season (Oct. to Mar.); at Phalaborwa: frequency much greater in
hot, dry conditions (58) than in warm cloudy conditions following light
rainfall (2), or, in warm sunny conditions soon after heavy rainfall (13).
Ecoregions Botswana, Zimbabwe, Mozambique: E Kalahari Acacia-
Baikiaea Woodlands (AT0709), Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725), Southern Miombo
Woodlands (AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp) (Savanna Biome); sporadically in warmer N
parts, Dry Highveld Grassland (Gh) and Sub-Escarpment Grassland
(Gs) (Grassland Biome); margins of four other bioregions.
Assessment rationale: EOO = 1 564 855 km2; AOO would be smaller,
restricted by soil and vegetation specialisation; however, habitat trans-
formation through clearance of woodland in main South African part
of range (SVcb: 0–58%; SVl: 2–49%) may not be a great threat owing
to grassland habitat bias; furthermore, clearly widespread, therefore, as-
sessed as Least Concern (LC).
Conservation measures: None recommended; widespread in more open
vegetation on finer-grained soils and well protected in reserves, includ- 2 mm
ing Kruger National Park and uMkhuze Game Reserve (South Africa). ♀

ONTHOPHAGINI
532 SURICATA 6 (2020)

Phalops pauliani
Barbero, Palestrini & Roggero, 2003

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Okosongomingo l49 Otjiwarongo SE 20l7 Ca’ [Namibia].

Taxonomy: Accepted species in the fimbriatus group; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size;
all known individuals iridescent green.
Distribution: Centred on dry upland savanna of N central Namibia with
one inexact record from Alto Cubal, Benguela Province, SW Angola.
♂ Locality data (mean ± SD, range): Altitude: 1 437 ± 122, 1 214–1 577 m;
annual rainfall: 400 ± 57, 296–480 mm; annual temperature (max. +
min. /2): 19.9 ± 1.0, 16.6–21.8°C (N=11).
Habitat: No quantitative assessment; DBRU collection records mainly
on finer-grained soils: sand (1), sandy loam (4), clay (1), in woody
vegetation: shrubland (1), open woodland (5).
Food types: No quantitative assessment; DBRU collection records on
cattle (5) and donkey (1) dung.
Temporal activity: Diurnal flight activity; recorded primarily in the late
summer rainy season (Dec. to May).
Ecoregions Namibia: NW Kalahari Xeric Savanna (AT1309), Angolan
Mopane Woodlands (AT0702).
Assessment rationale: EOO = 66 480 km2 (estimated); recently described
and known primarily from farm rangeland in open N Namibian sa-
vanna woodland; one further record suggests a wider range extending
into Angola; recorded from dung of farm livestock and, apparently, not
threatened at present; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by a quantitative survey to determine its full EOO and eco-
logical associations, particularly if it would be influenced by woodland
2 mm
clearance; not currently known to be protected within any reserve.

2 mm
♀

ONTHOPHAGINI
SURICATA 6 (2020) 533

Phalops plancus
(Erichson, 1843)

= Onthophagus (Phalops) densegranosus d’Orbigny, 1908

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus plancus: ‘Angola’, lectotype: ‘Angola’; O.

(P.) densegranosus: ‘Great Namaqualand’ [SW to W Namibia].
Taxonomy: Accepted iridescent cupreous species in the divisus group;
minor sexual dimorphism (head).
Distribution: Centred in open arid lowland savanna bordering the Namib
and Kaokoveld Deserts in W to NW Namibia and SW Angola.
Locality data (mean ± SD, range): Altitude: 885 ± 336, 283–1 458 m; ♂
annual rainfall: 232 ± 88, 44–375 mm; annual temperature (max. +
min. /2): 17.0 ± 1.2, 14.5–18.8°C (N=14).
Habitat: No quantitative assessment; DBRU collection records entirely on
sand (4) in scrub (2), shrubland (1) or open woodland (1).
Food types: No quantitative assessment; DBRU collection records on
cattle (3) or donkey (1) dung.
Temporal activity: Diurnal flight activity during the late summer rainy
season (Feb. to May).
Ecoregions Namibia: Centred on Namibian Savanna Woodlands
(AT1316), also, margins of Namib Desert (AT1315).
Assessment rationale: EOO = 39 635 km2; AOO may be restricted by soil
specialisation; found in little-transformed, arid savanna used primarily
for grazing of livestock; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; not currently
known from any national park although widely protected in commu-
nity conservation areas.

2 mm

♀
2 mm

ONTHOPHAGINI
534 SURICATA 6 (2020)

Phalops prasinus
(Erichson, 1843)

= Onthophagus adspersipennis Boheman, 1860

= Onthophagus (Phalops) prasinus var. aeraneus d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus prasinus: ‘Angola’, lectotype: ‘Angola’;

O. adspersipennis: ‘regione fluvii Svakop’ [Swakop River region, Namib-
ia]; var. aeraneus: Not stated.
Taxonomy: Accepted species in the prasinus group; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
♂ Distribution: Centred on dry savanna in N Namibia with a dry to mod-
erately moist savanna range continuing along a narrow W coastal belt
in Angola to Democratic Republic of the Congo (DRC); record from
wet tropical Gabon seems doubtful.
Locality data (mean ± SD, range): Altitude: 1 203 ± 313, 536–1 730 m;
annual rainfall: 324 ± 87, 171–507 mm; annual temperature (max. +
min. /2): 18.5 ± 1.3, 16.3–20.7°C (N=32).
Habitat: No quantitative assessment; DBRU collection records suggest
a bias to coarser-grained soils: sand (6), sandy loam (4), clay (1), in
woody vegetation: scrub (1), shrubland (5), open woodland (6).
Food types: No quantitative assessment; DBRU collection records from
cattle (8), horse (2) and donkey (2) dung.
Temporal activity: Diurnal flight activity during the late summer rainy
season (Dec. to Apr.).
Ecoregions Namibia: Centred on NW Kalahari Xeric Savanna (AT1309),
N Namibian Savanna Woodlands (AT1316); marginal in Angolan Mo-
pane Woodlands (AT0702), Namib Desert (AT1315).
Assessment rationale: EOO = 115 340 km2; widespread on sandy soils in
N Namibian savanna woodland; EOO primarily coincides with farm
2 mm
rangeland where it has been recorded on the dung of farm livestock;
probably not currently threatened by habitat transformation, therefore,
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Etosha National Park (Namibia).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 535

Phalops pyroides
(d’Orbigny, 1908)

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: As Onthophagus (Phalops) pyroides: ‘Damaraland (Okah-

andja; Windhoek)’, lectotype: ‘Okahandja’ [Namibia].
Taxonomy: Accepted iridescent cupreous species in the fimbriatus group;
sexually dimorphic (head, prothoracic disc), characters vary in promi-
nence with body size.
Distribution: Centred on dry N Namibian savanna woodland and arid ♂
Northern Cape Karoo, South Africa; apparent disjunction may be a
collecting artefact.
Locality data (mean ± SD, range): Altitude: 1 133 ± 224, 783–1 928 m;
annual rainfall: 242 ± 66, 133–439 mm; annual temperature (max. +
min. /2): 18.7 ± 0.9, 15.8–22.8°C (N=61).
Habitat: No adequate quantitative data; in Northern Cape: similar abun-
dance on sand (1.01) and sandy loam (0.98) in grassland, scrub or
shrubland; DBRU collection records from N Namibia on sand (1),
sandy loam (3), sandy clay loam (1), clay (1) in scrub (1), shrubland
(1), open woodland (4).
Food types: No quantitative data; DBRU collection records on cattle (5)
and donkey (1) dung.
Temporal activity: Diurnal flight activity in the late summer rainy season
(Jan. to Apr.).
Ecoregions Namibia: Nama Karoo (AT1314), NW Kalahari Xeric Sa-
vanna (AT1309), Namibian Savanna Woodlands (AT1316), S Angolan
Mopane Woodlands (AT0702).
2 mm
Bioregions South Africa: NE Bushmanland (NKb) (Nama Karoo Biome)
and incorporated outlier patches of Kalahari Duneveld (SVkd) (Savan-
na Biome).
Assessment rationale: EOO = 122 540 km2; observed habitat association
varies from dry woody savanna (N Namibia) to arid grass, scrub, and
shrubland Karoo of NE Bushmanland (W Griqualand, South Africa);
habitat transformation low as land use is dominated by grazing of do-
mestic livestock to whose dung it is readily attracted; therefore, assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by a survey to determine if its distribution is disjunct or contin-
uous; owing to the degree of variability in observations, a quantitative
study of ecological associations would also be useful in both the S and
the N; not known to occur in any protected area.

♀
2 mm

ONTHOPHAGINI
536 SURICATA 6 (2020)

Phalops rufosignatus
van Lansberge, 1885a

= Phalops euplynes Bates, 1888

= Phalops adspersipennis Boheman, 1857, sensu Péringuey, 1901 (pars)
= Onthophagus (Phalops) lansbergei d’Orbigny, 1902 (pars)
Global: LC

J A S O N D J F M A M J

Type localities: P. rufosignatus (pars): ‘l’Afrique méridionale’ [southern

Africa]; P. euplynes: ‘Damara Land’ [Namibia]; P. adspersipennis sensu
Péringuey (pars) ‘Cape Colony (Calvinia, Bushmanland, Namaqua­
land)’ [South Africa]; O. (P.) lansbergei (pars): invalidated nom nov. for
♂ P. rufosignatus.
Taxonomy: Accepted species in the divisus group, but designated type
series comprises one male P. dregei Harold, 1867, and one female P. ru­
fosignatus; sexually dimorphic (head, prothoracic disc), characters vary
in prominence with body size.
Distribution: Centred on the arid late summer rainfall region in SW
Africa: South Africa, Namibia, SW Angola, extreme SW Botswana;
Tanzania record requires further validation.
Locality data (mean ± SD, range): Altitude: 1 161 ± 348, 0–1 928 m;
annual rainfall: 311 ± 101, 59–486 mm; annual temperature (max. +
min. /2): 18.6 ± 2.2, 13.3–22.5°C (N=76).
Habitat: No quantitative assessment; DBRU collection records from sand
(31), sandy loam (17), sandy clay loam (23), clay (1) in grassland/pas-
ture (14), scrub (17), shrubland (16), open woodland (15).
Food types: In Botswana: bias to dung of omnivores (pig: 101), pellet-
dropping ruminant and monogastric herbivores (sheep: 74; elephant:
35) rather than pad-dropping ruminant herbivores (cattle: 9); DBRU
collection records reflect commonly sampled dung types: cattle (57),
buffalo (1), wildebeest (2), elephant (2), zebra (3), horse (3), donkey
2 mm (4).
Temporal activity: Diurnal flight activity, primarily, during the late sum-
mer rainy season (Dec. to May) with few records in July, Oct. and Nov.
Ecoregions Namibia: Nama Karoo (AT1314), W Kalahari Xeric Savanna
(AT1309), Namibian Savanna Woodlands (AT1316), S Angolan Mo-
pane Woodlands (AT0702).
Bioregions South Africa: Upper Karoo (NKu), Bushmanland (NKb)
(Nama Karoo Biome), Eastern Kalahari Bushveld (SVk), Kalahari
Duneveld (SVkd) (Savanna Biome); also Albany Thicket Biome (AT).
Assessment rationale: EOO = 591 895 km2; widespread across various
soil and vegetation types in a little-transformed region used primarily
for the grazing of domestic livestock to whose dung it is readily attract-
ed despite a measured bias to omnivore dung; therefore, assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations; protected in Kga-
lagadi Transfrontier Park (South Africa, Botswana), Etosha National
Park (Namibia).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 537

Phalops smaragdinus
(Harold, 1875)

= Onthophagus tarsatus Péringuey, 1888

= Onthophagus (Phalops) smaragdinus var. coerulosus d’Orbigny, 1902
= Onthophagus (Phalops) smaragdinus var. cuprinus d’Orbigny, 1902
= Onthophagus smaragdinus n. f. idiomelas Kuntzen, 1913
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus smaragdinus: ‘Abyssinia australis’ [?south-

ern Ethiopia], lectotype: ‘Holub’ [possibly refers to the 19th century
collector, E. Holub, who travelled South and central Africa]; O. tarsa­
tus: ‘Rustenburg and Potchefstroom, Transvaal’ [North-West Province, ♂
South Africa]; var. coerulosus: ‘Transvaal’ [South Africa]; var. cuprinus:
‘Zanzibar’ [Tanzania]; n. f. idiomelas: Not stated.
Taxonomy: Accepted species in the fimbriatus group, but cited NE Afri-
can occurrence of original type, compared to SE African occurrence of
lectotype and varieties, suggests re-evaluation might be warranted, par-
ticularly, in comparison with the close East African relative, P. lamin­
ifrons Fairmaire, 1882, described from Zanzibar; sexually dimorphic
(head, prothoraccic disc), characters vary in prominence with body
size; iridescent colour variation from cupreous to green to blue.
Distribution: Validated distribution centre in SE Africa: South Africa, E Bot­
swana, Zimbabwe, Malawi, Mozambique; also reported from Zambia, SE
Tanzania; records from Namibia, N Tanzania, Uganda, and original type
locality of Ethiopia (cited as Abyssinia australis) require further validation.
Locality data (mean ± SD, range): Altitude: 825 ± 473, 0–1 785 m; an-
nual rainfall: 731 ± 178, 304–1 259 mm; annual temperature (max. +
min. /2): 20.4 ± 2.1, 14.8–25.4°C (N=107).
Habitat: In Gauteng bushveld: extreme bias to sand (96) rather than san- 2 mm
dy clay loam (0) with very strong bias to open woodland (86) rather
than grassland (9) or shaded thickets (1); weakly supported by DBRU
collection records on sand (14), sandy loam (16), sandy clay loam (10)
in pasture/grassland (13), open shrub/woodland (17), thickets (1).
Food types: No adequate quantitative assessment; at Phalaborwa: only
recorded to pig dung (8), none to cattle or elephant dung; DBRU col-
lection records from dung of cattle (42), elephant (1), donkey (1), hu-
man/cattle mix (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.).
Ecoregions Botswana, Zimbabwe, Mozambique: E Kalahari Acacia-
Baikiaea Woodlands (AT0709), Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725), Southern Miombo
Woodlands (AT0719), Maputaland Coastal Forest Mosaic (AT0119),
margins of two other ecoregions.
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome); N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 1 479 555 km2; AOO possibly reduced by soil
and vegetation bias; in South Africa, habitat transformation across its range
varies from 0 to 58% (SVcb) and 2–49% (SVl), with clearance of wood-
land possibly detrimental to population density; currently assessed as Least
Concern (LC) on basis of high frequency of records across a large EOO.
Conservation measures: Assessment of conservation status would be impro­
ved by further quantitative ecological data to support soil, vegetation and 2 mm

dung type specialisation; protected in Tswaing Nature Reserve, Kruger Na-

tional Park (South Africa), Lake Mutirikwe Game Reserve (Zimbabwe). ♀

ONTHOPHAGINI
538 SURICATA 6 (2020)

Phalops wittei
(Harold, 1867c)

= Onthophagus boschimanus Péringuey, 1888

= Onthophagus adspersipennis Boheman, 1857, sensu Péringuey, 1901 (pars)
= Onthophagus (Phalops) rufatus d’Orbigny, 1902
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus wittei: ‘innere Südafrika, Owampo’, lec-

totype: ‘Owampo’ [South African interior; Ovamboland, Namibia];
O. boschimanus: ‘Bushmanland’ [Northern Cape, South Africa]; O. ad­
spersipennis (pars): ‘Ovampoland (Omrramba)’ [Omuramba, Namib-
♂ ia]; O. (P.) rufatus: ‘Angola: Benguella’ [Benguela].
Taxonomy: Accepted species in the fimbriatus group with black head and
prothoracic disc with brown elytra; sexually dimorphic (head, protho-
racic disc), characters vary in prominence with body size; synonymy of
Phalops rufatus is based on the single female type specimen comprising
parts of two different species (Barbero et al. 2003), however, this deci-
sion may require re-examination, as material from arid NW Namibia
appears to differ a little from P. wittei.
Distribution: Centred on deep sands in drier climates of southern Afri-
ca: N central South Africa, Namibia, Botswana, SW Zimbabwe, SW
Angola.
Locality data (mean ± SD, range): Altitude: 1 102 ± 217, 0–2 199 m;
annual rainfall: 300 ± 136, 59–741 mm; annual temperature (max. +
min. /2): 19.2 ± 1.6, 11.2–23.1°C (N=254).
Habitat: In Gauteng bushveld: extreme bias to sand (41) as opposed to
sandy clay loam (0) and grassland (39) as opposed to open woodland
(2) or shaded thickets (0); largely supported by DBRU collection re-
cords on sand (58), sandy loam (6), sandy clay loam (4) in grassland/
pasture (18), scrub (13), shrubland (20), open woodland (12).
Food types: In Botswana: bias to dung of omnivores (pig: 190), pellet-
dropping ruminant and monogastric herbivores (sheep: 103; elephant:
2 mm
77) rather than pad-dropping ruminant herbivores (cattle: 8); DBRU
collection records mostly reflect commonly sampled dung types: ba-
boon (1), cattle (59), wildebeest (1), elephant (1), horse (2), donkey
(2).
Temporal activity: Diurnal flight activity, primarily during the mid- and
late summer rainy seasons (Oct. to Apr./May).
Ecoregions Namibia, Botswana, Zimbabwe: Widespread across deep sands
of Namibian Savanna Woodlands (AT1316), Kalahari Xeric Savanna
(AT1309), Kalahari Acacia-Baikiaea Woodlands (AT0709), S Angolan
Mopane Woodlands (AT0702), margins of four adjoining ecoregions.
Bioregions South Africa: Deep sands of Bushmanland (NKb) (Nama
Karoo Biome); Dry Highveld Grassland (Gh) (Grassland Biome); Ka-
lahari Duneveld (SVkd), Eastern Kalahari Bushveld (SVk), Central
Bushveld (SVcb) (Savanna Biome); margins of three other bioregions.
Assessment rationale: EOO = 1 562 330 km2; AOO reduced by soil spe-
cialisation and bias to open vegetation, but much of range comprises
continuous, little-transformed, sparsely vegetated, deep Kalahari sands
used primarily for grazing of domestic livestock to which P. wittei is
widely attracted despite a recorded bias to omnivore dung; therefore,
assessed as Least Concern (LC).
Conservation measures: None recommended; protected in the private
Tswalu Kalahari Reserve, Kgalagadi Transfrontier Park (South Africa,
♀ Botswana).
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 539

Phalops zuninoi
Barbero, Palestrini & Roggero, 2003

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Bicuari N. P. Angola (Nr Campo)’ [near camp in Bicuar

National Park, Angola].
Taxonomy: Accepted species in the boschas group; sexually dimorphic
(head, prothoracic disc), characters vary in prominence with body size.
Distribution: Recently described from moist climate at S edge of Bié Pla-
teau and adjoining deep sands, S Angola; also recorded on deep sands
of Ovamboland floodplain, N Namibia, and Liuwa floodplain, SW ♂
Zambia.
Locality data (mean ± SD, range): Altitude: 1 158 ± 70, 1 109–1 207 m;
annual rainfall: 674 ± 197, 534–813 mm; annual temperature (max. +
min. /2): 21.8 ± 0.7, 21.4–22.3°C (N=2).
Habitat: No quantitative assessment; DBRU collection records primarily
on sand (3), but also sandy loam (1) in shrubland (2) or open wood-
land (2).
Food types: No quantitative assessment; DBRU collection records from
the dung of cattle (3) and elephant (1).
Temporal activity: Presumed to show diurnal flight activity; recorded in
the summer rainy season (Nov., Dec.).
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702).
Assessment rationale: EOO = 197 250 km2; AOO possibly more limited
due to soil and woody vegetation habitat bias, but very poorly known
ecologically; owing to limited data, assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative
survey is required to determine the full EOO, AOO and ecological
associations; protected in Bicuar National Park, Angola.

2 mm

2 mm
♀

ONTHOPHAGINI
540 SURICATA 6 (2020)

Genus Proagoderus van Lansberge, 1883

Type species and designation: Onthophagus (Proagoderus) ritsemae van Lansberge, 1883, by monotypy.
Synonyms: = Onthotrogus Motschulsky, 1860; Type species: Copris auratus Fabricius, 1801, by
monotypy (suppressed by ICZN 1993: Opinion 1708).
= Melinocerus Motschulsky, 1860; Type species: Copris harpax Fabricius, 1801, by
monotypy (nomen oblitum).
= Tauronthophagus Shipp, 1895c: Type species: Onthophagus rangifer Klug, 1855, by
original designation.
Last review: Reviewed as a subgenus of Onthophagus Latreille, 1802, by Palestrini (1992, all spe-
cies); with revision of a species complex (Moretto 2004) and addition of new Afrotrop-
ical species by Josso and Prévost (2001), Moretto and Nicolas (2002, 2004), Moretto
(2004, 2013b), Josso (2014).

Proagoderus van Lansberge, 1883, was described as a subgenus of Onthophagus Latreille, 1802. More recently, it has been
cited as a genus (Cambefort 1991; Davis et al. 2008), but only formally raised to generic status by Moretto (2013b).

On the basis of long-term usage, the name has been conserved as Proagoderus despite the precedence of Onthotrogus Mot-
schulsky, 1860, which has been suppressed and placed on the list of invalid names by the International Commission of
Zoological Nomenclature (ICZN 1993: Opinion 1708). As Proagoderus, thus, becomes a protected name (nomen protec­
tum), the forgotten name (nomen oblitum) of Melinocerus Motschulsky, 1860, may also be considered a synonym (Moretto
2013b) despite its precedence.

Proagoderus currently comprises 127 species primarily centred in the Afrotropical region (117) with a few centred in the
Oriental region (10). The genus is divided into nine species groups (see Palestrini 1992) with such a range in morphology
that each may deserve subgeneric or even generic status. A review of Proagoderus and its synonyms may thus be justified,
as the four type species belong to different species groups.

In South Africa, Botswana and Namibia, Proagoderus is represented by 14 species belonging to seven groups. Six of these
species belonging to Groups 2, 4, 5 or 7, are primarily associated with the dung of monogastric herbivores, e.g. elephant,
zebra, accounting for the patchy distributions centred around game reserves.

In southern Africa:
1. Group confossus: Comprises three species; two with E savanna ranges in open woodland or dense woodland on
clay soils; one other centred on SE coast and highland grassland of South Africa plus N Namib-
ian uplands where it is a rarity.
2. Group auratus: One species with an E savanna range extending N into the E tropics.
3. Group bicallosus: One species with a savanna range extending N into the tropics.
4. Group gibbiramus: Two rare species with savanna or W savanna ranges.
5. Group prostans: One species with an E savanna range extending N into the E tropics.
6. Group ramosicornis: One species with E savanna range extending N into the E tropics.
7. Group dives: Comprises five species, four with ranges centred on the E coast sands: E coast forest (1), E coast
grass plus E savanna (2), E coast plus E upland grasslands; the fifth centred on deep Kalahari sands.

Most Proagoderus species are assessed as Least Concern (LC) as they are widespread. However, with the exception of species
showing Kalahari or highland ranges, most are more abundant in reserves than in farmland, particularly those that are
frequently recorded on elephant or zebra dung. The members of the gibbiramus group are assessed as Data Deficient (DD)
as they are recorded very infrequently, and then only in elephant dung in reserves.

ONTHOPHAGINI
SURICATA 6 (2020) 541

Proagoderus aciculatus
(Fahraeus, 1857)

= Onthophagus (Proagoderus) aciculatus var. ahenus d’Orbigny, 1902

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus aciculatus: ‘in terra Natalensi et prope

fluvium Limpopo’ [KwaZulu-Natal, South Africa, and near Limpopo
River]; var. ahenus: ‘Mozambique: Delagoa-Bay’ [Maputo, Mozam-
bique].
Taxonomy: Accepted dives group species; muted metallic colour variation
from dull cupreous to dull green to dull blue.
Distribution: Vegetation offering shade on SE coastal sands: South Africa,
Mozambique.
Locality data (mean ± SD, range): Altitude: 43 ± 58, 0–273 m; annual
rainfall: 862 ± 120, 629–1 254 mm; annual temperature (max. + min.
/2): 21.8 ± 1.2, 18.4–24.8°C (N=74).
Habitat: In uMkhuze Game Reserve: extreme bias to coarse-grained soils,
deep sand (462.9), sand over clay (93.9), sandy clay loam (0.4), clay
(1.3); on deep sand in Maputo Elephant Reserve: bias to cooler coastal
dune forest (303.9), less abundant in warmer inland sand forest (large
patch: 41.2; medium: 38.7; small: 24.3), rare in grassland (2.2); across
a chronosequence of regenerating dune vegetation at Richards Bay:
uncommon in 0–5 yr grassland (4.7), more abundant in 3–15 yr dense
(18.3) or 12–33 yr open understorey woodland (38.0); largely support-
ed by DBRU collection records: sand (23), sandy loam (1), sandy clay
loam (1) in grassland (12), open woodland (7), forest (17).
Food types: No quantitative assessment; DBRU collection records on var-
ious dung types with possible bias to omnivore dung: human/human +
cattle mix (13), horse (2), elephant (1), cattle (9).
Temporal activity: Diurnal flight activity during much of the year (Aug.
to May); observed to pitch on leaves in forest.
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128) (not
shown on map). 2 mm
Bioregions South Africa: Centred on Indian Ocean Coastal Belt Biome
(CB); SE margins of Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 33 450 km2; AOO smaller, restricted
to shaded situations on moist E coastal sands; therefore, susceptible
to clearance of forest or dense woodland for coastal development; in
South Africa, vegetation units of the coastal belt are some 30–50%
transformed by plantations, cultivation and urban sprawl (CB1, CB2);
however, P. aciculatus is locally abundant and well protected in reserves;
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on food type associations; protected in
Hluhluwe–iMfolozi, uMkhuze game reserves (South Africa), Maputo
Elephant Reserve (Mozambique).

ONTHOPHAGINI
542 SURICATA 6 (2020)

Proagoderus aureiceps
(d’Orbigny, 1902)

= Onthophagus sapphyrinus Péringuey, 1901

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus (Proagoderus) aureiceps: ‘Mozambique:

district de Delagoa....Delagoa-Bay’ [Maputo, Mozambique], lecto-
type: Maputo; O. sapphyrinus: ‘Southern Rhodesia (Salisbury, Victoria
Falls, Limpopo River), Ovampoland, (Okovango River, Humbe Om-
rramba), Transvaal (Rustenburg), Mozambique (Rikatla)’ [Zimbabwe,
South Africa, Mozambique].
Taxonomy: Accepted dives group species, but confusion in literature in-
dicates that revision is required; P. aureiceps may be a junior synonym
of P. sapphirinus (Fahraeus, 1857), whose name suggests a blue type
specimen; details and cited localities for the synonym, O. sapphyrinus
Fahraeus sensu Péringuey, include a green/blue coastal form (Rikatla)
that would equate to P. aureiceps, but primarily comprises green indi-
viduals with yellow/red elytra that would belong to a different species
(see species page for P. sapphirinus (Fahraeus, 1857)).
Distribution: Abundant on coastal sands in SE Mozambique and NE
South Africa; also moister parts of Limpopo River Valley; uncommon
in the N interior of South Africa where scattered records border the
Soutpansberg and Waterberg and overlap with the closely related
species (green with red elytra) that is currently listed under the name,
P. sapphirinus (Fahraeus, 1857).
Locality data (mean ± SD, range): Altitude: 157 ± 303, 0–1 376 m; an-
nual rainfall: 763 ± 148, 422–1 006 mm; annual temperature (max. +
min. /2): 22.0 ± 1.0, 17.6–23.4°C (N=63).
Habitat: In uMkhuze Game Reserve: extreme bias to coarse-grained soils,
deep sand (279.3), sand over clay (23.7), sandy clay loam (0.1), clay
(0.2); on deep sand in Maputo Elephant Reserve: bias to grassland
(54.8), much less abundant in warmer inland sand forest (large patch:
3.6; medium: 4.6; small: 3.5) and cooler coastal dune forest (1.0);
across a chronosequence of regenerating dune vegetation at Richards
Bay: primarily in 0–5 yr grassland (3.5), rare in 3–15 yr dense (0.1)
2 mm or 12–33 yr open understorey woodland (0.2); largely supported by
DBRU collection records: sand (16), sandy loam (4) in grassland (11),
open woodland (9), forest (4).
Food types: No quantitative assessment; DBRU collection records on various dung types: human/human + cattle mix (4),
elephant (1), horse (2), cattle (1), wildebeest (2).
Temporal activity: Diurnal flight activity; recorded primarily in the summer rainy season (Oct. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119), Southern Zanzibar-Inhambane Coastal Forest Mosaic
(AT0128).
Bioregions South Africa: Centred primarily on the N Indian Ocean Coastal Belt Biome (CB); sand patches in Lowveld (SVl),
Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 192 750 km2; AOO much smaller, range restricted to coastal sands where it is abundant and
inland sand outliers where it is uncommon; in South Africa vegetation units of the coastal belt are some 30–50% transformed
by plantations, cultivation and urban sprawl (CB1, CB2); centred on grassland habitats, so perhaps not that influenced by
clearance of woody vegetation; abundant in reserves so assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be improved by quantitative data on food type associations;
protected in uMkhuze Game Reserve, iSimangaliso Wetland Park (World Heritage Site) (South Africa), Maputo Elephant
Reserve (Mozambique).

ONTHOPHAGINI
SURICATA 6 (2020) 543

Proagoderus bicallosus
(Klug, 1855)

= Onthophagus metallicus Fahraeus, 1857

= Onthophagus (Proagoderus) bicallosus var. olivicolor d’Orbigny, 1902
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Onthophagus bicallosus: ‘Sena’ [central Mozambique];

O. metallicus: ‘prope fluvium Limpopo’ [near Limpopo River]; var. ol­
ivicolor: ‘région du lac Tanganyika’ [Lake Tanzania region].
Taxonomy: Accepted bicallosus group species; metallic colour variation
from cupreous to green to olive; cupreous in southern Africa; minor
sexual dimorphism (prothoracic disc). ♂
Distribution: Widespread, but patchy occurrence across southern to E
African lowland savannas owing to dung specialisation: NE South
Africa, N Namibia, N Botswana, Angola, Zimbabwe, Mozambique,
Tanzania, Kenya; also reported from Malawi, Zambia.
Locality data (mean ± SD, range): Altitude: 517 ± 401, 0–1 207 m;
annual rainfall: 656 ± 122, 418–971 mm; annual temperature (max. +
min. /2): 22.3 ± 0.9, 20.5–25.7°C (N=52).
Habitat: No quantitative assessment; DBRU collection records suggest a
bias to coarser-grained soils: sand (7), sandy loam (6), in woody vegeta-
tion: open shrub-woodland (13), grassland (2).
Food types: In Botswana: strong bias to monogastric herbivore (elephant:
20) and omnivore dung (pig: 12), few on ruminant herbivore (cattle:
2; sheep: 2) and carrion (chicken livers: 0); partly supported by DBRU
collection records: elephant (15), rhinoceros (3), donkey (1), cattle (6).
Temporal activity: Diurnal flight activity; recorded primarily in the sum-
mer rainy season (Oct. to Apr.), also Aug. at the warmer coastline.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique:, Centred
on Angolan Mopane Woodlands (AT0702), Zambezian and Mopane
Woodlands (AT0725), Zambezian Baikiaea Woodlands (AT0726),
Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Centred on Lowveld (SVl), Mopane (SVmp) 5 mm
(Savanna Biome).
Assessment rationale: EOO = 3 227 420 km2; AOO would be much
smaller since modern records (52 out of 68) are primarily from game
reserves; recent records centred in two regions (see map); in South Afri-
ca: recent records only from reserves containing monogastric herbivores
despite a secondary strong association with omnivore dung; at present
assessed as Least Concern (LC) owing to widespread range and occur-
rence in many reserves.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations to determine if
AOO is influenced by soil and vegetation type specialisation; current
AOO primarily restricted to reserves containing elephants and rhinoc-
eros whose continuing protection would be of great importance for the
conservation of this species; protected in various national parks and
game reserves, including, Kruger, Hluhluwe–iMfolozi (South Africa),
Chobe (Botswana), Manyara (Tanzania).

ONTHOPHAGINI
544 SURICATA 6 (2020)

Proagoderus chalcostolus
(d’Orbigny, 1902)

= Onthophagus (Proagoderus) brucei var. panchlorus d’Orbigny, 1913

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus (Proagoderus) brucei var. chalcostolus:

‘Natal......Cap de Bonne-Espérance’, lectotype: ‘Natal’; var. panchlorus:
Not stated, lectotype: ‘Natal’ [South Africa: KwaZulu-Natal and Cape
of Good Hope; latter locality inexact].
Taxonomy: Accepted dives group species; metallic colour variation from
cupreus to more frequent green; sexually dimorphic (head, prothoracic
♂ disc), characters vary in prominence with body size.
Distribution: Centred on moist uplands and moist coastal regions of SE
Africa: NE South Africa, S Mozambique, Eswatini, E central Zimbab­
we; Botswana record requires validation.
Locality data (mean ± SD, range): Altitude: 1 026 ± 587, 0–2 392 m;
annual rainfall: 829 ± 150, 428–1 415 mm; annual temperature (max.
+ min. /2): 18.0 ± 2.6, 9.9–23.5°C (N=136).
Habitat: No quantitative assessment: DBRU collection records suggest a
bias to finer-grained soils: clay (1), sandy clay loam (11), sandy loam
(14), sand (5) in open vegetation: pasture/grassland (23), open wood-
land (5), forest (2); one record from a fallow crop field.
Food types: No quantitative assessment: DBRU collection records pri-
marily from cattle dung (39), also dung of rhinoceros, horse, zebra,
human, buck pellets (all 1, except horse: 2).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.); also active in May at the warmer E coastline.
Ecoregions Zimbabwe, Mozambique: Maputaland Coastal Forest Mosa-
ic (AT0119), Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Centred on Indian Ocean Coastal Belt
Biome (CB); warmer NE Mesic Highveld Grassland (Gm), N Sub-
Escarpment Grassland (Gs) (Grassland Biome); also moist upland is-
2 mm lands (Waterberg, Soutpansberg) in Central Bushveld (SVcb) (Savanna
Biome); margins of two adjacent bioregions.
Assessment rationale: EOO = 232 570 km2; widespread in warm, moist
upland grasslands dominated by farmland where it readily colonises
the dung of domestic livestock, especially cattle; in South Africa: ap-
preciable transformation due to tree plantations and cultivation in up-
land bioregions (0–69%, Gm; 10–50%, Gs), but also found along the
coastline, much of which is protected; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, particularly pos-
sible effects of pasture improvement; protected along the SE coastline
in iSimangaliso Wetland Park (World Heritage Site) (South Africa).

ONTHOPHAGINI
SURICATA 6 (2020) 545

Proagoderus dives
(Harold, 1877b)

= Onthophagus deyrollei Raffray, 1877

= Onthophagus optivus Péringuey, 1901
= Onthophagus (Proagoderus) dives var. funereus d’Orbigny, 1902
= Onthophagus (Proagoderus) dives var. semicuprinus d’Orbigny, 1913
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus dives: ‘Nyassa’, lectotype: Nyassa [Lake

Malawi region]; O. deyrollei: ‘Zanguebar, Bagamoyo’ [Bagamoyo, NE
Tanzania]; O. optivus: ‘Southern Rhodesia (Mtoko)’ [Zimbabwe]; var.
funereus: ‘M’honda dans l’Ouzigoua’ [Mhonda, Tanzania]; var. semicu­
prinus: Not stated.
Taxonomy: Accepted dives group species, metallic colour variation from
cupreous to green to blue, even black; in southern Africa: cupreous
inland, green to blue on the coast.
Distribution: Widespread, but somewhat patchy distribution across moist
savanna from southern to E Africa, probably due to soil and vegetation
specialisation: South Africa, Mozambique, Zimbabwe, Zambia, Tan-
zania; also reported from Malawi; Kenyan record requires validation;
black square represents a questionable record.
Locality data (mean ± SD, range): Altitude: 411 ± 436, 0–1 410 m; an-
nual rainfall: 805 ± 211, 383–1 270 mm; annual temperature (max. +
min. /2): 22.1 ± 2.1, 17.1–25.9°C (N=57).
Habitat: In uMkhuze Game Reserve: extreme bias to coarse-grained soils,
deep sand (125.0), sand over clay (22.5), sandy clay loam (0.1), clay
(0.1); no quantitative assessment of vegetation association; DBRU col-
lection records support a bias to coarse-grained soil association: sand
(16), sandy loam (3), sandy clay loam (4), in shaded vegetation: forest
(14), open woodland (16), grassland/pasture (6).
Food types: No quantitative assessment; DBRU collection records on var-
ious dung types with possible bias to omnivore dung: human/human
+ cattle mix (14), baboon (3), elephant (2), rhinoceros (2), horse (3),
cattle (9), wildebeest (1).
Temporal activity: Diurnal flight activity during the summer rainy season 2 mm
(Oct. to Apr.)
Ecoregions Zimbabwe, Mozambique: Zambezian and Mopane Wood-
lands (AT0725), Southern Miombo Woodlands (AT0719), Maputa-
land Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Centred primarily on N Indian Ocean Coastal
Belt Biome (CB); sand patches with shady vegetation in Lowveld (SVl),
Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 1 444 830 km2; AOO presumably much
smaller due to sand and shade specialisation; widespread, but cupreous va-
riety uncommon near Tzaneen (South Africa) where there has been much
vegetation transformation; also in South Africa, green variety abundant
in coastal shade on deep sand; therefore, assessed as Least Concern (LC).
Conservation measures: Presumably, protection of dense woody vege-
tation on deep sand patches would be essential for the conservation
of this species; however, assessment of conservation status would be
improved by quantitative data on vegetation and food type association;
cupreous variety protected in the N Kruger National Park, green variety
protected in uMkhuze Game Reserve and iSimangaliso Wetland Park
(World Heritage Site) (South Africa).

ONTHOPHAGINI
546 SURICATA 6 (2020)

Proagoderus furcifer
(Boheman, 1860)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: As Onthophagus furcifer: ‘juxta lacum N’Gami’ [near Lake

Ngami, Botswana].
Taxonomy: Accepted gibbiramus group species; Onthophagus plato Bates,
1888, made a synonym by Péringuey (1901), but considered incorrect;
strong sexual dimorphism (head, prothoracic disc), prominence of
characters varies with body size.
♂ Distribution: Dry southern African savanna woodlands: South Africa,
Zimbabwe, Botswana, Namibia.
Locality data (mean ± SD, range): Altitude: 951 ± 280, 429–1 364 m;
annual rainfall: 555 ± 85, 414–672 mm; annual temperature (max. +
min. /2): 22.2 ± 0.7, 21.2–23.1°C (N=7).
Habitat: No quantitative assessment; limited DBRU and UP collection
records inconclusive; sand (2), sandy loam (3), clay (1) in shrubland
(1), open woodland (3), fallow crop field (1).
Food types: No quantitative assessment; limited DBRU collection records
primarily on monogastric herbivore dung: elephant (3), rhinoceros (1),
cattle (1).
Temporal activity: Diurnal flight activity during the summer rainy season
(Nov to Mar.); observed flying early in the day.
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian and Mopane Woodlands (AT0725),
Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Deep sand patch in Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 267 405 km2; AOO probably much
smaller; rarely observed; of six recent records, five in national parks,
5 mm
one caught flying and four on elephant or rhinoceros dung; probably
a monogastric dung specialist, possibly on coarse-grained sandy soils
in woodland; owing to its rarity and likely specialisation, probably de-
serves a threat category, but currently assessed as Data Deficient (DD)
owing to limited records.
Conservation measures: A quantitative survey is required to better de-
termine the EOO, ecological associations and AOO of this species
before an accurate assessment may be made of its conservation status;
however, currently protected in several national parks: N Kruger (South
Africa), Hwange (Zimbabwe), Chobe (Botswana).

♀
5 mm

ONTHOPHAGINI
SURICATA 6 (2020) 547

Proagoderus lanista
(Castelnau, 1840)

= Onthophagus tubericollis Castelnau, 1840

= Onthophagus aulicus Fahraeus, 1857
= Onthophagus laniger Chevrolat in litt. Harold, 1867c
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: As Ontophagus lanista: ‘Cap de Bonne-Espérance’ [Cape of

Good Hope, South Africa]; O. tubericollis: ‘Cap de Bonne-Espérance’;
O. aulicus: ‘in terra Natalensi’ [KwaZulu-Natal, South Africa]; O. lan­
iger: Not stated. ♂
Taxonomy: Accepted confossus group species; strong sexual dimorphism
(head, prothoracic disc); prominence of characters varies with body
size.
Distribution: Highveld and SE coastline of South Africa with limited
scattered records on the plateau of N Namibia; records reported for
Botswana and Zambia considered of doubtful validity.
Locality data (mean ± SD, range): Altitude: 1 193 ± 491, 0–2 199 m;
annual rainfall: 672 ± 173, 305–982 mm; annual temperature (max. +
min. /2): 16.7 ± 2.3, 11.2–22.3°C (N=73).
Habitat: No adequate quantitative assessment; at Foggy Valley: primar-
ily in natural Themeda grassland (21), few in disturbed regenerating
vegetation of crop fields (4) or improved Kikuyu pasture (0); DBRU
collection records suggest bias to finer-grained soils: sand (3), sandy
loam (11), sandy clay loam (14), clay (3) in open vegetation: pasture/
grassland (19), scrub/shrubland (3), open woodland (1); few in fallow
crop fields (2).
Food types: No quantitative assessment; DBRU collection records pri-
marily on cattle dung (32); also horse (1) and zebra dung (2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.); also Aug. and May on warmer NE coastline. 2 mm
Ecoregions Namibia: NW Kalahari Xeric Savanna (AT1309), Angolan
Mopane Woodlands (AT0702).
Bioregions South Africa: Centred on Indian Ocean Coastal Belt Biome
(CB); Sub-Escarpment Savanna (SVs) (Savanna Biome), E Dry High-
veld Grassland (Gh), Mesic Highveld Grassland (Gm), Sub-Escarpment
Grassland (Gs) (Grassland Biome); marginal occurrence in two adja-
cent bioregions.
Assessment rationale: EOO = 300 285 km2; uncommon on N plateau
of Namibia but commonly found in cattle dung on farms across the
Highveld grasslands of South Africa; results from Foggy Valley suggest
that habitat transformation, including pasture improvement, are det-
rimental; transformation through cultivation and development of tree
plantations varies from 4 to 60% (Gh), 6–69% (Gm) and 2–50% (Gs);
nevertheless, widespread in SE South Africa; assessed as Least Concern
(LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; lim-
ited data suggest protection from pasture improvement within nature
reserves would be advantageous to the conservation of this species; pro-
tected in Ithala Game Reserve, Suikerbosrand Nature Reserve (South
Africa); Etosha National Park (Namibia).
♀ 2 mm

ONTHOPHAGINI
548 SURICATA 6 (2020)

Proagoderus loricatus
(Klug, 1855)

= Onthophagus collaris Fahraeus, 1857

= Onthophagus harpax Fabricius, 1801, sensu Gestro, 1895
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus loricatus: Not stated; O. collaris: ‘juxta

fluvium Gariep’ [near Orange River, South Africa]; O. harpax sensu
Gestro: ‘Boran Galla, Auata’ [Mount Auata, Boran Galla, Ethiopia].
Taxonomy: Accepted auratus group species, but validation of NE African
synonym would be useful; limited sexual dimorphism (head).
♂ Distribution: Widespread from SE to NE Africa; occurrence now patchy,
mainly centred on game reserves in dry savanna: South Africa, N Bot­
swana, Zimbabwe. Mozambique, Kenya; also reported from Tanzania,
Uganda; records from Somalia, Ethiopia overlap with range of close
relative, P. harpax (Fabricius, 1801).
Locality data (mean ± SD, range): Altitude: 595 ± 367, 92–1 524 m;
annual rainfall: 604 ± 94, 428–900 mm; annual temperature (max. +
min. /2): 21.4 ± 1.8, 15.6–25.9°C (N=51).
Habitat: No conclusive data: in open woodland at Leeuwfontein Nature
Reserve: sand (10), sandy loam (2), stony sandy loam (2); DBRU
collection records: sand (2), sandy loam (5), sandy clay loam (1) in
shrubland (1), open woodland (3), shaded thicket (1).
Food types: At Phalaborwa: strong bias to omnivore dung (pig: 32) com-
pared to herbivore dung (elephant: 2; cattle: 1); similar in Botswana:
pig (21), elephant (3), cattle (2), sheep (3); however, DBRU collection
records support strong influence of monogastric herbivore dung on dis-
tribution pattern: elephant (8), donkey (1); few records on ruminant
herbivore dung: cattle (2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.); at Phalaborwa: frequency much greater in hot, dry con-
2 mm ditions (28) than in warm cloudy conditions following light rainfall (2)
or in warm sunny conditions soon after heavy rainfall (5).
Ecoregions Botswana, Zimbabwe: Zambezian and Mopane Woodlands
(AT0725), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl), Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 1 895 725 km2; despite measured food
type bias, AOO probably much reduced due to range contraction by
monogastric herbivores; 65% of DBRU collection records (11/17) were
from game reserves; in South Africa, most modern records are from two
centres, environs of Kruger National Park and bushveld nature/game
reserves containing zebra; assessed as Least Concern (LC) due to wide
distribution in many reserves.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations, owing
to the inconclusive and conflicting information currently available;
protected in various national parks, including Kruger (South Africa),
Hwange (Zimbabwe), Chobe (Botswana).

2 mm ♀

ONTHOPHAGINI
SURICATA 6 (2020) 549

Proagoderus plato
(Bates, 1888)

No synonyms
Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type locality: As Onthophagus plato: ‘Damaraland’ [inexact, N Namib-

ia – defined at the time of collection as lying between 19–23°S and
14–20°E (Bates 1888)].
Taxonomy: Accepted gibbiramus group species described from a single
male specimen; male specimen cited here as P. plato requires validation
by comparison with the type; female specimens need to be confirmed
as those of P. plato; they show arcuate clypeal margins that differ from ♂
those of P. furciger by lacking a mid-clypeal projection; also see notes on
former synonymy of P. plato on species page for P. furcifer (Boheman,
1860); strong sexual dimorphism (head, prothoracic disc); prominence
of characters varies with body size.
Distribution: Dry savanna from N Namibia to W Zimbabwe; black
line represents inexact origin of a male specimen (Eiseb Omuramba –
Omuramba = a dry watercourse), all other data points and shaded areas
represent female specimens.
Locality data (mean ± SD, range): Altitude: 1 023 ± 111, 945–1 102 m;
annual rainfall: 613 ± 21, 598–628 mm; annual temperature (max.
+ min. /2): 21.6 ± 0.6, 21.2–22.1°C (N=2); Evari and Eiseb similar
except drier (annual rainfall: 385 ± 14, 371–398 mm).
Habitat: No quantitative assessment; one DBRU record for females from
sandy loam in open mopane woodland.
Food types: No quantitative assessment; one DBRU record for females
from elephant dung.
Temporal activity: Presumed to show diurnal flight activity during the
summer rainy season like other Proagoderus species in southern Africa
(Nov., Apr.).
Ecoregions Namibia, Zimbabwe: Zambezian Baikiaea Woodlands
(AT0726: red squares), Angolan Mopane Woodlands (AT0702), Kala- 2 mm

hari Acacia-Baikiaea Woodlands (AT0709).

Assessment rationale: EOO = 235 670 km2 (tentative); species of the
gibbiramus group are usually recorded as rarities from elephant dung;
the few individuals treated here as P. plato may equally represent the
same or a different species; two specimens were collected from within
the area of the inexact type locality, the male and one old record for
a female (Evari River =?upper reaches of Etaka River, NW of Etosha
Pan); assessed as Data Deficient (DD).
Conservation measures: Specimens cited here need to be validated as
P. plato by comparison with the type and by acquisition of further ref-
erence material from Damaraland eastwards to W Zimbabwe; such a
survey is also required to determine the EOO, AOO and ecological
associations before an accurate assessment may be made of conserva-
tion status; possibly protected in Hwange National Park along with
P. furcifer.

ONTHOPHAGINI
550 SURICATA 6 (2020)

Proagoderus quadrituber
(d’Orbigny, 1908)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Onthophagus (Proagoderus) quadrituber: ‘Zambèze:

Chindé...Mozambique: vallée de Pungoué...Sikumba’, lectotype:
Guengère [all central Mozambique].
Taxonomy: Accepted confossus group species; strong sexual dimorphism
(head, prothoracic disc); prominence of characters varies with body
size.
♂ Distribution: Widespread, but very patchy SE African occurrence due
to soil and woody vegetation specialisation: South Africa, Zimbabwe,
Mozambique, Malawi.
Locality data (mean ± SD, range): Altitude: 315 ± 447, 2–1 377 m; an-
nual rainfall: 858 ± 178, 634–1 177 mm; annual temperature (max. +
min. /2): 21.6 ± 2.2, 17.3–25.2°C (N=11).
Habitat: In uMkhuze Game Reserve: strongly biased to clay (2.3) rather
than sandy clay loam (0.0), or sand (0.3); some support from DBRU
collection records on clay (3), sandy clay loam (2), sand (1) in forest
(1), woodland (2), shrubland (1), grassland (2).
Food types: No quantitative assessment; DBRU collection records on
various dung types: human/cattle mix (2), rhinoceros (3), cattle (2).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.); also Aug. in warm coastal KwaZulu-Natal.
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719); also
riverine locality in Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Moist S Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 134 250 km2; AOO probably a small frac-
tion of the EOO reflecting restricted extent of clay; if this species is
associated with dense woody vegetation, clearance would be detrimen-
tal; in South Africa, vegetation units within the known species range
2 mm are 22–26% transformed; however, currently assessed as Least Concern
(LC) owing to protection offered by reserves.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data to better support association with dense
woody vegetation, and a survey of densely vegetated clay patches in SE
Africa to better determine the EOO and AOO; existing records mostly
from protected game reserves, such as Lake Mutirikwe (Zimbabwe),
uMkhuze, Hluhluwe–iMfolozi (South Africa).

2 mm ♀

ONTHOPHAGINI
SURICATA 6 (2020) 551

Proagoderus rangifer
(Klug, 1855)

= Onthophagus bradshawi Péringuey, 1885

Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus rangifer: ‘in den ebenen von Sena und in
Caya häufig’ [common on the plains of Sena and Caia, central Mozam-
bique]; O. bradshawi: ‘Zambezi River’ [southern Africa].
Taxonomy: Accepted ramosicornis group species; N limits of distribution
reported to be in Tanzania, presumably to the S of the range occupied
by the very closely related P. ramosicornis d’Orbigny, 1902 in Kenya and
Tanzania; reconsideration of the differences between these two species ♂
would be useful; strong sexual dimorphism (head, prothoracic disc);
prominence of characters varies with body size; iridescent colour varia-
tion from cupreous to green.
Distribution: Widespread in SE Africa; occurrence now patchy, centred
in dry savanna, mainly in game reserves or areas with sizeable equine
populations: South Africa, NW Botswana, Zimbabwe, Mozambique;
also reported from Namibia, Malawi, Tanzania.
Locality data (mean ± SD, range): Altitude: 586 ± 341, 61–1 478 m;
annual rainfall: 598 ± 139, 317–1 100 mm; annual temperature (max.
+ min. /2): 21.9 ± 1.7, 16.7–25.7°C (N=35).
Habitat: No quantitative assessment; DBRU collection records from sand
(1), sandy loam (6), sandy clay loam (1) in pasture (1), shrubland (2),
open woodland (3).
Food types: At Phalaborwa: much more abundant on omnivore dung
(pig: 41) than herbivore dung (elephant: 2; cattle: 2); DBRU collection
records from dung of elephant (2), rhinoceros (1), donkey (2), cattle
(4), waterbuck (1).
Temporal activity: Diurnal flight activity during the summer rainy season 2 mm

(Oct. to Mar.); at Phalaborwa: frequency much greater in warm sunny

conditions soon after heavy rainfall (33) than in hot, dry conditions
(11) or warm cloudy conditions following light rainfall (2).
Ecoregions Botswana, Zimbabwe: Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725), margins of Southern
Miombo Woodlands (AT0719).
Bioregions South Africa: N Lowveld (SVl), Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 1 303 445 km2; AOO probably much
smaller owing to contraction in the range of indigenous monogastric
herbivores despite quantitative data that suggest a bias to omnivore
dung or collection records that suggest equal numbers from monogas-
tric and ruminant herbivore dung; in South Africa, distribution centred
around and within the Kruger National Park; however, currently as-
sessed as Least Concern (LC) owing to wide distribution and adequate
protection.
Conservation measures: Conservation status difficult to assess owing to
inconclusive and conflicting data; further quantitative data on ecolog-
ical associations are required to support possible effects of range con-
traction of monogastric herbivores and clearance of woody vegetation
with which it may be associated; protected in Kruger National Park
(South Africa).

♀
2 mm

ONTHOPHAGINI
552 SURICATA 6 (2020)

Proagoderus rectefurcatus
(Fairmaire, 1891)

= Onthophagus porrectus Fahraeus, 1857 [pre-occupied name]

= Onthophagus (Proagoderus) rectefurcatus var. metallarius d’Orbigny, 1913
Global: LC

J A S O N D J F M A M J

Type localities: As Onthophagus rectefurcatus: ‘Mrogoro’ [Morogoro, Tan-

zania]; O. porrectus Fahraeus: ‘prope fluvium Limpopo’ [near Limpopo
River]; var. metallarius: Not stated.
Taxonomy: Accepted prostans group species; strong sexual dimorphism
(head, prothoracic disc); prominence of characters varies with body
♂ size; iridescent colour variation from cupreous to green; usually green.
Distribution: Widespread from SE to E Africa; occurrence now patchy,
mainly centred on game reserves in dry savanna: South Africa, Zim-
babwe, Tanzania; also reported from Botswana, Malawi, Zambia, Mo-
zambique; record from Kenya requires validation.
Locality data (mean ± SD, range): Altitude: 630 ± 365, 266–1 247 m;
annual rainfall: 580 ± 80, 435–697 mm; annual temperature (max. +
min. /2): 22.5 ± 1.5, 19.3–25.9°C (N=20).
Habitat: No quantitative assessment; DBRU collections records from
sand (1), sandy loam (5) in shrubland (2), open woodland (2).
Food types: No quantitative assessment; DBRU collections records from
dung of elephant (13), rhinoceros (1), cattle (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Mar.).
Ecoregions Zimbabwe: Centred on Zambezian and Mopane Woodlands
(AT0725); marginal in two adjoining ecoregions.
Bioregions South Africa: N Lowveld (SVl), Mopane (SVmp) (Savanna
Biome).
Assessment rationale: EOO = 847 660 km2; AOO much more range
restricted, probably due to association with dung of large monogastric
2 mm herbivores; 78% of DBRU collection records (14 out of 18) from game
reserves; in South Africa, known only from Kruger National Park;
widespread, but regularly found in dry savanna reserves containing
elephants; therefore, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; however,
continued protection of large monogastric herbivores (elephant, rhi-
noceros) and woodland habitat on finer-grained soils probably essential
for the conservation of this species; protected in several national parks
including Kruger (South Africa), Hwange (Zimbabwe).

♀
2 mm

ONTHOPHAGINI
SURICATA 6 (2020) 553

Proagoderus sapphirinus
(Fahraeus, 1857)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Onthophagus sapphirinus: ‘juxta fluvium Limpopo’ [near

Limpopo River, southern Africa].
Taxonomy: Accepted dives group species, but confusion in literature in-
dicates that revision is required; name possibly invalid as the Limpopo
type specimen is blue and possibly a senior name for the green/blue
Limpopo/E coastal P. aureiceps d’Orbigny, 1902; this Kalahari species
is green with yellow/cupreous elytra, has more extensive and denser
punctation than P. aureiceps, and may have no valid name (see species
page for P. aureiceps d’Orbigny).
Distribution: Centred on continuous S Kalahari deep sands and outliers;
NW South Africa, Botswana, Namibia; S Angola, S Zambia, central
Mozambique; in the E: occurs inland in Kalahari outliers and on E
coast at mouth of the Zambezi Valley; in the W: extends to edge of
Namib Desert; in S Angola and S Zambia: overlaps with blue P. speculi­
collis Quedenfeldt, 1884; in NE South Africa and on SE Mozambique
coast: overlaps with green/blue P. aureiceps d’Orbigny.
Locality data (mean ± SD, range): Altitude: 1 111 ± 187, 0–1 624 m;
annual rainfall: 411 ± 176, 56–1 254 mm; annual temperature (max. +
min. /2): 19.6 ± 1.2, 16.2–24.6°C (N=131).
Habitat: In Gauteng bushveld: extreme sand specialist (381), sandy clay
loam (0) in grassland (151), open woodland (172), shaded thickets (58);
largely supported by DBRU collection records: sand (19), sandy loam
(6) in grassland/pasture (12), scrub/shrubland (7), open woodland (7).
Food types: In Botswana: bias to omnivore dung (pig: 48) rather than
carrion (chicken livers: 14) and herbivore dung (elephant: 12; cattle:
9; sheep: 7); similar in Gauteng bushveld: carrion (54), dung of pig
(87), horse (35), cattle (36), also rotted grass clippings (5) and overripe
banana (8); DBRU collection records reflect common dung types: ele-
phant (3), zebra (1), horse (1), donkey (1), cattle (13), waterbuck (1).
Temporal activity: Diurnal flight activity during the summer rainy season
2 mm
(Oct. to Apr.).
Ecoregions Namibia, Botswana, Zimbabwe: Centred on deep sands in
Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Wood-
lands (AT0709), Zambezian Baikiaea Woodlands (AT0726); marginal
in Namibian Savanna Woodlands (AT1316), Angolan Mopane Wood-
lands (AT0702), Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Centred on E Kalahari Duneveld (SVkd), W
Eastern Kalahari Bushveld (SVk), sand patches in Central Bushveld
(SVcb) (Savanna Biome).
Assessment rationale: EOO = 1 117 105 km2; as a sand specialist, AOO
would be limited to areas and patches of deep sands, which are, nev-
ertheless, extensive across its EOO; much of range coincides with
little-transformed regions used primarily for grazing of domestic live-
stock; data indicate a widespread vegetation and food generalist; there-
fore, assessed as Least Concern (LC).
Conservation measures: It is necessary to ensure that the species name is
valid as it is possible that past errors may have resulted in this taxon being
represented by a type specimen belonging to a different species; protect-
ed in the private Tswalu Kalahari Reserve (South Africa), Central Ka-
lahari Game Reserve (Botswana), Hwange National Park (Zimbabwe).

ONTHOPHAGINI
554 SURICATA 6 (2020)

Proagoderus tersidorsis
(d’Orbigny, 1902)

No synonyms
Global: LC
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type locality: As Onthophagus (Proagoderus) tersidorsis: ‘Natal’ [KwaZu-

lu-Natal, South Africa].
Taxonomy: Accepted confossus group species; strong sexual dimorphism
(head, prothoracic disc); prominence of characters varies with body
size.
♂ Distribution: Centred on moist lowland savanna in NE South Africa and
Eswatini.
Locality data (mean ± SD, range): Altitude: 532 ± 421, 0–1 694 m;
annual rainfall: 697 ± 108, 448–932 mm; annual temperature (max. +
min. /2): 20.7 ± 1.9, 15.1–23.1°C (N=74).
Habitat: In uMkhuze Game Reserve: biased to sandy clay loam (32.3) as
opposed to deep sand (3.4), sand over clay (8.0) or clay (1.0); large-
ly supported by DBRU collection records: sand (3), sandy loam (7),
sandy clay loam (5), clay (3) in grassland/pasture (3), open woodland
(8), shaded thickets (1).
Food types: No quantitative assessment; DBRU collection records on
various dung types: baboon (1), elephant (4), rhinoceros (2), zebra (2),
horse (1), donkey (1), cattle (12), wildebeest (2), impala (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.).
Bioregions South Africa: Centred on Lowveld (SVl), scattered records
from Central Bushveld (SVcb) (Savanna Biome); marginal in three
other bioregions.
Assessment rationale: EOO = 85 690 km2; AOO probably more re-
stricted due to soil type specialisation and possible woody vegetation
2 mm association; the possibility of reduced population density in response to
clearance of woodland needs to be tested; however, as 66% (25 out of
38) of DBRU collection records are from game reserves, this species is
currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tected in uMkhuze and Hluhluwe–iMfolozi game reserves plus Kruger
National Park.

♀
2 mm

ONTHOPHAGINI
556 SURICATA 6 (2020)

TRIBE SCARABAEINI
Latreille, 1802

As Scarabaeïdes; Type genus: Scarabaeus Linnaeus, 1758.

Synonyms:
Pachysomides Ferreira, 1953a; Type genus: Pachysoma Macleay, 1821.
Actinophorini Ádám, 2003; Type genus: Actinophorus Creutzer, 1799 [=Scarabaeus Linnaeus, 1758, senior name].

The tribe Scarabaeini is probably monophyletic. It shows a distribution centred on the winter rainfall, arid
summer rainfall and savanna regions of the Afrotropical, S to E Palaearctic and W Oriental regions. Most
species show either diurnal or nocturnal ball-rolling activity, although some show other specialisations.
Recent generic and subgeneric subdivision of the tribe has been subject to different treatments by different
authors.

Forgie et al. (2005) reduced 11 valid genera to only two, Pachylomera Griffith & Pidgeon, 1831, and
Scarabaeus Linnaeus, 1758. Of the nine genera synonymised with Scarabaeus, some were made full syn-
onyms: Neateuchus Gillet, 1911; Drepanopodus Janssens, 1940b; plus most flightless genera: Mnemati-
um Macleay, 1821; Mnematidium Ritsema, 1888; Neomnematium Janssens, 1938a; Madateuchus Paulian,
1953). Others were reduced to subgeneric status: Pachysoma Macleay, 1821; Sceliages Westwood, 1837b;
Kheper Janssens, 1940b. In addition, Scarabaeolus Balthasar, 1965a, was revalidated at subgeneric level.

Subsequent to 2005, nomenclature was, again, radically revised. Some taxa were revalidated as genera
(Pachysoma; Parateuchus Shipp, 1895e) or raised to generic level (Scarabaeolus) (Forgie et al. 2006; Moretto
2016b). Some taxa were cited as subgenera (Scarabaeus s. str. Linnaeus, 1758), created as new subgenera
(Escarabaeus Zídek & Pokorný, 2011; Pachylosoma Zídek & Pokorný, 2008) or revalidated as subgenera
(Mnematium; Ateuchetus Bedel, 1892b) (Zídek & Pokorný 2008, 2011). Together with Mnematidium and
Kheper, all of these subgenera were, thereafter, raised to generic status by Ziani and Gudenzi (2012), except
for Pachylosoma. Thus, the tribe currently comprises ten genera (Pachylomera, Kheper, Escarabaeus, Scarabae-
us s. str., Ateuchetus, Parateuchus, Scarabaeolus, Mnematium; Mnematidium, Pachysoma) and two subgenera
of Scarabaeus s. str. (Pachylosoma, Sceliages).

In the present work, we have followed the most recent rulings on status for each higher taxon (as of Nov.
2017) with one exception (Sceliages is treated as a genus). Thus, representatives of the tribe in South Africa,
Botswana and Namibia are treated as seven genera. They are listed below in rank order of derivation ac-
cording to Forgie et al. (2005), which suggests Pachylomera is more basal, Pachysoma more derived and the
remainder, essentially, sister genera.
SURICATA 6 (2020) 557

(1) Pachylomera Griffith & Pidgeon, 1831: Genus comprising two S Afrotropical species centred on
deep sands, particularly, the Kalahari and outliers.
(2) Kheper Janssens, 1940b: An Afrotropical and Oriental genus including ten southern African species
with widespread or restricted distributions centred in savanna (6), N upland grassland (1), SW
Kalahari deep sand (1) SW winter rainfall region (1), NW arid late summer rainfall region (1)
(3) Escarabaeus Zídek & Pokorný, 2011: An Afrotropical and Palaearctic genus with two southern Af-
rican species centred in the SW Arid late summer rainfall region or alluvial soils in the N Botswana
Kalahari.
(4) Sceliages Westwood, 1837b: S Afrotropical genus with specialised breeding using millipede body
contents; not yet formally revalidated at generic level; six species in South Africa and Botswana
centred on winter and bimodal rainfall regions (2), Kalahari deep sand (1) or E savanna (3).
(5) Scarabaeolus Balthasar, 1965a: S Afrotropical centred genus comprising 27 South African, Botswa-
nan and Namibian species with widespread or restricted distributions centred in the Namib Desert
(4), winter and bimodal rainfall regions (2), Kalahari deep sands (9), SW Arid late summer rainfall
region (5), E savanna (6), savanna and Karoo (1). However, further species were added by Zídek &
Pokorný (2018) (see Preamble).
(6) Scarabaeus s. str. Linnaeus, 1758: Afrotropical, Palaearctic and W Oriental genus; revision required
as the 31 or 33 South African, Botswanan and Namibian species would probably belong to several
different genera in addition to Scarabaeus s. str. according to the principles of subdivision used by
Ziani and Gudenzi (2012); species with widespread or restricted distributions centred in the Namib
Desert (2), Kalahari deep sands (2), Kalahari and Namib Desert (2), winter and bimodal rainfall
regions (7), Upper Karoo (1), upland grassland (8), savanna (10), E forest (1).
(7) Pachysoma Macleay, 1821: Flightless SW Afrotropical genus comprising 13 species endemic to the
arid W coasts of South Africa and S Namibia.

SCARABAEINI
558 SURICATA 6 (2020)

Genus Escarabaeus Zídek & Pokorný, 2011

Type species and designation: Scarabaeus cristatus Fabricius, 1775, by original designation.
Synonyms: = Mesoscarabaeus Zídek & Pokorný, 2008: pre-occupied name; type species: Scarabaeus
cristatus Fabricius, 1775, by original designation.
Last review: Formally created and reviewed as Mesoscarabaeus (Zídek & Pokorný 2008); renamed
Escarabaeus with updated species list (Zídek & Pokorný 2011); raised to generic status
(Ziani & Gudenzi 2012).

Escarabaeus Zídek & Pokorný, 2011, is a nom nov. for the pre-occupied, subgeneric name of Mesoscarabaeus Zídek
& Pokorný, 2008, which has subsequently been raised to generic level (Ziani & Gudenzi 2012). It comprises eight
confirmed species with combined distributions primarily in arid to dry areas of the Afrotropical, S Palaearctic and
W Oriental regions.

Generic membership of Palaearctic species is supported by a clade in the morphological phylogeny of Barbero et al.
(1998). Only two species from southern Africa are officially designated as members of the genus at present: E. satyrus (Bo-
heman, 1857) and the recently described close relative, E. remii Davis & Deschodt, 2017. These two validated members
of the genus in southern Africa show distributions centred on the arid SW and alluvial soils in N Botswana where habitat
transformation is mostly limited. Therefore, both are assessed as Least Concern (LC).

SCARABAEINI
SURICATA 6 (2020) 559

Escarabaeus remii
Davis & Deschodt, 2017

No synonyms
Global: LC
Endemic: Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Botswana, Okavango, Thamalakane riv.’.

Taxonomy: Accepted species; newly described, smaller-bodied, close rela-
tive of Escarabaeus satyrus (Boheman, 1860).
Distribution: Centred on alluvial soils of the Okavango Delta and Mak-
gadikgadi Pan: N Botswana, NE Namibia.
Locality data (mean ± SD, range): Altitude: 937 ± 60, 871–1 075 m;
annual rainfall: 503 ± 93, 399–647 mm; annual temperature (max. +
min. /2): 22.4 ± 0.2, 22.1–22.7°C (N=9).
Habitat: No quantitative assessment; one DBRU collection record from
grey sand in open mopane woodland.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Flight activity in darkness, primarily during the sum-
mer rainy season (Sept. to Mar.).
Ecoregions Namibia, Botswana: Primarily Zambezian Flooded Grass-
lands (AT0907), Zambezian Halophytics (AT0908) embedded within
other ecoregions (see summary table).
Assessment rationale: EOO = 74 550 km2; occupies a fairly large EOO
in an area with a high degree of protection; unclear if AOO would be
proportionately much smaller as ecologically poorly known; on basis of
wide range and protection, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations and AOO; 5 mm
protected in Makgadikgadi Pans Game Reserve.

SCARABAEINI
560 SURICATA 6 (2020)

Escarabaeus satyrus
(Boheman, 1860)

= Scarabaeus burchelli Gillet, 1911

Global: LC

J A S O N D J F M A M J

Type localities: As Ateuchus satyrus: ‘juxta fluvios Svakop et Nolagi’ [near

Swakop River (W Namibia) and Nolagi, which is an untraced locality,
possibly in Namibia or Botswana]; S. burchelli: ‘tous d’Afrique australe’
[all southern Africa].
Taxonomy: Accepted species; habitus close in appearance to S. deludens
zur Strassen, 1961, but aedeagus differs radically.
Distribution: Arid to dry regions of SW Africa, often on sandy soils;
SW Angola, Namibia, Botswana, South Africa; centred in the arid late
summer rainfall region, also on Kalahari deep sands, present only as a
rarity in the vicinity of Cape Town; record from Somalia considered
an error due to interpretation of untraced type locality of ‘Nolagi’ as a
misspelling of ‘Nogali’, Somalia; citations from Somalia and Ethiopia
presumably errors.
Locality data (mean ± SD, range): Altitude: 1 017 ± 304, 1–2 074 m;
annual rainfall: 230 ± 101, 22–655 mm; annual temperature (max. +
min. /2): 18.2 ± 2.0, 12.9–23.3°C (N=342).
Habitat: No adequate quantitative assessment; in Northern Cape: most
abundant in warm arid Bushmanland (3.3 per sample in 57 of 67 sam-
ples = 85.1%), but less abundant on deep Kalahari sands (1.1 in 64 of
98 = 65.3%), cooler uplands to S of Orange River (0.8 in 95 of 121
= 78.5%), and cool Upper Karoo (0.3 in 38 of 68 = 55.9%); DBRU
collection records mainly from sandy soils: sand (20), sandy loam (4),
sandy clay loam (1), in open vegetation, grassland (8), scrub/shrubland
(12), open woodland (5).
Food types: No quantitative assessment; DBRU collection records from
herbivore dung: cattle (16), horse (1), donkey (1), goat + sheep pellets (1).
Temporal activity: Flight activity during darkness in warmer months of
5 mm
the year across the mid-summer, late summer, winter and bimodal rain-
fall regions (Sept. to May); attracted to light.
Ecoregions Namibia, Botswana: Centred on arid ecoregions; Nama
Karoo (AT1314), N Succulent Karoo (AT1322), Kaokoveld Desert
(AT1310), Namib Desert (AT1315), Namibian Savanna Woodlands
(AT1316), Kalahari Xeric Savanna (AT1309), marginal in three savan-
na ecoregions to NE.
Bioregions South Africa: Centred on Bushmanland (NKb), Upper
Karoo (NKu) (Nama Karoo Biome); also Eastern Kalahari Bushveld
(SVk), Kalahari Duneveld (SVkd) plus W margins of Central Bushveld
(SVcb), Mopane (SVmp) (Savanna Biome); marginal in Succulent
Karoo (SK) and Fynbos (F) biomes.
Assessment rationale: EOO = 1 491 750 km2; widespread across an arid
range used primarily for grazing of domestic livestock and conserva-
tion; high frequency of records in farm rangeland where it is readily
attracted to herbivore dung of farm livestock; assessed as Least Concern
(LC) owing to large, mostly little-transformed EOO.
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; protected in Kga-
lagadi Transfrontier Park (South Africa, Botswana), Namib-Naukluft
and Etosha national parks (Namibia).

SCARABAEINI
SURICATA 6 (2020) 561

Genus Kheper Janssens, 1940b

Type species and designation: Ateuchus aegyptiorum Latreille, 1823, by original designation.
Synonyms: None.
Last review: Entire genus reviewed by Janssens (1940b); three new valid species described subsequent-
ly (Davis 1986; Deschodt et al. 2011; Pokorný & Zídek 2015), one revalidated from
synonymy (Moretto 1998), and one transferred from Scarabaeus (present publication).

Kheper Janssens, 1940b, was described as a full genus. Although it was recently reduced to subgeneric status (Forgie et al.
2005), it has since been revalidated at generic level (Ziani & Gudenzi 2012). Furthermore, the most recently discovered
new species (Pokorný & Zídek 2015) was described under Kheper at generic level. Unlike other members of the tribe
Scarabaeini, some species show metallic colouration.

It currently comprises 25 valid, large-bodied species primarily characteristic of African savannas south of the Sahara (22
species), but also the W Oriental region (3 species: India, Sri Lanka) and margins of the Palaearctic region (Afghanistan).
Exceptionally, of the 22 African species, three are found in arid and winter rainfall regions in the SW.

The genus is divisible into three species groups on the basis of the presence of two, one or no spines on the dorsal surface
of the fore tibia at the base of tibial teeth 3 and/or 2 (Davis 1986). This may or may not parallel the three forms of the
scutellum that have been noted for Kheper: shield-shaped, triangular or sub-triangular (Ziani & Gudenzi 2012).

A total of ten species occur within the study area in southern Africa including one species, Scarabaeus vethi (van Lansberge,
1886), here transferred to Kheper.
Group 1: Two spines on dorsal surface of protibiae; one each at the bases of tibial teeth 2 and 3. Comprises three
species with distributions in all savanna (2 spp.) or E savanna only (1); all showing distributions into the
tropics. Two showing associations with monogastric herbivore dung, one a night-flier.
Group 2: One spine on dorsal surface of protibiae at base of tibial tooth 2. Comprises six diurnal species with
distributions in E savanna (1), Maputaland (1), N Highveld (1), W Cape coast (1), SW Kalahari (1), N
Namibia coast (1).
Group 3: No spines on dorsal surface of protibiae at bases of tibial teeth. Comprises one night-flying species in
savanna with a distribution into the tropics.

Grouping matches behaviour. In Group 1, pheromone may be released by males to attract females to a dropping. Also,
balls intended for breeding are very large as 2–3 broods (or more) are constructed per breeding chamber. Group 2 roll
smaller balls and may construct only a single brood per breeding chamber, which may explain why some species are able to
survive in arid areas. If rolling in pairs, the female sits on the side of the ball rolled by the male. If rolled only by a male, he
releases a pheromone at the burial site to attract a female. Females may breed once or twice a year raising breeding success
by tending the brood during development of immatures. They are able to breed following a single rainfall event (refers
K. nigroaeneus (Boheman, 1857): Edwards 1988; Edwards & Aschenborn 1988).

Seven species are currently assessed as Least Concern (LC) and three are Data Deficient (DD). However, clearance of nat-
ural shrubland in the Western Cape is a cause for concern (K. bonellii (Macleay, 1821)) as well as small range (700 km2) in
a highly transformed area of NE KwaZulu-Natal (K. clericus (Boheman, 1857)). Furthermore, K. zurstrasseni Davis, 1986,
is known by only few museum specimens and has not been recorded since 1936.

SCARABAEINI
562 SURICATA 6 (2020)

Kheper bonellii
(Macleay, 1821)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus bonellii: ‘Cap Bon. Spei’ [Cape of Good

Hope, South Africa].
Taxonomy: Accepted Group 2 species.
Distribution: Restricted to the winter rainfall region, South Africa, where
it is centred on deep sands of the West Coast from the Cape Peninsula
as far as the Richtersveld; a few in adjacent dry uplands.
Locality data (mean ± SD, range): Altitude: 162 ± 226, 0–975 m; annual
rainfall: 223 ± 103, 88–429 mm; annual temperature (max. + min. /2):
17.1 ± 1.0, 15.7–18.5°C (N=36).
Habitat: In Western Cape: quantitative data inconclusive; abundant on
deep sand in shrubland at Modderrivier (89); also in shrubland at Pam-
poenvlei on deep sand (13) or sandy loam (31) and on sandy soils in
rough pasture at Oranjefontein (48); rare in sparse pasture at Groote
Post (2) and Kikuyu pasture at Waylands (0); DBRU collection records
primarily from deep sands (8) in natural scrub/shrubland (7); few re-
cords from sandy loam (2), sandy clay loam (1), or disturbed habitats,
fallow crop fields (2), sparse pasture (1).
Food types: No quantitative assessment; DBRU collection records from
the dung of cattle (10) and horse (2).
Temporal activity: Diurnal flight activity; in the Western Cape: narrow
Aug. to Oct. peak in seasonal activity tailing off into Nov. and Dec.;
tenerals only in Aug. and Sept. suggesting an annual breeding cycle
with seasonal dormancy between spring activity peaks.
Bioregions South Africa: Centred on Namaqualand Sandveld (SKs)
5 mm
(Succulent Karoo Biome); W vegetation units in West Strandveld (FS),
Sand Fynbos (FFd), Sandstone Fynbos (FFs) (Fynbos Biome).
Assessment rationale: EOO = 25 000 km2; little threatened in N part of
range that is used primarily for grazing of livestock, (transformation:
3–10%); assessment of threats from cultivation and urbanisation in
southern range (transformation: 25–50% (FS), 40–55% (FFd), 75%
(FFs)) not possible due to inconclusive quantitative results; currently
assessed as Least Concern (LC) on basis of low transformation in N
of range.
Conservation measures: Assessment of conservation status would be im-
proved by additional quantitative data on ecological associations, par-
ticularly further assessment of threats due to vegetation transformation
in the S; protected in West Coast National Park and the coastal part of
Namaqua National Park; also Cape of Good Hope Nature Reserve and
the private Grotto Bay Nature Reserve where it was frequently recorded
in 1987 when the natural shrubland of the coastal strip was part of the
Farm Modderrivier and was used for grazing cattle.

SCARABAEINI
SURICATA 6 (2020) 563

Kheper clericus
(Boheman, 1857)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Ateuchus clericus: ‘Caffraria interiore’ [probably interior

of SE South Africa].
Taxonomy: Accepted Group 2 species; type collected by J. Wahlberg; la-
belled ‘Natal, Caffraria’.
Distribution: Localised on the narrow coastal plain in the Maputaland
Centre of Endemism, KwaZulu-Natal, South Africa.
Locality data (mean ± SD, range): Altitude: 161 ± 81, 55–273 m; annual
rainfall: 833 ± 85, 738–904 mm; annual temperature (max. + min. /2):
21.2 ± 0.6, 20.5–22.0°C (N=5).
Habitat: No adequate quantitative assessment; in uMkhuze Game Re-
serve, recorded only on clay (4), not on sandy clay loam, duplex soil
(sand on sandy clay loam) or sand; DBRU collection records only from
finer-grained soils (one each on clay, sandy clay loam, sandy loam) in
savanna woodland (4).
Food types: No quantitative assessment; DBRU collection records from
the dung of square-lipped (white) rhinoceros (1) and zebra (1).
Temporal activity: Diurnal flight activity in early summer (Oct. to Dec.).
Bioregions South Africa: Vegetation units: Zululand Lowveld (SVl 23),
Western Maputaland Clay Bushveld (SVl 20) (SE extremity of Lowveld
(SVl), Savanna Biome).
Assessment rationale: EOO = 750 km2; collection records exclusively on
finer-grained soils and monogastric herbivore dung in wooded areas of
game reserves are too limited to draw clear conclusions, therefore, cur-
rently assessed as Data Deficient (DD); however, possible specialisation
and restricted known occurrence within a very small range may warrant 2 mm

allotment of a threat category of at least Vulnerable (VU); although

habitat transformation by cultivation is 26–34% in SVl 20 and SVl 23,
possible effect of woodland loss is unknown.
Conservation measures: Owing to limited information, assessment of
conservation status would be improved by further quantitative data
on soil, dung and vegetation associations both within and outside of
reserves on the coastal plain; protected in uMkhuze and Hluhluwe–
iMfolozi game reserves.

SCARABAEINI
564 SURICATA 6 (2020)

Kheper cupreus
(Castelnau, 1840)

= Ateuchus aeruginosus Klug, 1855

= Ateuchus metallicus Boheman, 1857
= Kheper namibicus Krajcik, 2006
Global: LC (see IUCN Red List – LC as Scarabaeus namibicus)

J A S O N D J F M A M J

Type localities: As Ateuchus cupreus: ‘Caffrerie’ [probably South Africa];

A. aeruginosus: ‘Tette’ [Tete, central Mozambique]. A. metallicus: ‘in
tractus fluvii Limpoponis’ [Limpopo River course, southern Africa];
K. namibicus: ‘NAMIBIA, road C20 10 km S of Gobabis’.
Taxonomy: Accepted Group 3 species; colour variation from black with
iridescent sheen to iridescent cupreous.
Distribution: Centred on hot, dry savanna in southern central Africa: N
South Africa, Botswana, N Namibia, Zimbabwe; also reported from
Mozambique, S Democratic Republic of the Congo (DRC) and S
Tanzania.
Locality data (mean ± SD, range): Altitude: 866 ± 465, 55–1 924 m;
annual rainfall: 487 ± 134, 171–743 mm; annual temperature (max. +
min. /2): 21.0 ± 1.8, 16.8–25.9°C (N=59).
Habitat: No adequate quantitative assessment; DBRU collection records
shared between sand (5), sandy loam (4) and sandy clay loam (1), pri-
marily in shrub/woodland (8), rare in grassland (1).
Food types: At Phalaborwa, attracted to small baits of pig (9) or cattle (9)
dung, but not elephant (0); DBRU collection records from dung of
cattle (9), hook-lipped (black) rhinoceros (1) and wildebeest (1).
Temporal activity: Flight activity in darkness as indicated by tan-coloured
pilosity; mid-summer activity from Nov. to Feb.; at Phalaborwa: more
abundant immediately following substantial rainfall (15) compared to
drier conditions (2); attracted to light.
5 mm
Ecoregions Namibia, Botswana, Zimbabwe: N Namibia Savanna
Woodlands (AT1316), Kalahari Xeric Savanna (AT1309), Kalahari
Acacia-Baikiaea Woodlands (AT0709), Angolan Mopane Woodlands
(AT0702), Zambezian and Mopane Woodlands (AT0725), Southern
African Bushveld (AT0717).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 865 025 km2; widespread on various
soil types in dry savannas of southern central Africa within both game
reserves and farmland as it is readily attracted to the dung of both
omnivores and ruminant herbivores; assessed as Least Concern (LC),
although a possible association with woodland savanna would render
it susceptible to habitat transformation, which stands at 0–58% (SVl),
2–49% (SVcb) or 0–20% (SVmp) across the South African part of its
range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
vegetation associations; protected in various South African nature re-
serves including the Kruger National Park plus Hwange National Park,
Zimbabwe.

SCARABAEINI
SURICATA 6 (2020) 565

Kheper kalaharicus
(Davis, Deschodt & Scholtz, 2011)

No synonyms
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: As Scarabaeus (Kheper) kalaharicus: ‘Kgalagadi Transfrontier

Park’ [Botswana].
Taxonomy: Accepted Group 2 species.
Distribution: Probably restricted to the SW Kalahari dune field strad-
dling the borders of Namibia, Botswana and South Africa.
Locality data (mean ± SD, range): Altitude: 953 ± 85, 908–1 124 m;
annual rainfall: 209 ± 19, 188–239 mm; annual temperature (max. +
min. /2): 20.4 ± 0.8, 19.0–21.1°C (N=6).
Habitat: No quantitative data, but all specimens (45) sampled from
shrub/grassland on arid deep sands of a dune field, an association that
may account for the elongate body shape.
Food types: In Botswana: not attracted to carrion, but shows a very strong
bias to pig dung with much lower attraction to other dung types: carri-
on (0), pig (31), elephant (1), cattle (3), sheep dung (8).
Temporal activity: Diel periodicity not known, but presumably diurnal
flight as indicated by black pilosity; active during the late summer rainy
season (Feb., Mar.).
Ecoregions Botswana: Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Kalahari Duneveld (SKvd) (Savanna Biome).
Assessment rationale: EOO = 6 000 km2; undoubtedly underestimated;
probably faces few threats in a dune field that is used primarily for
grazing domestic livestock or is under conservation; assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
5 mm
proved by quantitative data to confirm habitat associations; protected
in Kgalagadi Transfrontier Park (Botswana and South Africa).

SCARABAEINI
566 SURICATA 6 (2020)

Kheper lamarcki
(Macleay, 1821)

= Ateuchus infernalis Klug, 1855

Global: LC
J A S O N D J F M A M J

Type localities: As Scarabaeus lamarckii: ‘Senegallia’ [Senegal]; A. infer-

nalis: ‘In Mossambique verbreitet, eben so, gewöhnlich kleiner, in Port
Natal’ [evenly distributed in Mozambique, usually localised in Durban,
South Africa].
Taxonomy: Accepted Group 1 species.
Distribution: Widespread, but localised on deep sands in hot, dry sa-
vanna from W to S central and southern Africa; mega-Kalahari, E
coast and intervening sand outliers: South Africa, Botswana, Namibia,
Zimbabwe, Mozambique, Angola, Democratic Republic of the Congo
(DRC) and Senegal; also reported from Guinea (Conakry).
Locality data (mean ± SD, range): Altitude: 849 ± 454, 0–1 628 m; an-
nual rainfall: 548 ± 218, 220–1 275 mm; annual temperature (max. +
min. /2): 20.7 ± 1.6, 16.1–25.2°C (N=153).
Habitat: In Gauteng: exclusively on coarse-grained soils, deep sand (261),
sandy clay loam (0) with a bias to wooded savanna, grassland (48),
open woodland (170), shaded thickets (43); in uMkhuze Game Re-
serve: equally strong bias to coarse-grained soils, sand (8.8), duplex soils
(sand over clay) (2.0), sandy clay loam (0.0), clay (0.5); largely support-
ed by DBRU collection records: sand (29), sandy loam (14), sandy clay
loam (1) in grassland (12) or shrub/woodland (31).
Food types: In both Gauteng and Botswana: strong bias to omnivore
dung; in Gauteng bushveld: carrion (0), pig (69), horse (4), cattle dung
(9); in Botswana: carrion (115), pig (3 336), elephant (996), cattle
(576), sheep dung (484); DBRU collection records on various dung
types: cattle (35), buffalo (1), wildebeest (1), elephant (5), square-
5 mm lipped (white) rhinoceros (1), horse (1), waterbuck (1).
Temporal activity: Diurnal flight activity in both sunny and cloudy con-
ditions as long as temperatures are sufficiently high; in Gauteng bush-
veld: active in the summer rainy season (Oct. to Apr.), also May, Aug.
and Sept. on the warmer E coastline.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Deep sands:
Kalahari Xeric Savanna (AT1322), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Southern African Bushveld (AT0717), Zambezian and Mopane
Woodlands (AT0725), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Deep sands in E Kalahari Duneveld (SVkd),
Eastern Kalahari Bushveld (SVk), Central Bushveld (SVcb), Mopane
(SVmp), Lowland (SVl) (Savanna Biome); also Indian Ocean Coastal
Belt Biome (CB).
Assessment rationale: EOO = 2 173 065 km2; widespread on deep sands
in both farm rangeland and conserved areas as it is readily attracted to
a range of dung types despite a bias to omnivore dung; vegetation bias
suggests that clearance of woodland would result in a fourfold reduc-
tion in population density; in South Africa, transformation low in W
vegetation units of SVk (0–2%), but often higher in SVcb, SVmp and
SVl (0–58%); assessed as Least Concern (LC) on basis of high frequen-
cy across a wide range.
Conservation measures: None recommended; protected in various re-
serves containing deep sands, including uMkhuze Game Reserve, Kru-
ger National Park and iSimangaliso Wetland Park (World Heritage
Site) (South Africa), Chobe National Park (Botswana) and Maputo
Special Reserve (Mozambique).

SCARABAEINI
SURICATA 6 (2020) 567

Kheper nigroaeneus
(Boheman, 1857)

= Scarabaeus usurpator Péringuey, 1901

Global: LC (see IUCN Red List – LC as Scarabaeus)

J A S O N D J F M A M J

Type localities: As Ateuchus nigroaeneus: ‘Caffraria interiore’ [interior of

SE South Africa]; S. usurpator: ‘Manica, Mozambique’.
Taxonomy: Accepted Group 2 species; colour variation from black to iri-
descent cupreous; green iridescence in some black individuals.
Distribution: Centred on hot, dry to moist savanna in E southern Africa:
N South Africa, E Botswana, Zimbabwe, Mozambique; Namibian re-
cord requires validation.
Locality data (mean ± SD, range): Altitude: 840 ± 433, 1–1 670 m; an-
nual rainfall: 672 ± 158, 306–1 415 mm; annual temperature (max. +
min. /2): 20.1 ± 2.0, 14.2–25.2°C (N=207).
Habitat: In uMkhuze Game Reserve: no clear soil type bias, sand (31.7),
duplex soil (sand over clay) (27.6), sandy clay loam (46.6), clay (8.8);
no quantitative assessment of vegetation association; DBRU collection
records from sand (10), sandy loam (19), sandy clay loam (13), clay (0)
in pasture/grassland (13), shrub/woodland (25) and thickets/forest (2).
Food types: At Phalaborwa: clear bias to omnivore dung (pig, 736; cattle,
406; elephant, 144), although prominent on fresh elephant droppings
in the Kruger National Park; DBRU collection records from dead
cricket (1) and various dung types: baboon (1), elephant (2), rhinoceros
(3), donkey (1), cattle (32), buffalo (2), wildebeest (2), waterbuck (1).
Temporal activity: Diurnal flight activity, especially in the cooler early morn-
ing during the summer rainy season (mainly Oct. to Apr.); ball intended
for breeding is rolled by male on his own or with female clinging to the
side; if alone, male releases pheromone at burial site to attract female; fe-
males tend a single brood in a chamber in the soil from early in the seasonal 5 mm
activity period; tenerals appear in Mar. (Edwards 1988); females may or
may not breed again and tend a brood during winter; at Phalaborwa: much
greater activity in warm conditions after heavy rainfall (981) compared
to hot, dry (190) or warm cloudy conditions after light rainfall (110).
Ecoregions Botswana, Zimbabwe: Centred on Southern Miombo
Woodlands (AT0719), Southern African Bushveld (AT0717), marginal
in moister parts of Zambezian and Mopane Woodlands (AT0725), E
Kalahari Xeric Savanna (AT1309), E Kalahari Acacia-Baikiaea Wood-
lands (AT0709).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); also sporadically in East Kalahari
Bushveld (SVkb) (Savanna Biome) plus Sub-Escarpment Grassland
(Gs) (Grassland Biome).
Assessment rationale: EOO = 662 125 km2; widely distributed in both
farm rangeland and conserved areas; recorded on a range of soil and
dung types in both grassland and the open woodland that dominates its
range; transformation in South Africa by cultivation and urbanisation
(0–58% (SVl), 2–49% (SVcb) or 0–20% (SVmp)) would presumably
be detrimental; however, assessed as Least Concern (LC) because of
its abundance in the partially protected NE lowlands of South Africa.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and vegetation associations; pro-
tected in various reserves including, uMkhuze and Hluhluwe–iMfolozi
game reserves and Kruger National Park (South Africa).

SCARABAEINI
568 SURICATA 6 (2020)

Kheper prodigiosus
(Erichson, 1843)

= Scarabaeus pubiventris van Lansberge, 1874b

Global: LC

J A S O N D J F M A M J

Type localities: As Ateuchus prodigiosus: Not stated; S. pubiventris: ‘Mo-

zambique’.
Taxonomy: Accepted Group 1 species.
Distribution: Widespread in hot, dry savanna; primarily on deep sands,
of the summer rainfall region in southern central Africa: N South Af-
rica, Botswana, Namibia, Angola, Zimbabwe, Zambia, Mozambique;
Democratic Republic of the Congo (DRC), Tanzania records require
validation.
Locality data (mean ± SD, range): Altitude: 891 ± 352, 102–1 730 m;
annual rainfall: 484 ± 136, 171–813 mm; annual temperature (max. +
min. /2): 21.3 ± 1.7, 16.8–25.9°C (N=76).
Habitat: No adequate quantitative assessment; on deep sands in Chobe
National Park, Botswana: more abundant in grassland with trees (113)
than open woodland (35) or dense woodland (33); DBRU collection
records biased to sand (8), also sandy loam (3) in grassland (4) and
open woodland (6).
Food types: In Botswana: bias to monogastric herbivore and omnivore
dung: carrion (2), pig (62), elephant (143), cattle (26), sheep dung
(18); DBRU collection records from dung of elephant (6), cattle (6),
warthog (1).
Temporal activity: Flight activity in darkness during the summer rainy
season (Nov. to May) as suggested by tan pilosity; attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe: Centred on sandy soils in
Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Wood-
lands (AT0709), Zambezian and Mopane Woodlands (AT0725), also
5 mm
Angolan Mopane Woodlands (AT0702) and margins of three other
ecoregions.
Bioregions South Africa: Sandy areas of Eastern Kalahari Bushveld
(SVk), Central Bushveld (SVcb), Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 2 045 625 km2 (possibly underestimated);
widespread although AOO presumably influenced by soil, vegetation
and dung type bias; abundant on deep sands in NW Botswana where
elephants are abundant (249), rare in the arid SW where elephants are
absent (2); many collection records from game reserves, therefore, as-
sessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on soil and vegetation associations as obser-
vations suggest these would be influential to AOO; protected in various
national parks where elephants are conserved, including Kruger (South
Africa), Chobe (Botswana), Etosha (Namibia), Hwange (Zimbabwe).

SCARABAEINI
SURICATA 6 (2020) 569

Kheper subaeneus
(Harold, 1869b)

= Ateuchus lamarcki Macleay, 1821, sensu Castelnau, 1840

= Scarabaeus metallicus Boheman, 1857, sensu Péringuey, 1901
= Scarabaeus subaeneus var. orientalis Gillet, 1907
= Scarabaeus subaeneus var. atratus Gillet, 1911
= Kheper subaeneus angolensis Janssens, 1940b
Global: LC (see IUCN Red List – LC as Scarabaeus)

J A S O N D J F M A M J

Type localities: As Scarabaeus subaeneus: ‘Senegal’; as A. lamarckii sensu

Castelnau: ‘Senegal’; S. metallicus sensu Péringuey: ‘Zambesia (Victoria
Falls)’ [Zimbabwe]; S. subaeneus orientalis: ‘Mozambique’; S. subaeneus
atratus: ‘l’Oubangui et de la Chari’ [Central African Republic (CAR)];
K. subaeneus angolensis: ‘Angola’.
Taxonomy: Accepted Group 1 species; colour variation from black to
iridescent cupreous.
Distribution: Widespread in hot, dry to moist savannas in southern,
southern central, E central and W Africa; possibly localised by soil
specialisation; validated for South Africa, Botswana, Zimbabwe, Mo-
zambique, Angola; synonyms in CAR, Senegal; also reported from
Democratic Republic of the Congo (DRC), Côte d’Ivoire, Ghana,
Mauritania; possibly also Uganda, South Sudan.
Locality data (mean ± SD, range): Altitude: 750 ± 464, 55–1 635 m;
annual rainfall: 641 ± 137, 317–1 177 mm; annual temperature (max.
+ min. /2): 20.8 ± 2.4, 16.3–25.9°C (N=88).
Habitat: No adequate quantitative assessment; DBRU collection records
indicate a bias to finer-grained soils: sand (3), sandy loam (9), sandy
clay loam (6) with similar numbers of records from pasture/grassland
(7) and open woodland (7), also in forest (1). 5 mm

Food types: At Phalaborwa: clear bias to dung of omnivores (pig, 60;

elephant, 3; cattle, 4), but prominent on fresh elephant and rhinoceros
droppings in game reserves; this bias is supported by DBRU collection
records for elephant (16), rhinoceros (3) and buffalo dung (1) in re-
serves compared to cattle (7) and donkey dung (2) in farmland.
Temporal activity: Diurnal flight activity during the summer rainy season (Oct. to Apr.); at Phalaborwa: activity much greater
in warm conditions after substantial rainfall (58) compared to hot, dry conditions (6) or warm cloudy conditions after light
rainfall (0).
Ecoregions Botswana, Zimbabwe: E Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern African Bushveld (AT0717),
Zambezian and Mopane Woodlands (AT0725), Zambezian Baikiaea Woodlands (AT0726), Southern Miombo Woodlands
(AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 1 257 850 km2 (probably underestimated); extremely widespread, but may be localised on
finer-grained soils with greater population density in reserves that protect elephants and rhinoceros although this is not
supported near the South African Wildlife College in the Kruger National Park (3.0) compared to adjoining farm rangeland
(17.8); assessed as Least Concern (LC) owing to widespread occurrence and large number of records from reserves (61 out
of 103).
Conservation measures: Assessment of conservation status would be improved by quantitative data on habitat and dung type
associations both inside and outside of reserves that protect large monogastric herbivores on finer-grained soils; protected
especially in the Kruger National Park and game reserves including Hluhluwe–iMfolozi, Roodeplaat (South Africa) and Lake
Mutirikwe (Zimbabwe).

SCARABAEINI
570 SURICATA 6 (2020)

Kheper vethi
(van Lansberge, 1886)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: As Scarabaeus vethi: ‘Benguela’ [Angola].

Taxonomy: Accepted species, but here transferred to Kheper from
Scarabaeus; classified in Group 2; identification of Kaokoveld speci-
mens as K. vethi requires validation.
Distribution: Arid to dry coastal plain: N Namibia, SW Angola.
Locality data (mean ± SD, range): Altitude: 519 ± 224, 361–677 m;
annual rainfall: 193 ± 37, 167–219 mm; annual temperature (max. +
min. /2): 16.9 ± 0.3, 16.7–17.2°C (N=2).
Habitat: Not known.
Food types: Not known.
Temporal activity: Diel flight periodicity unknown, but presumably diur-
nal flight based on black pilosity; seasonal activity recorded during the
late summer rainy season (Dec. to Feb.).
Ecoregions Namibia: On the border of the arid Kaokoveld Desert
(AT1310) and N Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 35 400 km2 (estimated); rarity may or
may not be a collection artefact; two N Namibian localities lie outside
the boundary of the Skeleton Coast Park and the Tora Conservancy
although they do lie in an arid area used primarily for grazing of live-
stock; currently assessed as Data Deficient (DD).
Conservation measures: Current data is insufficient to determine con-
servation status; a survey needs to be conducted in N Namibia and
particularly Iona National Park northwards to Benguela, SW Angola;
not currently known from any officially protected area.

5 mm

SCARABAEINI
SURICATA 6 (2020) 571

Kheper zurstrasseni
Davis, 1986

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Transvaal, Pretoria’ [Gauteng, South Africa].

Taxonomy: Accepted Group 2 species.
Distribution: Locality records only from uplands in North-West, Gauteng
and Limpopo provinces, South Africa.
Locality data (mean ± SD, range): Altitude: 1 206 ± 196, 1 013–1 356 m;
annual rainfall: 626 ± 43, 588–672 mm; annual temperature (max. +
min. /2): 18.2 ± 1.0, 17.3–19.2°C (N=3).
Habitat: Unknown.
Food types: Unknown.
Temporal activity: Diel flight periodicity unknown, but presumably di-
urnal based on black pilosity; a single temporal record in the summer
rainy season (Feb.).
Bioregions South Africa: Difficult to assess; localities coincide with Mesic
Highveld Grassland (Gm) (Grassland Biome) or lie close to patches
of the same grassland type isolated in the adjoining Central Bushveld
(SVcb) (Savanna Biome).
Assessment rationale: EOO = 1 450 km2, probably underestimated,
based on only three known localities; very poorly represented by only
six specimens in the type series despite a distribution centred in an
intensively sampled region; available data suggest it is highly threatened
as the last known dated collection was made in 1936; qualifies for En-
dangered (EN) status on basis of EOO < 5 000 km2 at < five localities,
but is currently assessed as Data Deficient (DD).
Conservation measures: In view of its rarity in collections, K. zurstrasseni
could be a relative specialist; however, as nothing is known of its habits,
a range of habitat and dung types needs to be tested for its occurrence, 5 mm

starting around the three known, but relatively inexact localities, none
of the known localities coincide with protected areas.

SCARABAEINI
572 SURICATA 6 (2020)

Genus Pachylomera Griffith & Pidgeon, 1831

Type species and designation: Scarabaeus femoralis Kirby, 1828, by monotypy.
Synonym: = Pachylomerus Bertolini, 1849: Type species: Scarabaeus femoralis Kirby, 1828, by
monotypy.
Last review: Genus: Ziani & Branco (2011); Species: Janssens (1940b).

Although this genus has been long accepted as comprising only two species described in the 19th century, there has been
much confusion over the validity of two available generic names (Pachylomerus or Pachylomera) and their authorship (Kir-
by 1828; Hope 1832 (unpublished); Bertolini 1849). A recent consideration of the complicated published history (Ziani
& Branco 2011) has shown that none of the previous combinations is correct and that the valid name and authorship is
Pachylomera Griffith & Pidgeon, 1831.

The genus shows a widespread EOO in the summer rainfall region of southern and S central Africa. Its AOO is much
more restricted as it is centred on deep sands, particularly the mega-Kalahari from South Africa northwestwards to W
Democratic Republic of the Congo (DRC), as well as deep sand outliers and coastal sands in the E from South Africa,
reportedly as far N as coastal Tanzania. As both species are either centred on or restricted to Kalahari sands, neither is
seriously threatened by habitat transformation although one species also occurs in areas of greater transformation.

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Pachylomera femoralis
(Kirby, 1828)

= Ateuchus (Pachylomera) horridus Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type localities: As Scarabaeus femoralis: ‘Soudan’ [presumably an error;

never since found in Sudan]; A. horridus: ‘juxta fluvium Limpopo’ [near
Limpopo River, southern Africa].
Taxonomy: Accepted species; see discussion of generic name on genus
page.
Distribution: Centred on moister deep sands of the mega-Kalahari and
its outliers from South Africa across Botswana, Namibia, Zimbabwe,
SW Zambia, Angola to W Democratic Republic of the Congo (DRC);
also deep sands on E seaboard from South Africa across Mozambique
to Tanzania.
Locality data (mean ± SD, range): Altitude: 796 ± 481, 0–1 696 m;
annual rainfall: 583 ± 217, 194–1 270 mm; annual temperature (max.
+ min. /2): 20.9 ± 1.7, 16.6–25.5°C (N=197).
Habitat: In Gauteng bushveld: recorded exclusively on deep sand (70),
absent from sandy clay loam (0); equally abundant in grassland (30)
and open woodland (37) rare in shaded thickets (3); in uMkhuze Game
Reserve: similar soil bias: deep sand (208.7), duplex soil (sand over
clay) (22.2), sandy clay loam (0.3), clay (0.4); supported by DBRU
collection records: sand (17), sandy loam (6), sandy clay loam (1) in
grassland (9), shrubland (10), open woodland (8), forest (1).
Food types: In Botswana: bias to omnivore dung (pig: 5 131), but also
abundant on carrion (1 149) and herbivore dung (elephant: 759, cattle:
607, sheep: 1 303); similar bias in Gauteng bushveld: (carrion: 3, pig:
15, horse: 2, cattle dung: 4); DBRU records from various dung types:
elephant (3), horse (1), donkey (2), cattle (16), buffalo (1), wildebeest
(1).
Temporal activity: Diurnal flight activity, even under cloudy conditions
as long as temperatures are sufficiently high; in Gauteng bushveld: sea-
sonal activity Oct. to Apr.; at least Aug. to May on warmer E coastline.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Deep
5 mm
sands, particularly in Kalahari Xeric Savanna (AT1309), Kalahari
Acacia-Baikiaea Woodlands (AT0709), Angolan Mopane Woodlands
(AT0702), Zambezian Baikiaea Woodlands (AT0726), Southern
African Bushveld (AT0717), Zambezian and Mopane Woodlands
(AT0725), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Deep sands of Eastern Kalahari Bushveld (SVk),
Central Bushveld (Svcb), Mopane (SVmp), Lowveld (SVl) (Savanna
Biome); also Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 2 663 660 km2; AOO restricted by the
distribution of deep sands, but widespread in farm rangeland although
vegetation units in NE part of South African range are subject to
0–58% transformation; circa 20% of southern African range protected
in national parks and game reserves, therefore, assessed as Least Con-
cern (LC).
Conservation measures: None recommended; widespread on deep sands
in farm rangeland and protected in various national parks and game
reserves, especially Chobe, Khutse (Botswana); Kruger, Hluhluwe–
iMfolozi, Tswalu Kalahari (South Africa).

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Pachylomera opaca
van Lansberge, 1874b

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Lac N’gami’ [Lake Ngami, Botswana].

Taxonomy: Accepted species; see discussion of generic name on genus
page.
Distribution: Centred on the arid southwest Kalahari deep sands; rare
occurrence in mainland Kalahari sands: N Botswana, SW Zambia;
however, also occurs in cooler deep sand outliers of Gauteng and Lim-
popo provinces, South Africa.
Locality data (mean ± SD, range): Altitude: 1 076 ± 151, 776–1 401 m;
annual rainfall: 275 ± 91, 172–636 mm; annual temperature (max. +
min. /2): 19.3 ± 0.9, 17.6–21.2°C (N=78).
Habitat: No quantitative assessment; DBRU collection records exclusively
on deep sand (19) in pasture/grassland (9), scrub (3) or open shrubland
(8); most records from arid localities, but occurs as a rarity at moister
localities supporting open grassland.
Food types: In Botswana: attracted primarily to omnivore dung (pig: 28)
less commonly to herbivore dung (elephant: 3, cattle: 1, sheep: 6);
DBRU collection records on dung of cattle (12), horse (2), donkey (2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.).
Ecoregions Botswana: Scattered localities in Kalahari Acacia-Baikiaea
Woodlands (AT0709).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), Kalahari
Duneveld (SVkd) (Savanna Biome); N limits of Bushmanland (NKb),
Upper Karoo (NKu) (Nama Karoo Biome); sand outliers in Central
Bushveld (SVcb) (Savanna Biome).
Assessment rationale: Full EOO = 800 000 km2; SW Kalahari EOO =
120 000 km2; widespread, although AOO is limited by distribution of
5 mm deep sand and open vegetation; not abundant on dung of domestic her-
bivore livestock, but frequently recorded on farms in the SW Kalahari
in an area subject to only 0–2% transformation (W SVk); assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by by quantitative data on habitat associations; circa 10% of
the SW Kalahari range protected within Kgalagadi Transfrontier Park
(South Africa and Botswana); also protected in Telperion Nature Re-
serve (Gauteng in South Africa).

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SURICATA 6 (2020) 575

Genus Pachysoma Macleay, 1821

Type species and designation: Pachysoma hippocrates Macleay, 1821, by original designation.
Synonyms: = Irrorhotides Shipp, 1896a: Type species: Irrorhotides fryi Shipp, 1896a, by monotypy.
= Neopachysoma Ferreira, 1953a. Type species: Pachysoma denticolle Péringuey, 1888,
by original designation.
Last review: Entire genus reviewed by Harrison et al. (2003) as Scarabaeus (Pachysoma) with the
inclusion of distributional and ecological data.

The long-established, flightless genus Pachysoma Macleay, 1821, differs strongly in appearance to winged members of
the tribe Scarabaeini. However, it was recently reclassified as a subgenus of Scarabaeus Linnaeus, 1758, on the basis that
the two genera cannot be differentiated morphologically after excluding characters associated with flightlessness, which
include constriction between the reduced thorax and the anteriorly rounded abdomen that results from the loss of the
humeral angles on the elytra. Subsequent combined morphological and molecular phylogenetic analysis resulted in the
revalidation of Pachysoma as a full genus (Forgie et al. 2006).

Pachysoma is considered to be the most derived of the genera and subgenera in the tribe Scarabaeini. It currently comprises
13 valid species described between 1789 and 2003. Both molecular and morphological phylogenetic analyses support
division of the genus into three species groups associated with particular soil type and climatic factors.

The genus is endemic to the dry to arid area below the escarpment along the West Coast from Cape Town in South Africa
to Walvis Bay in central Namibia. Species have diversified primarily along a north–south axis between deep coastal sands
of South Africa and southern Namibia (eight species), dune fields of the central Namib Desert in Namibia (three species),
and the pre-Namib gravel plains in Namibia (two species).

Pachysoma is considered to have a Pliocene origin (2.9 MY) (Sole et al. 2005) associated with increased upwelling of
the cold Benguela current and reduced rainfall followed by subsequent Plio–Pleistocene speciation events in response to
cyclic climatic changes. The three resulting species groups show northern, central or mostly southern bias in ranges, but
exceptionly, distributions of two species of the southern group are disjunct in the north centre of the generic range. Spe-
ciation has probably been influenced by vicariance across river barriers, environmental tracking during climatic change,
or morphological adaptation to different landscapes and soil types from gravel to sands, either compacted or loose dune
sands.

Behavioural differences to winged species of Scarabaeini support the treatment of Pachysoma as a full genus. All species
are flightless and show diurnal foraging activity walking on the soil surface. Loss of flight and lower metabolic rate were
possibly driven by increasing aridity on the West Coast and reduced availability of food. Whilst collection of dung
pellets and dry portions of other dung types apparently dominates in five species, three species are characterised by
mixed collection of dung pellets and vegetation detritus, and four species by a strong bias to collection of vegetation
detritus, which may be more generally available than dung (Harrison et al. 2003). Several species have been observed to
grip forage between the hind legs and drag it forwards to pre-constructed tunnels, which may be more energy efficient
than the ball-rolling behaviour shown by their closest relatives and/or better suited to collection of detritus. It is unclear
how the forage is treated for feeding purposes, but the triturating mouthparts of adults are able to ingest larger particles
than most dung beetles. Pachysoma shows unique nesting behaviour within the subfamily Scarabaeinae, since larvae
are free living in a cup-shaped mass of particles within a chamber in the soil. With the loss of brood ball construction
that typifies ball-rolling dung beetles, larvae lack the characteristic dorsal hump found in species that develop within
a dung ball.

Owing to relative aridity across the generic range, land usage is primarily grazing in natural vegetation and conservation,
particularly in Namibia where almost the entire range of Pachysoma is conserved within national parks. In South Africa,
only small parts of the ranges occupied by Pachysoma species coincide with national parks, or other reserves, and several
species are comparatively unprotected. In the arid north, there is extensive grazing by livestock with threats from diamond
mining along the coastline. In the moister south, cultivation, invasive exotic shrubs and urban development constitute

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the greatest threat, particularly to the southernmost species, P. aesculapius Macleay, 1821. This species and two others
with very restricted ranges are considered to deserve IUCN threat categories from Vulnerable (VU) to Endangered (EN).
Further protection of these and four other species found in South Africa is desirable considering the restricted ranges and
low vagility shown by the genus.

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SURICATA 6 (2020) 577

Pachysoma aesculapius
(Olivier, 1789)

= Ateuchus barbatus Thunberg, 1818

= Pachysoma validum Boheman, 1857
Global: VU (see IUCN Red List – VU)
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus aesculapius: ‘Cap’ [Cape of Good Hope,

South Africa], lectotype: ‘no locality’; A. barbatus: Not stated; P. val-
idum: ‘Caffraria’ [presumably South Africa].
Taxonomy: Accepted species; N–S cline in decreasing body size, but
morphological variation does not support separation at species level; N
individuals bear limited elytral indument.
Distribution: Restricted to a small area of coastal and inland sands in SW
South Africa from the south bank of the Olifants River (N) to the Cape
Flats, E of Cape Town (S).
Locality data (mean ± SD, range): Altitude: 134 ± 137, 0–502 m; annual
rainfall: 259 ± 84, 178–466 mm; annual temperature (max. + min. /2):
17.0 ± 0.8, 15.8–18.4°C (N=26).
Habitat: No quantitative assessment; open shrubland on firm deep sands
of coastal hummocks, river banks and vegetated dunes.
Food types: No quantitative assessment; seven excavated burrows con-
tained only dry dung pellets.
Temporal activity: Flightless; diel activity diurnal; in SW Cape: seasonal
activity in the moist spring (Aug. to Oct.) and dry summer (Nov. to
Feb.); in mid-summer (Dec.): forages for short periods in the morning
(07:00–09:00) and late afternoon (16:00–18:00) when radiant heat is
lower than at midday.
Bioregions South Africa: Vegetation units of West Strandveld (FS 1, FS
5, FS 6), Sand Fynbos (FFd 2–5) (Fynbos Biome).
Assessment rationale: EOO = 10 750 km2; AOO = ± 2 000 km2; S part of
range threatened by urban expansion around Cape Town (FS 6: 40%;
FFd 5: 80% transformed) with no records from the City (Salt River),
nearby Cape Flats and Somerset West since 1882–1886; much of the
centre and N of range is threatened by conversion to arable lands and 5 mm
pasturage (FS 1, 5: 25–35%, FFd 2–4: 40–55% transformed); assessed
as Vulnerable (VU).
Conservation measures: A quantitative survey to determine the exact
AOO is advisable since distribution is probably localised on firmer
deep sands and negatively influenced by loss of indigenous shrubland,
through both agricultural and urban development, as well as invasive
exotic vegetation; occurs in the provincial Rocherpan Nature Reserve
and the coastal part of the Farm Modderrivier (60 km N of Cape
Town), now run as the private Grotto Bay Nature Reserve; not found
on looser coastal dune sands on the Farm Geelbek, now part of West
Coast National Park; known occurrence within reserves is limited, so
surveys should include the inland portion of West Coast National Park
and the municipal protected coastal strip along False Bay.

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Pachysoma bennigseni
Felsche, 1907

= Pachysoma granulatum Ferreira, 1953b

Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: P. bennigseni: ‘Oranjefluss’ [presumably lower reaches of

Orange River, SW Namibia]; P. granulatum: ‘Daberas Dunes’ [SW
Namibia].
Taxonomy: Accepted species; Orange River is a geographical barrier
between N and S populations although morphological differences are
insufficient to justify separation at species level (N: larger, granular
prothoracic disc; S: smaller, smooth prothoracic disc).
Distribution: Restricted to the arid N Succulent Karoo from the Hol-
gat River mouth (NW coast, South Africa) to Spencer Bay, just N
of Luderitz (Namibia); occurs no more than 60 km inland from the
coastline in the S and central parts of its range; in the N, inland extent
of the range narrows to the coastline along the S edge of the Namib
Desert dune field.
Locality data (mean ± SD, range): Altitude: 287 ± 282, 11–991 m; an-
nual rainfall: 41 ± 20, 2–75 mm; annual temperature (max. + min. /2):
15.5 ± 0.8, 14.0–16.9°C (N=24).
Habitat: No quantitative assessment; sparsely vegetated coastal sand dunes
and sand flats that vary in nature from unstable to compacted sand.
Food types: No quantitative assessment; a single nest contained both
dung pellets and detritus.
Temporal activity: Flightless; diel activity diurnal; recorded sporadically
across all seasons.
Ecoregions Namibia: Centred on Succulent Karoo (AT1322).
Bioregions South Africa: Centred on two vegetation units: Richtersveld
Coastal Duneveld (SKs 1), Richtersveld Red Duneveld (SKs 5) (Nam-
aqualand Sandveld (SKs), Succulent Karoo Biome).
Assessment rationale: EOO = 8 500 km2; known from 24 sites across a
5 mm restricted but largely undisturbed and mostly protected range; there-
fore, assessed as Least Concern (LC).
Conservation measures: Assessement of conservation status would be
improved by quantitative data on ecological associations; ± 90% of
range protected by the Namib-Naukluft and Sperrgebiet national parks
(Namibia) with the latter having restricted access since 1908; no pro-
tection in South Africa; coastal strip in S and centre of range heavily
disturbed by open cast mining; unknown if there is any coastal bias in
population density.

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SURICATA 6 (2020) 579

Pachysoma denticolle
Péringuey, 1888

= Neopachysoma penrithae Zunino, 1977

Global: LC (see IUCN Red List – LC)
Endemic: Namibia

J A S O N D J F M A M J

Type localities: P. denticolle: ‘Walfish Bay’ [Walvis Bay, central Namibia],

lectotype: ‘Walfish Bay, Namibia’; N. penrithae: ‘Lüderitz’ [SW Namib-
ia].
Taxonomy: Accepted species; as variation in morphology shows no con-
sistent geographical trend, the various populations are considered to
represent a single species without subspeciation; variation comprises
granular elytral interstriae in some S individuals and a N–S cline from
red to melanic elytra.
Distribution: Restricted to the Namib Desert dune field, SW Namibia;
dunes extend inland from the coastline to the gravel plains or base of
the W escarpment from just N of Luderitz to just S of Walvis Bay.
Locality data (mean ± SD, range): Altitude: 589 ± 352, 0–1 145 m; an-
nual rainfall: 47 ± 31, 0–89 mm; annual temperature (max. + min. /2):
16.1 ± 0.6, 14.9–17.2°C (N=58).
Habitat: No quantitative assessment; forages amongst sparse vegetation at
the bases of dunes in the cooler early morning, which is consistent with
other observations made in dune streets and on semi-stable sand; not
observed on sand dunes.
Food types: No quantitative assessment; forage picked up by hind legs
and held under the abdomen whilst using front and middle legs for
walking; collects mainly oryx and hare dung pellets, also droppings of
chameleons, mice, dead insects and vegetable matter.
Temporal activity: Flightless; diel activity diurnal; no recorded seasonality
in activity; collection records throughout the year.
Ecoregions Namibia: Centred on Namib Desert (AT1315).
Assessment rationale: EOO = 40 000 km2; known from 68 sites across a
2 mm
somewhat restricted, but largely undisturbed, mostly protected range;
no known threats, therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; however,
± 95% of the range protected in the Namib-Naukluft National Park
(Namibia).

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Pachysoma endroedyi
(Harrison, Scholtz & Chown, 2003)

No synonyms
Global: EN (see IUCN Red List – EN)
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus (Pachysoma) endroedyi: ‘Namaqualand, Kom-

mandokraal Farm’ [South Africa].
Taxonomy: Accepted species; smaller than the related P. hippocrates Ma-
cleay, 1821, from which it differs only by shorter metatarsal claws and
elytral indument.
Distribution: Localised to an extremely small known range just north of
the Olifants River, possibly, a little underestimated.
Locality data (mean ± SD, range): Altitude: 87 ± 34, 40–124 m; annual
rainfall: 157 ± 2, 154–159 mm; annual temperature (max. + min. /2):
18.3 ± 0.1, 18.1–18.5°C (N=6).
Habitat: No quantitative assessment; open shrubland on the firm sand of
coastal hillocks and dunes.
Food types: No quantitative assessment; of seven excavated nests, one con-
tained dung pellets and six a mixture of pellets and vegetation detritus.
Temporal activity: Flightless; diel activity diurnal; recorded in the moist
spring (Aug., Sept.) and dry summer (Dec., Jan.).
Bioregions South Africa: Margins of the Succulent Karoo (SK) and Fyn-
bos (F) Biomes in only two vegetation units: Namaqualand Strandveld
(SKs 7), Namaqualand Sand Fynbos (FFd 1).
Assessment rationale: EOO = 130 km2; recorded from 12 sites across
an extremely restricted known range with little official protection, al-
though there are some small private reserves; the range is only 2% (FFd
1) to 10% (SKs 7) transformed, but is extensively grazed by livestock
on farms; as no part of the small range is known to occur within a
protected area, assessed as Endangered (EN).
5 mm Conservation measures: A quantitative survey is required to determine
its ecological associations, AOO and population density within the re-
stricted area of remaining suitable dune habitat to assess extinction risk;
effects of recreational development around Strandfontein and Koeke-
naap require investigation. Not currently known from any protected
area.

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SURICATA 6 (2020) 581

Pachysoma fitzsimonsi
Ferreira, 1953a

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Namtib, 70 km NW of Aus’ [farm in SW Namibia].

Taxonomy: Accepted species.
Distribution: Centred on pre-Namib gravel plains, which border the
inland margin of the Namib Desert dune field at the base of the W
escarpment, SW Namibia.
Locality data (mean ± SD, range): Altitude: 1 010 ± 255, 598–1 488 m;
annual rainfall: 83 ± 26, 46–146 mm; annual temperature (max. + min.
/2): 15.6 ± 0.4, 13.3–16.0°C (N=13).
Habitat: No quantitative assessment; arid, sparsely grassed, gravel plains
or gravelly sands, with or without scattered shrubs.
Food types: No quantitative assessment; predicted to be dung pellets on
the basis of morphological characters and supported by excavations of
nests in a sparse-grass pasture grazed by sheep on the farm Weissen-
born; also observations of tunnels clustered around concentrations of
dung pellets that were being collected by P. fitzsimonsi in Namibrand
Nature Reserve.
Temporal activity: Flightless; diurnal activity in the dry winter/spring
(June to Oct.) and late summer (Feb., Mar.).
Ecoregions Namibia: Namib Desert (AT1315), marginal in Namibian
Savanna Woodlands (AT1316).
Assessment rationale: EOO = 8 250 km2; known from 16 sites across
a somewhat restricted range, but would be supported by the dung of
both wild herbivores and domestic livestock; assessed as Least Concern
(LC) owing to the degree of protection in reserves. 5 mm
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; however,
± 70% of the species range is protected within the Namib-Naukluft
National Park and the private Namibrand Nature Reserve (Namibia);
the remainder occurs on farms in the pre-Namib used for the grazing
of livestock.

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582 SURICATA 6 (2020)

Pachysoma gariepinum
Ferreira, 1953a

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: ‘Namaqualand: Holgat’ [dry riverbed, NW South Africa].

Taxonomy: Accepted species; presence, relative prominence, or absence
of a mesepisternal protuberance considered to represent a N–S cline in
morphological variation within a single species.
Distribution: Distributed N and S of the Orange River [Gariep]; S range
limits at the Buffels River (NW South Africa); northernmost locality at
Agub Mountain (SW Namibia); present in coastal regions but 80% of
records centred 20–35 km inland in South Africa; occurs up to 100 km
inland in Namibia.
Locality data (mean ± SD, range): Altitude: 397 ± 273, 2.5–1 137 m;
annual rainfall: 57 ± 17, 22–99 mm; annual temperature (max. + min.
/2): 15.6 ± 0.8, 14.0–17.4°C (N=79).
Habitat: No quantitative assessment; centred on firm, consolidated deep
sands of river banks, inland flats and at the base of rocky outcrops;
found on dunes to the N of the Orange River, but not to the S.
Food types: Limited quantitative assessment; pre-constructs a tunnel then
forages for food; of 17 excavated nests, nine contained dung pellets,
three vegetation detritus and seven a mixture of the two; dry oryx
(gemsbok) dung pellets are collected one at a time by dragging them
forwards gripped between the hind legs; will also use dry horse dung
from which it breaks a piece using the front legs and clypeus then drags
it as for pellets.
Temporal activity: Flightless; diel activity diurnal; no recorded seasonality
in activity; collection records throughout most of the year (Apr. to Jan.).
Ecoregions Namibia: Centred on Succulent Karoo (AT1322).
Bioregions South Africa: Centred on arid sands in the S Namib Desert
(Desert Biome plus Succulent Karoo Biome: Richtersveld (SKr), N
Namaqualand Sandveld (SKs)).
5 mm Assessment rationale: EOO = 17 650 km2; known from 92 sites across a
somewhat restricted range; South African range mostly coincides with
farmland, but Namibian range largely undisturbed and protected; as-
sessed as Least Concern (LC) owing to protection in Namibia.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations; in Namibia: ±
90% of the range is protected within the Sperrgebiet National Park; the
entire South African part of the range is unprotected with the most arid
vegetation units considered in need of conservation due to sensitivity to
degradation through grazing by livestock although possible effects on
population density of P. gariepinum have not been tested.

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SURICATA 6 (2020) 583

Pachysoma glentoni
(Harrison, Scholtz & Chown, 2003)

No synonyms
Global: VU (see IUCN Red List – VU)
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus (Pachysoma) glentoni: ‘Nortier Farm, SW

Cape, South Africa’.
Taxonomy: Accepted species; very close to P. hippocrates Macleay, 1821,
from which it is externally indistinguishable.
Distribution: Very restricted range extending along the coastal belt from
just N of Lamberts Bay to just S of the Olifants River along whose
banks the distribution extends inland almost to Clanwilliam.
Locality data (mean ± SD, range): Altitude: 136 ± 91, 39–342 m; annual
rainfall: 191 ± 15, 171–214 mm; annual temperature (max. + min. /2):
17.9 ± 0.3, 17.5–18.3°C (N=9).
Habitat: No quantitative assessment; open shrubland on firm sand of river
banks (Olifants, Groot Sandleegte) or coastal hummocks; not recorded
in the vicinity of large dunes.
Food types: No quantitative assessment; grips detritus between hind legs
and drags it to a pre-constructed burrow; one male observed in typical
head-down, hind-legs-extended, pheromone-release stance, which may
be used to attract females to the burrow entrance.
Temporal activity: Flightless; diurnal activity; recorded in the moist
spring (July to Sept.) and dry summer (Nov., Dec.).
Bioregions South Africa: Occurs at the N edge of the Fynbos Biome in
single vegetation units of Sand Fynbos (FFd) and Western Strandveld
(FS): Leipoldtville Sand Fynbos (FFd 2), Lamberts Bay Strandveld (FS
1).
Assessment rationale: EOO = 700 km2; known from only 12 sites that 5 mm

are centred on two vegetation units lacking official protection; however,

there are some small private reserves; the range is 25% (FS 1) to 55%
(FFd 2) transformed by cultivation and grazing by domestic livestock;
assessed as Vulnerable (VU) owing to very small, largely unprotected,
partly transformed range.
Conservation measures: A quantitative survey is required to determine
the ecological associations and the AOO, which may reflect the amount
of untransformed suitable habitat remaining within its range; the as-
sessment should determine how much of the AOO coincides with pri-
vate reserves, land grazed by livestock or land threatened by clearance
for cultivation.

SCARABAEINI
584 SURICATA 6 (2020)

Pachysoma hippocrates
Macleay, 1821

= Pachysoma macleayi Castelnau, 1840

= Pachysoma hessei Ferreira, 1953a
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type localities: P. hippocrates: ‘Cap Bonae Spei’ [Cape of Good Hope,

South Africa]; P. macleayi: ‘Cap’; P. hessei: ‘Wallekraal, Namaqualand’
[farm in W South Africa].
Taxonomy: Accepted species; morphological and molecular variation
along a N–S cline insufficient for division at species level either be-
tween four populations separated by major rivers or between S (typical
P. hippocrates) and N populations (P. hessei) in which the inner protibial
margin is elbowed and serrated S of the Olifants River with this char-
acter absent to the N.
Distribution: Restricted to arid W coastal strip, South Africa, from Blou-
bergstrand, N outskirts of Cape Town, to Port Nolloth in N Namaqua-
land; occupies a very narrow coastal strip in the N.
Locality data (mean ± SD, range): Altitude: 96 ± 77, 0–232 m; annual
rainfall: 166 ± 78, 63–368 mm; annual temperature (max. + min. /2):
17.1 ± 1.1, 14.5–18.6°C (N=64).
Habitat: No quantitative assessment; soft to firm deep sand of vegetated
coastal hummocks and hillocks, the periphery of dune systems, as well
as river beds and banks. Found in open shrubland, but not in adjacent
sparse-grass pasture cleared of shrubs (farm Geelbek, now part of West
Coast National Park).
Food types: No quantitative assessment; of 36 excavated nests, contents
comprised vegetation detritus (25), dung pellets (5) or a mixture of the
two (6).
5 mm Temporal activity: Flightless, diel activity diurnal; seasonal activity in the
moist spring (Aug. to Oct.) and dry summer (Nov. to Mar.).
Bioregions South Africa: Namaqualand Sandveld (SKs) (Succulent
Karoo Biome), vegetation units of Western Strandveld (FS) and Sand
Fynbos (FFd) (Fynbos Biome).
Assessment rationale: EOO = 8 500 km2; only ± 10% of range protect-
ed; in the S: range threatened by urban development and agricultural
interests (25–35% transformed); in the N, most unprotected areas little
transformed (2–10%); currently assessed as Least Concern (LC) on
basis of little transformation in the N.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; in the S: re-
corded in the 1990s in West Coast National Park (gazetted 1998); also
occurs in the Groen and Spoeg rivers area of Namaqua National Park
(gazetted 2008); probably not seriously threatened over much of the
range, but further protection desirable in both N and S, particularly as
the EOO is under 20 000 km2.

SCARABAEINI
SURICATA 6 (2020) 585

Pachysoma rodriguesi
(Ferreira, 1953a)

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type locality: As Neopachysoma rodriguesi: ‘Namtib’ [farm, SW Namibia].

Taxonomy: Accepted species; some between-individual variation in the
size of the protuberance on the frons.
Distribution: Restricted to Namib Desert dune field, SW Namibia; dunes
extend inland from the coastline to the gravel plains or base of W es-
carpment from just N of Luderitz to just S of Walvis Bay.
Locality data (mean ± SD, range): Altitude: 555 ± 215, 2–1 145 m; an-
nual rainfall: 40 ± 20, 0–89 mm; annual temperature (max. + min. /2):
16.2 ± 0.4, 15.6–17.0°C (N=44).
Habitat: No quantitative assessment; extreme psammophilous conditions
on sparsely vegetated sand dunes as suggested by long legs with broad-
ened or strongly setose tibiae for increased traction.
Food types: No quantitative assessment; collects dung pellets of hare and
oryx (gemsbok).
Temporal activity: Flightless; diel activity diurnal; active in autumn, win-
ter and spring (Mar. to Oct.), few records in the hot summer (only
Jan.).
Ecoregions Namibia: Namib Desert (AT1315).
Assessment rationale: EOO = 24 000 km2; known from 48 sites across a
somewhat restricted, but largely undisturbed, mostly protected range;
no known threats, therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; however,
± 95% of the range protected in the Namib-Naukluft National Park
(Namibia).
10 mm

SCARABAEINI
586 SURICATA 6 (2020)

Pachysoma rotundigena
Felsche, 1907

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Sinclair, SWA’ [farm, SW Namibia].

Taxonomy: Accepted species.
Distribution: Restricted to the inland edge of the Namib Desert dune
field, which extends inland from the coastline to the gravel plains or
base of the W escarpment from just N of Luderitz to just S of Walvis
Bay, SW Namibia.
Locality data (mean ± SD, range): Altitude: 1 021 ± 205, 665–1 319 m;
annual rainfall: 89 ± 21, 45–124 mm; annual temperature (max. + min.
/2): 16.1 ± 0.7, 14.9–17.5°C (N=18).
Habitat: No quantitative assessment; vegetated dunes at the inland mar-
gins of the dune field.
Food types: Unknown, but predicted to be a detritus feeder on the basis
of morphology and habitat.
Temporal activity: Flightless; diel activity diurnal; no recorded seasonality
in activity; active throughout most of the year.
Ecoregions Namibia: Namib Desert (AT1315), marginal in Namibian
Savanna Woodlands (AT1316).
Assessment rationale: EOO = 9 500 km2; known from 19 sites across a
restricted, but largely undisturbed range; no known threats, therefore,
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; although ± 50%
of the range is protected along the E margin of the Namib-Naukluft
National Park and in the private Namibrand Nature Reserve (Namib-
ia), a survey of its status on dunes in adjacent farmland would be useful.

5 mm

SCARABAEINI
SURICATA 6 (2020) 587

Pachysoma schinzi
Fairmaire, 1888

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Namaqua-Land’ [probably Great Namaqualand, southern

Namibia, which is the origin of most known individuals]; the collector
(Dr Hans Schinz) sampled ‘d’Owambo, de Damara et de Namaqua’.
Taxonomy: Accepted species; marked sexual dimorphism in the morphol-
ogy of the head; male has two large bifid clypeal projections that are
reduced in the female to clypeal teeth, but prominence varies with body ♂
size.
Distribution: Restricted to upland plains of Huib-Hoch Plateau near Aus,
SW Namibia, along W escarpment edge; full EOO uncertain.
Locality data (mean ± SD, range): Altitude: 1 397 ± 134, 1 176–1 533 m;
annual rainfall: 89 ± 20, 71–131 mm; annual temperature (max. + min.
/2): 14.0 ± 0.5, 13.3–14.9°C (N=9).
Habitat: No quantitative assessment; sparse grassland and open scrubland
on gravel plains straddling the boundary between two ecoregions.
Food types: No quantitative assessment; recorded in farmland grazed by
cattle, sheep and horses, but has been reported to feed on vegetation
detritus.
Temporal activity: Flightless; diel activity diurnal; active in spring (Sept.
to Nov.) and late summer (Feb.); possibly a collecting artefact, possibly
related to hot mid-summer temperatures.
Ecoregions Namibia: Margins of Nama Karoo (AT1314) and Namib
Desert (AT1315).
Assessment rationale: EOO = 1 400 km2; known from only 11 sites across 5 mm
a very restricted known range; most specimens recorded close to Aus;
one further locality is interpreted as the nearby Tiras Plains having been
wrongly cited as Tinkasflache; would probably qualify for an IUCN
threat category of at least Vulnerable (VU) on the basis of small range
in an unprotected region; however, currently assessed as Data Deficient
(DD) owing to limited supporting information.
Conservation measures: Assessment of conservation status would be im-
proved by a quantitative survey to determine the full EOO, AOO and
ecological associations; currently known only from unprotected farm-
land used for the grazing of livestock where only 16% of the known
reference material has been recorded since a large serial collection was
made in 1950 (158 specimens; Harrison et al. 2003); low numbers
may be an artefact related to low collecting effort or they may represent
population decline in response to habitat modification resulting from
farming operations.

5 mm

SCARABAEINI
588 SURICATA 6 (2020)

Pachysoma striatum
Castelnau, 1840

= Pachysoma marginatum Péringuey, 1888

= Irrorhotides fryi Shipp, 1896a
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: P. striatum: ‘Cape’ [Cape of Good Hope, South Africa];

P. marginatum: ‘Port Nolloth, Namaqualand’; I. fryi: ‘Port Nolloth’
[both Port Nolloth, NW South Africa].
Taxonomy: Accepted species; differences in body size, elytral sculpture
and relative development of elytral indument are considered to repre-
sent intraspecific morphological variation.
Distribution: Centred from three to 14 km from the coastline along W
coastal zone, South Africa, between Holgat River in the N and Olifants
River in the S.
Locality data (mean ± SD, range): Altitude: 113 ± 88, 0–609 m; annual
rainfall: 114 ± 26, 52–159 mm; annual temperature (max. + min. /2):
17.3 ± 1.0, 14.5–18.5°C (N=85).
Habitat: No quantitative assessment; firm, consolidated deep sands on
river banks, vegetated coastal hummocks, hillocks and dunes.
Food types: No quantitative assessment; forages for dung pellets (rodent,
sheep) and portions of ostrich dung; of 28 excavated nests, 26 con-
tained dry dung pellets and only two a mixture of pellets and vegetation
detritus.
Temporal activity: Flightless; diel activity diurnal; seasonal activity re-
corded from autumn through winter and spring to early summer (May
to Dec.).
Bioregions South Africa: Centred on Namaqualand Sandveld (SKs) (Suc-
culent Karoo Biome) with southern limits at the northern tip of Sand
Fynbos (FFd) (Fynbos Biome).
Assessment rationale: EOO = 6 000 km2; known from 150 sites across
5 mm a somewhat restricted range of which a small part has received official
protection since 2008; range subject to only 2–10% transformation,
primarily through coastal mining, although there is extensive sheep
grazing in SKs 7 and FFd 1; currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; ± 5% of the
range is protected in the Groen and Spoeg river section of Namaqua
National Park; of the remainder, the N range lies largely within the
De Beers diamond concession and is subject to restricted access; the
S range occurs on farms used primarily for the grazing of livestock;
nevertheless, further protection would be desirable.

SCARABAEINI
SURICATA 6 (2020) 589

Pachysoma valeflorae
Ferreira, 1953a

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘10m W of Haalenberg, Gt Namaqualand’ [coastal moun-

tain range in SW Namibia].
Taxonomy: Provisionally accepted species, but very close to P. schinzi
Fairmaire, 1888; marked sexual dimorphism in the morphology of the
head; male has two large clypeal projections that are reduced in the
female to clypeal teeth. ♂
Distribution: All but one of the specimens was recorded from around the
coastal mountains of the Haalenberg and Rotkop, but the true extent of
the range is uncertain; the suggestion of an E–W cline in morphological
variation from Aus (P. schinzi) to the coast (P. valeflorae) is not currently
supported since the species distributions are disjunct other than for one
joint record from Tinkasflache, a clearly suspect and likely erroneous
citation that has been interpreted as the Tiras Plains near Aus.
Locality data (mean ± SD, range): Altitude: 388 ± 98, 295–490 m; an-
nual rainfall: 30 ± 4, 26–33 mm; annual temperature (max. + min. /2):
15.2 ± 0.1, 15.2–15.3°C (N=3).
Habitat: Unknown; possibly sand dunes.
Food types: Unknown.
Diel activity: Flightless; diel activity assumed diurnal like all other
Pachysoma species; active in spring (Sept., Oct.), but seasonality, if any,
unknown.
Ecoregions Namibia: Succulent Karoo (AT1322).
Assessment rationale: EOO = 200 km2 (validated range); known by
eight specimens from four sites (three validated) across an extremely
restricted, but largely undisturbed known range; no known threats, but
a poorly recorded species possibly because of low collecting intensity;
assessed as Data Deficient (DD).
Conservation measures: A study is required to resolve questions on spe-
cies status; assuming this is supported, a quantitative survey is required 5 mm

to assess conservation status; this should determine the full EOO, AOO
and ecological associations; although known by very few specimens, no
conservation actions are currently required as the entire known range is
protected at the extreme N limits of the Succulent Karoo in the Sper-
rgebiet National Park.

SCARABAEINI
590 SURICATA 6 (2020)

Genus Scarabaeolus Balthasar, 1965a

Type species and designation: Scarabaeus (Scarabaeolus) laevifrons Fairmaire, 1884a, by original designation.
Synonyms: None.
Last review: Subgeneric status of Scarabaeolus (Balthasar 1965a) not recognised in a review (zur
Strassen 1967) nor a checklist (Zídek & Pokorný 2004) of the genus, Scarabaeus Lin-
naeus, 1758; new species added to subgenus with synonymies (Deschodt et al. 2015b);
raised to generic status (Moretto 2016b); new species and species list of Scarabaeolus,
again, treated as a subgenus (Zídek & Pokorný 2018).

Scarabaeolus Balthasar, 1965a, was created as a subgenus of Scarabaeus Linnaeus, 1758, using a combination of characters.
These include the absence of an overhanging elytral edge; two widely separated, parallel, lateral carinae on the elytra; small
size (length: 10–14 mm); elytra tapering posteriorly, and the presence of two terminal mesotibial spurs (one vestigeal)
as opposed to one. However, the subgenus was not recognised as a valid taxon in some subsequent taxonomic papers
since both Scarabaeus s. str. and some Scarabaeolus possess a single mesotibial terminal spur, including the type species of
Scarabaeolus. Nevertheless, following years of controversy, Scarabaeolus has now been raised to generic level on the basis of
possessing an entire upper lateral elytral carena compared to an incomplete carena in other taxa (Moretto 2016b). There-
fore, Scarabaeolus is treated here as a full genus.

When this book was completed at the end of 2017, Scarabaeolus comprised 33 valid species found in the Afrotropical
region, although one NE African flightless taxon may be misplaced (S. scholtzi Mostert & Holm, 1982). Subsequently,
further species were added by Zídek & Pokorný (2018) who, again, treated this taxon as a subgenus of Scarabaeus (see
comments in Preamble). Most species show southern African centres of distribution although there are three exceptions.
Two species are centred in NE Africa (Somalia), including the type species, S. laevifrons. A further newly described species,
S. davisianus Moretto, 2016b, is centred in the Sahel (W and N central Africa). As a close, tropical, W African relative has
an incomplete upper carena laterally on each elytron, it has recently (2016) been removed from Scarabaeolus and assigned
to the revalidated genus Parateuchus Shipp, 1895e, as P. palemo (Olivier, 1789).

Both morphological and molecular phylogenies (Forgie et al. 2005, 2006) separate three Scarabaeolus species into a clade
that is sister to those of other taxa in the tribe Scarabaeini. Behavioural differences to Kheper and Scarabaeus s. str. have
been noted in pairs of Scarabaeolus bohemani (Harold, 1869b) where a female follows a ball rolled by the male as opposed
to clinging to the ball rolled by the male. Available quantitative data suggest a bias to carrion (3), carrion + omnivore dung
(3) or omnivore dung (3) with fewer found equally on omnivore and ruminant herbivore dung (2).

Of the 30 southern African species, 27 are found south of 15°S. Distribution patterns are primarily centred on deep sands
although there are exceptions (S. bohemani, S. megaparvulus (Davis & Deschodt, 2015b)). Predominantly diurnal activity
in hot, relatively dry systems may favour relatively smaller body size than in species of Kheper and Scarabaeus s. str. Further-
more, peak activity occurs especially after cooling rainfall events. Only two night-flying species are known (S. canaliculatus
(Fairmaire, 1888); S. flavicornis (Boheman, 1860)).

Many species show distribution patterns centred on deep Kalahari sands (2) with bias to the arid SW (3), moister N (1),
SE edge bordering the Highveld of South Africa (1) or additional occurrence in the pre-Namib (2) southwards to the SW
coast (1). Other species have a distinct SW distributional bias in the SW Arid late summer rainfall region (1) with a bias
to the S (4), or they are centred along the SW coast in the Western and Northern Cape (2), S Namib (2) or N Namib (2).
A number of species show an E distributional bias in savanna bordering the Soutpansberg (2) or on the S Mozambique
Coastal Plain (4). Exceptionally, S. bohemani shows an E, S and W circum-Kalahari distribution in the pre-Namib, Karoo
and Savanna.

Threats to most of the 27 most southerly species are probably low as their distributions are centred on or partly extend
into little-transformed or protected regions. Conservation status has been assessed as Least Concern (LC: 20 spp.) or Data
Deficient (DD: 7 spp.).

SCARABAEINI
SURICATA 6 (2020) 591

Scarabaeolus afronitidus
Davis, Stals & Deschodt, 2015

= Scarabaeus (Scarabaeolus) nitidus Davis & Deschodt, 2015b

Global: DD
Endemic: Botswana

J A S O N D J F M A M J

Type locality: S. afronitidus, nom. nov. for S. (S.) nitidus Davis & Deschodt,
2015b (pre-occupied name): ‘Kutse Game Reserve’ [Khutse Game Re-
serve, S central Botswana].
Taxonomy: Accepted species.
Distribution: Known from three localities in the SW Kalahari, SW Bot­
swana.
Locality data (mean ± SD, range): Altitude: 1 004 ± 26, 987–1 035 m;
annual rainfall: 347 ± 56, 283–386 mm; annual temperature (max. +
min. /2): 20.5 ± 0.1, 20.4–20.6°C (N=3).
Habitat: No adequate quantitative assessment; in Botswana: sampled in
low numbers (12) on deep sand in grassland with scattered shrubs.
Food types: In Botswana: sampled primarily from omnivore dung: pig
(10), rather than herbivore dung: elephant (1), sheep (1).
Temporal activity: Diel flight periodicity unknown, but presumably di-
urnal on the basis of black pilosity; recorded during the summer rainy
season (Jan., Feb.).
Ecoregions Botswana: Kalahari Xeric Savanna (AT1309).
Assessment rationale: EOO = 9 250 km2 (presumably underestimated);
limited data suggest an arid, Kalahari sand specialist on omnivore
dung, but further information is required to fully support the findings;
therefore, assessed as Data Deficient (DD).
Conservation measures: Although all known records are from game re-
serves (Botswana: Khutse, Mabuasehube), this species remains poorly
known; for a comprehensive assessment of conservation status, a quan- 2 mm
titative survey of the SW Kalahari region is required to determine the
full EOO and AOO, and to provide support for the currently limited
data on ecological associations.

SCARABAEINI
592 SURICATA 6 (2020)

Scarabaeolus anderseni
(Waterhouse, 1890)

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus anderseni: cited as ‘Lake Nyassa’ [Lake Ma-

lawi], but type labelled ‘Lake Ngami’; the collector, ‘Andersson’ (not
‘Andersen’) sampled only from Damaraland, Ovamboland and Lake
Ngami, and never went near Lake Nyassa.
Taxonomy: Accepted species; NW specimens show a green sheen becom-
ing melanic to the SW.
Distribution: Known distribution centred on the SW Kalahari with ad-
ditional records from the edge of the Namib Desert in Namibia, N
Botswana, and deep sand outliers in Limpopo Province, South Africa;
citations from Malawi and Zimbabwe require validation.
Locality data (mean ± SD, range): Altitude: 1 070 ± 164, 714–1 730 m;
annual rainfall: 321 ± 138, 56–672 mm; annual temperature (max. +
min. /2): 19.8 ± 1.3, 16.6–22.4°C (N=55).
Habitat: No adequate quantitative assessment; in Northern Cape: mostly
restricted to SW Kalahari study sites (16) as opposed to Nama Karoo
(1); recorded on deep sand (13) and sandy loam (4); essentially a deep
sand specialist in open shrubland or grassland.
Food types: In Botswana: very strong bias to carrion (chicken livers: 322),
attraction to dung much weaker (pig: 41, elephant: 5, cattle: 7, sheep:
15); one DBRU collection record of this species making a ball from
the congealing blood of a dead snake supports carrion specialist habits.
Temporal activity: Diurnal flight activity during the late summer rainy
season (Dec. to Mar.).
Ecoregions Namibia, Botswana: Centred on Kalahari Xeric Savanna
(AT1309); also margins of three adjoining ecoregions.
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd) and the
2 mm
deep sands that dominate the west of the Eastern Kalahari Bushveld
(SVk) (Savanna Biome).
Assessment rationale: EOO = 627 900 km2; AOO restricted to an ex-
tensive area of dry savanna on deep sands that is used primarily for the
grazing of domestic livestock; in South Africa, there is little transfor-
mation in the two Kalahari Bioregions, particularly in the west; about
15% of the EOO coincides with reserves; currently assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations; protected
within the boundaries of game reserves in Botswana and South Africa,
including the Central Kalahari Game Reserve and Kgalagadi Transfron-
tier Park.

SCARABAEINI
SURICATA 6 (2020) 593

Scarabaeolus andreaei
(zur Strassen, 1963)

= Scarabaeus xavieri Ferreira, 1968

Global: LC (see IUCN Red List – DD for Scarabaeus xavieri)

J A S O N D J F M A M J

Type localities: As Scarabaeus andreaei: ‘Inhambane, Mozambique’;

S. xavieri: ‘Província de Moçambique: Inharrime’ [Mozambique].
Taxonomy: Accepted species with S. xavieri a recent synonymy.
Distribution: Known from coastal sands of S Mozambique and an inland
sand outlier up the Limpopo River Valley (Nyandu Sandveld), which is
the only known locality in South Africa.
Locality data (mean ± SD, range): Altitude: 83 ± 171, 0–429 m; annual
rainfall: 835 ± 197, 479–1 084 mm; annual temperature (max. + min.
/2): 23.6 ± 0.8, 22.9–25.2°C (N=6).
Habitat: No quantitative assessment; only recorded from deep sands;
sampled in an open area within shrubland on coastal dune sand near
Maçiene, Mozambique.
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Diurnal flight activity during the summer rainy season
(Dec., Jan.).
Ecoregions Mozambique: Southern Zanzibar-Inhambane Coastal Forest
Mosaic (AT0128) (not shown on map).
Bioregions South Africa: Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 34 050 km2; AOO probably limited by
distribution of deep sands; protected in South Africa, but status in Mo-
zambique uncertain; currently recorded as Least Concern (LC) on the
IUCN Red List but essentially Data Deficient (DD).
Conservation measures: None required in South Africa where the extreme
W edge of its range is protected in the N of the Kruger National Park;
however, a survey is required to determine its full EOO, AOO and 2 mm

ecological associations on the Mozambique Coastal Plain, particularly

the population sizes on deep sand patches compared to finer-grained
soil types and the possible effects on conservation status of land usage
along the coastline.

SCARABAEINI
594 SURICATA 6 (2020)

Scarabaeolus bohemani
(Harold, 1868b)

= Ateuchus cicatricosus Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type locality: S. bohemani, nom. nov. for A. cicatricosus: ‘Caffraria tota’

[inexact, ?all of SE or southern Africa].
Taxonomy: Accepted species, but part of a species complex that includes
the Sahel species, S. davisanus Moretto, 2016b, type locality ‘St LOUIS
Senég.’ [Senegal, W Africa], and S. obsoletepunctatus Balthasar, 1940,
type locality ‘Benguela’ [Angola]; original distribution cited for the
related tropical W African species, Parateuchus palemo (Olivier, 1789),
type locality, ‘Senegal au Cap’ [Senegal to the Cape of Good Hope],
presumably includes data for S. davisianus and S. bohemani.
Distribution: South Africa, Lesotho, Namibia, Botswana, Zimbabwe;
largely restricted to the summer rainfall region, but mostly absent from
centre of the subcontinent occupied by deep Kalahari sands; elytral
punctation variable from large pits with basal microgranulation, visible
at × 10, through medium to fine pits with basal microgranulation invis-
ible at macroscale; large pitted individuals in arid Upper Karoo and dry
Highveld (black squares); medium pitted individuals widespread with
cline to fine pitted individuals on arid W fringe of range (red squares);
both varieties (blue square).
Locality data (mean ± SD, range): Red square: Altitude: 1 091 ± 273,
344–1 998 m; annual rainfall: 338 ± 204, 65–846 mm; annual tem-
perature (max. + min. /2): 18.7 ± 1.6, 14.1–23.4°C (N=228). Black
square: Altitude: 1 326 ± 175, 797–1 765 m; annual rainfall: 309 ±
109, 197–639 mm; annual temperature (max. + min. /2): 15.4 ± 1.0,
13.4–17.5°C (N=62).
Habitat: In Gauteng bushveld: primarily on sandy clay loam (79) as op-
posed to deep sand (1) and mainly in grassland (76) as opposed to open
2 mm
woodland (4) or shaded thickets (0); in Northern Cape: data suggest a
soil generalist: sand (3.2), loamy sand (5.4), sandy loam (1.8), sandy clay
loam (3.2); DBRU collection records also suggest a soil generalist and
a bias to open vegetation: sand (10), sandy loam (9), sandy clay loam
(5) in pasture/grassland (6), scrub/shrubland (12), open woodland (6).
Food types: No adequate quantitative assessment, but, in Botswana, biased to carrion (5) and pig dung (6) as opposed to
herbivore dung: elephant (2), cattle (1), sheep (1); DBRU collection records on dung of human (2), baboon (1), horse (1),
donkey (1), cattle (17), wildebeest (1).
Temporal activity: Diurnal flight activity; seasonal activity primarily during the summer rains (Oct. to Apr.); at Phalaborwa:
activity greatest immediately in warm weather after heavy (24) or light rainfall (18), but lower under hot dry conditions (8).
Ecoregions Namibia, Botswana, Zimbabwe: Nama Karoo (AT1314), Namibian Savanna Woodlands (AT1316), SW Kalahari
Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands (AT0709), Southern African Bushveld (AT0717), Zambezian
Baikiaea Woodlands (AT0726), Southern Miombo Woodlands (AT0719), marginal occurrence in two other ecoregions.
Bioregions South Africa: Red square: centred on Bushmanland (NKb), Kalahari Duneveld (SVkd), Eastern Kalahari Bushveld
(SVk), Central Bushveld (SVcb), Mopane (SVmp); Black square: centred on Upper Nama Karoo (NKu), Lower Karoo
(NKl); overlap zone in Dry Highveld Grassland (Gh); marginal in four other bioregions.
Assessment rationale: EOO = 1 063 280 km2; widespread; mainly in more open vegetation at moderate altitude on various
soil types; some vegetation units in NE part of South African range subject to transformation (Gh: 4–63%, SVcb: 2–49%,
SVmp: 0–20%), but often very abundant after rainfall in the SW part of the range, which is little transformed (0–2% NKb,
NKl, SVkd, W SVk), therefore, assessed as Least Concern (LC).
Conservation measures: None recommended as it is widespread and often abundant in both farmland and reserves; protected
in various nature reserves, e.g. Roodeplaat, Abe Bailey, Leeuwfontein (South Africa); however possible taxonomic implica-
tions of the morphological variation should be investigated.

SCARABAEINI
SURICATA 6 (2020) 595

Scarabaeolus canaliculatus
(Fairmaire, 1888)

No synonyms
Global: LC
Endemic: Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus canaliculatus: ‘Namaqua’ [interpreted as Great

Namaqualand, southern Namibia, as it was described together with
Pachysoma schinzi Fairmaire, 1888, a Great Namaqualand endemic].
Taxonomy: Accepted species; not to be confused with the recently reval-
idated S. reichei Waterhouse, 1890, from the SW Cape, South Africa
(but see comments in Taxonomy section for S. reichei).
Distribution: Sporadically recorded in the southern Namib Desert, Na-
mibia, which is consistent with a Great Namaqualand type locality.
Locality data (mean ± SD, range): Altitude: 894 ± 160, 782–1 007 m;
annual rainfall: 67 ± 21, 52–81 mm; annual temperature (max. + min.
/2): 15.3 ± 0.4, 15.0–15.6°C (N=2).
Habitat: No quantitative assessment; one record from dunes, another
from the edge of a pan.
Food types: Not known.
Temporal activity: Diel flight periodicity unknown, but tan-coloured
pilosity is indicative of flight activity during darkness; recorded during
the late summer rainy season (Mar.).
Ecoregions Namibia: Range straddles the boundary between Namib Des-
ert (AT1315) and Succulent Karoo (AT1322).
Assessment rationale: EOO = 910 km2 (presumably underestimated);
although this species may occupy a relatively small EOO and AOO
below the W escarpment in SW Namibia, the two known exact records
probably represent a collecting artefact rather than any particular rarity
within this range; assessed as Least Concern (LC) rather than Data De-
ficient (DD) since the few known localities occur within a large reserve. 5 mm

Conservation measures: Assessment of conservation status would be

much improved by a survey to determine the full EOO and AOO
including quantitative data on ecological associations; both known,
exact records occur within the boundaries of the Namib-Naukluft or
Sperrgebiet National Parks (Namibia).

SCARABAEINI
596 SURICATA 6 (2020)

Scarabaeolus carniphilus
(Davis & Deschodt, 2015b)

No synonyms
Global: DD
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) carniphilus: ‘Mabuasehube

Game Reserve, SW Botswana’.
Taxonomy: Accepted species.
Distribution: Known only from three SW Kalahari localities in SW Bot­
swana and the Northern Cape, South Africa; probably poorly collected
due to dietary specialisation.
Locality data (mean ± SD, range): Altitude: 1 135 ± 226, 987–1 395 m;
annual rainfall: 334 ± 86, 283–433 mm; annual temperature (max. +
min. /2): 19.4 ± 1.5, 17.7–20.4°C (N=3).
Habitat: No quantitative assessment; all three known localities occur on
deep sands in pasture or arid grassland.
Food types: In Botswana: sampled to carrion (7) and sheep dung (2); one
DBRU collection record from cattle dung.
Temporal activity: Diel flight periodicity unknown, but probably diurnal
based on black pilosity; all specimens recorded during the late summer
rainy season (Feb.).
Ecoregions Botswana: Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk) (Savanna
Biome).
Assessment rationale: EOO = 4 285 km2 (underestimated); likely a carri-
on specialist restricted to the deep sands and open vegetation of the SW
Kalahari; however, known by only 10 specimens from three localities,
so must be assessed as Data Deficient (DD).
Conservation measures: A survey of the SW Kalahari is required to assess
2 mm the conservation status of this species; this should determine its full
EOO, AOO and provide quantitative data on its ecological habits; pro-
tected in Mabuasehube Game Reserve (Kgalagadi Transfrontier Park),
Botswana.

SCARABAEINI
SURICATA 6 (2020) 597

Scarabaeolus clanceyi
(Ferreira, 1954d)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Scarabaeus clanceyi: ‘Manguzi River, near Maputa, Zulu-

land’ [KwaZulu-Natal, South Africa].
Taxonomy: Accepted species.
Distribution: Localised in open vegetation on particularly sandy soils at
the S tip of the Mozambique Coastal Plain: NE South Africa, S Mo-
zambique.
Locality data (mean ± SD, range): Altitude: 167 ± 195, 0–443 m; annual
rainfall: 673 ± 98, 529–780 mm; annual temperature (max. + min. /2):
22.3 ± 0.3, 21.5–22.6°C (N=14).
Habitat: No adequate quantitative assessment of soil association; at Pha­
laborwa: observed on sandy soils primarily in open grassy patches with
scattered shrubs and trees (111), few in dense vegetation (5); on deep
sand at Maputo Elephant Reserve: primarily recorded in grassland of
saline pans (53), uncommon in small forest patches (4) and absent
from large forest patches (0).
Food types: At Phalaborwa: very strong bias to omnivore dung (pig: 110),
rare on cattle dung (6), absent from elephant dung (0).
Temporal activity: Diurnal flight activity during the summer rainy season
(Dec. to Feb.); at Phalaborwa: primarily active early on a dry sunny day
two days after light rainfall (88), less active on a sunny day following
very heavy rainfall (21), least on a cloudy day following light rainfall (7).
Ecoregions Mozambique: Localised in Maputaland Coastal Forest Mo-
saic (AT0119).
Bioregions South Africa: Localised in Mopane (SVmp), Lowveld (SVl)
(Savanna Biome); Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 20 500 km2; much of the known distribu-
tion occurs within conserved areas although it has also been recorded
in relative abundance in communal grazing lands where the woodland
has been opened up by collection of firewood; it would, therefore, face
few current threats and is assessed as Least Concern (LC). 2 mm

Conservation measures: Assessment of conservation status would be

improved by quantitative data on soil associations, particularly to de-
termine if the potential AOO is reduced by soil specialisation; however,
most of the known AOO is protected in Maputo Special Reserve (Mo-
zambique) and the central Kruger National Park (South Africa).

SCARABAEINI
598 SURICATA 6 (2020)

Scarabaeolus damarensis
(Janssens, 1940b)

= Scarabaeus azureus Ferreira, 1952a

Global: LC (see IUCN Red List – LC as Scarabaeus)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Scarabaeus damarensis: ‘Damara’ [Damaraland, Namib-

ia]; S. azureus: ‘Ovampoland (Oshikango)’ [Ovamboland, Namibia].
Taxonomy: Accepted species.
Distribution: Centred on the southern Kalahari, but several peripheral
records suggest potentially wider occurrence that is possibly limited by
specialisation to open vegetation on deep sands; much less abundant in
wooded NE Botswana (27) than open vegetation of arid SW (1072);
known distribution only South Africa, Botswana, Namibia.
Locality data (mean ± SD, range): Altitude: 1 078 ± 129, 596–1 402 m;
annual rainfall: 297 ± 86, 174–599 mm; annual temperature (max. +
min. /2): 19.7 ± 1.1, 17.9–22.7°C (N=113).
Habitat: No quantitative assessment; DBRU collection records primarily
on deep sand (46), sandy loam (1) and primarily in open vegetation,
pasture/grassland (16), scrub (12), shrubland (13), open woodland (6);
northern outlier records mostly on floodplain grasslands.
Food types: In Botswana: primarily attracted to the dung of omnivores
(pig: 341) and ruminant herbivores (cattle: 177, sheep: 297), less
abundant on monogastric herbivore dung (elephant: 21) and carrion
(chicken livers: 63); DBRU collection records on dung of human (1),
cattle (23), wildebeest (1), sheep (1), horse (3), donkey (3).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.).
Ecoregions Namibia, Botswana: Centred on Kalahari Xeric Savanna
(AT1309), also open vegetation in Angolan Mopane Woodlands
5 mm
(AT0702), Kalahari Acacia-Baikiaea Woodlands (AT0709), Zambezian
and Mopane Woodlands (AT0725).
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk) (Savanna Biome).
Assessment rationale: EOO = 952 000 km2; widely distributed in little-
transformed (0–2%, SVkd, W SVk), dry areas of the SW Kalahari that
are used primarily for conservation or the grazing of domestic livestock
to which S. damarensis is readily attracted; therefore, assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on habitat associations, particularly to
demonstrate how these influence the EOO and AOO; protected within
the borders of Chobe National Park, Central Kalahari Game Reserve
(Khutse), and Kgalagadi Transfrontier Park (Botswana and South Af-
rica).

SCARABAEINI
SURICATA 6 (2020) 599

Scarabaeolus flavicornis
(Boheman, 1860)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Ateuchus flavicornis: ‘prope fluvium Svakop’ [near Swa-

kop River, W central Namibia].
Taxonomy: Accepted species.
Distribution: Centred on S Damaraland (Namibia), SW Kalahari, W
coast and E marginal, Kalahari sand outliers (Botswana, South Africa);
abundant on Liuwa floodplain grasslands (Zambia), but rare on lacus-
trine sands (Savuti, N Botswana), range probably dictated by speciali-
sation to drier, open vegetation on deep sands; much less abundant in
wooded NE Botswana (80) than open vegetation of arid SW (2 971).
Locality data (mean ± SD, range): Altitude: 988 ± 341, 0–1 782 m; an-
nual rainfall: 272 ± 143, 9–799 mm; annual temperature (max. + min.
/2): 19.1 ± 1.5, 15.2–22.4°C (N=178).
Habitat: In Gauteng: recorded exclusively on deep sand (11) not on sandy
clay loam (0), primarily in grassland (10) as opposed to open woodland
(1) or shaded thickets (0); supported by DBRU collection records: all
records from deep sand (12) mainly in open vegetation: grassland (3),
scrub (2), shrubland (4), open woodland (3).
Food types: In Botswana: extreme bias in attraction to carrion (chicken
livers: 1 072) and pig dung (1 873) compared to herbivore dung, el-
ephant (48), cattle (32), sheep (26); DBRU collection records from
human (1), cattle (10) and donkey dung (1).
Temporal activity: Flight activity in darkness as indicated by the tan-
coloured pilosity; recorded mainly during the summer rainy season in
the NE (Oct. to May), also spring (Aug., Sept.) in the SW (winter
rainfall region).
Ecoregions Namibia, Botswana: Centred on Kalahari Xeric Savanna 2 mm
(AT1309), central Namibian Savanna Woodlands (AT1316) with pe-
ripheral occurrence in four other ecoregions.
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk) (Savanna Biome); Namaqualand Sandveld
(SVs) (Succulent Karoo Biome); N Upper Karoo (NKu), N Bushman-
land (NKb) (Nama Karoo Biome); plus E sand outliers in Grassland
and Savanna Biomes.
Assessment rationale: EOO = 844 500 km2; wide distribution centred in
little-transformed, arid areas used primarily for the grazing of domestic
livestock to which it is attracted even though there is a much stronger
bias to carrion and omnivore dung; assessed as Least Concern (LC).
Conservation measures: None required; about 5% of the range protected
in the Central Kalahari Game Reserve (Botswana), Kgalagadi Trans-
frontier Park (Botswana, South Africa) and Namaqua National Park
(South Africa).

SCARABAEINI
600 SURICATA 6 (2020)

Scarabaeolus fritschi
(Harold, 1868c)

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus fritschi: ‘Afric. Austral inter.’ [interior of

southern Africa].
Taxonomy: Accepted species.
Distribution: A wide E–W pattern of occurrence across the arid uplands
in the centre of the Northern Cape, South Africa that swings north-
wards along the upper edge of the western escarpment into S Namibia;
range mainly within the arid late summer rainfall region.
Locality data (mean ± SD, range): Altitude: 1 047 ± 228, 308–1 598 m;
annual rainfall: 192 ± 55, 58–267 mm; annual temperature (max. +
min. /2): 17.6 ± 1.4, 14.1–19.9°C (N=63).
Habitat: No quantitative assessment; observed on patches of arid deep or
loamy sands, some piled against mountains, occurring as outlier dunes,
fringing dunes of pans, or in beds of dry saline pans supporting grass-
land or a mixture of open scrub, shrubs and grassland.
Food types: No quantitative assessment; attracted to various dung types
including those of cattle, sheep, horse.
Diel activity: Diurnal flight activity; spring (Aug. to Oct.) and autumn
activity recorded (Feb., Mar.); absence of mid-summer records may be
a collecting artefact.
Ecoregions Namibia: Nama Karoo (AT1314).
Bioregions South Africa: Centred on N Upper Karoo (NKu), Bushman-
land (NKb) (Nama Karoo Biome); also Richtersveld (SKr), Namaqua-
land Hardeveld (SKn) (Succulent Karoo Biome) above W escarpment.
Assessment rationale: EOO = 110 000 km2; AOO would be somewhat
smaller owing to soil type specialisation; nevertheless the widespread
occurrence of patches of sandy soils and attraction to the dung of do-
mestic livestock on farms used primarily for grazing would mitigate for
5 mm the small part of the range that occurs within protected areas; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; range little
transformed (0–4% in South Africa); a small part is protected in the
Richtersveld National Park and the Riemvasmaak Conservancy (South
Africa).

SCARABAEINI
SURICATA 6 (2020) 601

Scarabaeolus gracai
(Ferreira, 1952a)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Moçambique, Província do Sul do Save: Massinga’ [Mo-

zambique].
Taxonomy: Accepted species.
Distribution: Coastal sands of S Mozambique and sand outliers strad-
dling the border of the Kruger National Park (South Africa) and Mo-
zambique.
Locality data (mean ± SD, range): Altitude: 337 ± 158, 101–429 m;
annual rainfall: 615 ± 267, 482–1 015 mm; annual temperature (max.
+ min. /2): 23.1 ± 0.2, 22.9–23.4°C (N=4).
Habitat: No quantitative assessment; recorded in unknown vegetation
types on deep sand patches and coastal sands.
Food types: Not known.
Temporal activity: Diel flight periodicity unknown, but assumed to be
diurnal on the basis of black pilosity; recorded during the summer rainy
season (Nov. to Apr.).
Ecoregions Mozambique: Sand localities: Southern Zanzibar-Inhambane
Coastal Forest Mosaic (AT0128) (not shown on map).
Bioregions South Africa: Deep sand patches in Mopane (SVmp), N
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 3 040 km2 (underestimated); known from
only four localities; likely a deep sand endemic, but otherwise very
poorly known; therefore, assessed as Data Deficient (DD).
Conservation measures: To assess its conservation status, a survey of the
S Mozambique Coastal Plain is required; this should determine its full
EOO, AOO and ecological associations; in South Africa, occurs only
in entirely protected, deep sand outliers in the N Kruger National Park.

2 mm

SCARABAEINI
602 SURICATA 6 (2020)

Scarabaeolus inoportunus
(Ferreira, 1953c)

No synonyms
Global: LC (see IUCN Red List – LC as Scarabaeus)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus inoportunus: ‘Província do Cabo: Griqualand

West, Niekerk’s Hope’ [Niekerkshoop, Northern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on the SW Kalahari with a bias to the arid SW;
peripheral records in S Damaraland (Namibia) plus W Free State and
North-West Province (South Africa).
Locality data (mean ± SD, range): Altitude: 1 055 ± 119, 714–1 349 m;
annual rainfall: 257 ± 77, 56–555 mm; annual temperature (max. +
min. /2): 19.4 ± 0.9, 16.6–22.0°C (N=67).
Habitat: No adequate quantitative assessment; observed in open shrub­
land and grassland on deep sands of plains, dunes and saline pans; at
Brulpan: recorded primarily in grassland on an outlier Kalahari dune
(966), rare in the surrounding Karoo shrubland on stony sand (1).
Food types: In Botswana: bias shown to omnivore dung (pig: 147) and ru-
minant herbivore pellets (sheep: 211) with carrion (chicken livers: 41),
and dung of other herbivores (cattle: 64, elephant: 13) less attractive.
Temporal activity: Diurnal flight activity recorded in most months of
the year (Dec. to Sept.); at Brulpan more abundant immediately after
substantial rainfall (967) than under hot dry conditions (342).
Ecoregions Namibia, Botswana: Centred on the Kalahari Xeric Savanna
(AT1309), also, SW margins of Kalahari Acacia-Baikiaea Woodlands
(AT0709).
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk) (Savanna Biome); N Upper Karoo (NKu), N
Bushmanland (NKb) (Nama Karoo Biome).
Assessment rationale: EOO = 510 500 km2; wide distribution and ready
2 mm
attraction to the range of dung types available in the little-transformed,
arid SW Kalahari (transformation 0–2% SVkd, W SVk) where grazing
of domestic livestock and conservation dominate land usage; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations; widespread
on farms and protected within the Kgalagadi Transfrontier Park (South
Africa, Botswana).

SCARABAEINI
SURICATA 6 (2020) 603

Scarabaeolus inquisitus
(Péringuey, 1908)

No synonyms
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type localities: ‘Transvaal (Potchefstroom, Plat River), Cape Colony

(Vryburg)’ [South Africa].
Taxonomy: Accepted species.
Distribution: Centred along the SE border of the S Kalahari (South Afri-
ca) with a few records from the Kalahari in central Botswana; unclear if
the disjunct pattern is real or a collection artefact.
Locality data (mean ± SD, range): Altitude: 1 141 ± 154, 923–1 395 m;
annual rainfall: 390 ± 134, 229–636 mm; annual temperature (max. +
min. /2): 18.9 ± 1.6, 15.6–22.1°C (N=22).
Habitat: No adequate quantitative assessment; in Northern Cape: more
abundant at four study sites on sand (27) than at four on sandy loam
(4); all known records of vegetation type in grassland or shrubland
(North Cape Survey, DBRU collection records).
Food types: No adequate quantitative assessment; in Botswana: recorded
most abundantly to carrion (chicken livers: 9), and uncommonly to
dung (pig: 1, elephant: 1, cattle: 2, sheep: 0).
Temporal activity: Diel flight periodicity unknown, but assumed to be
diurnal on the basis of black pilosity; recorded primarily during the
summer rainy season (Sept. to Mar., also May).
Ecoregions Botswana: Kalahari Acacia-Baikiaea Woodlands (AT0709).
Bioregions South Africa: Straddling parts of the Nama Karoo (NK),
Grassland (G) and Savanna (SV) biomes: N Upper Nama Karoo
(NKu); East Kalahari Bushveld (SVk), N Dry Highveld Grassland
(Gh), Central Bushveld (SVcb).
Assessment rationale: EOO = 446 500 km2; suspected to be a carrion 2 mm
specialist, primarily on deep sand patches in open vegetation, but range
and ecological data inconclusive owing to limited records; apparent
low population density may be an artefact of limited collection from
carrion; assessed as Least Concern (LC) on basis of wide range but
somewhat Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
improved by a survey to determine the full EOO and AOO as well as
further quantitative data on ecological associations; protected in the N
of the Central Kalahari Game Reserve (Botswana).

SCARABAEINI
604 SURICATA 6 (2020)

Scarabaeolus intricatus
(Fabricius, 1801)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus intricatus: ‘Cap Bon. Spei’ [Cape of Good

Hope, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on the coastal lowlands and western mountain es-
carpment in the winter rainfall region of South Africa, with one outlier
record from the Eastern Cape that may indicate a wider continuous
distribution in the bimodal rainfall region.
Locality data (mean ± SD, range): Altitude: 224 ± 269, 0–1 173 m;
annual rainfall: 233 ± 100, 63–466 mm; annual temperature (max. +
min. /2): 17.1 ± 1.1, 14.0–18.9°C (N=69).
Habitat: No adequate quantitative assessment; abundance in shrubland
at Modderrivier (1 406) and Pampoenvlei (814) greater on deep sand
than on sandy loam at Pampoenvlei (171); on arid deep sand at Geel-
bek: abundance in shrubland (247) strongly impacted by its clearance
and replacement by sparse pasture grass (19); DBRU collection records:
deep sand (5), sandy loam (2) in scrub (5) and pasture (2).
Food types: No quantitative assessment; DBRU collection records on
dung of human (1), cattle (8).
Temporal activity: Diurnal flight recorded in all months (Aug. to May)
except the coolest in mid-winter (June, July); in the SW Cape: sclero-
tised adults show a seasonal peak in early summer (Oct., Nov.), teneral
adults in Jan. and Apr.
Bioregions South Africa: Centred on sandy soils of the Succulent Karoo
(SK) and Fynbos (F) Biomes, especially: Namaqualand Sandveld (SKs),
vegetation units of Western Strandveld (FS), Sand Fynbos (FFd), Sand-
stone Fynbos (FFs); also, margins of Rainshadow Valley Karoo (SKv).
Assessment rationale: EOO = 48 000 km2 (W and SW localities); limited
data and observations suggest a deep sand specialist in natural shrub­
land with a strong negative response to invasive exotic shrubs as well
as the clearance of natural shrubland and its replacement by pasture
5 mm grassland; probably threatened in S parts of its range, particularly in
SW units of Sand Fynbos (FFd 2–7) and Western Strandveld (FS 1–6)
where there is 25–80% transformation; currently assessed as Least
Concern (LC) owing to little-transformed N of range.
Conservation measures: Assessment of conservation status would be
improved by further quantitative data on ecological associations; pro-
tection is provided across ± 10% of its range in Cape of Good Hope
Nature Reserve plus West Coast and Namaqua national parks (South
Africa).

SCARABAEINI
SURICATA 6 (2020) 605

Scarabaeolus karrooensis
(zur Strassen, 1961b)

No synonyms
Global: LC (see IUCN Red List – LC as Scarabaeus)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus karrooensis: ‘Noupoort (=Naauwpoort) im

Colesberg Distrikt’ [Noupoort, Eastern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: S of the arid late summer rainfall region and arid inland
part of the bimodal rainfall region: SW South Africa, S Namibia.
Locality data (mean ± SD, range): Altitude: 1 019 ± 128, 617–1 527 m;
annual rainfall: 210 ± 49, 81–348 mm; annual temperature (max. +
min. /2): 18.5 ± 1.3, 13.9–21.3°C (N=45).
Habitat: No quantitative assessment; known from both sand dunes and
finer-grained soils in scrub, but data too limited to determine if this
species is a habitat generalist or specialist.
Food types: No quantitative data; a few records from mixed cattle/sheep
dung baits.
Temporal activity: Diurnal flight activity during most months of the year
(not mid-summer: Jan.; not mid-winter: July) reflecting a range across
regions with either bimodal (spring/autumn) or late summer rainfall.
Ecoregions Namibia: SW Kalahari Xeric Savanna (AT1309), S Namibian
Savanna Woodlands (AT1316).
Bioregions South Africa: Rainshadow Valley Karoo (SKv) (Succulent
Karoo Biome); Upper Nama Karoo (NKu), N uplands Bushmanland
(NKb) (Nama Karoo Biome); Kalahari Duneveld (SVkd) (Savanna
Biome).
Assessment rationale: EOO = 359 300 km2; widespread, but relatively
poorly known, possibly because of confusion with close relatives; lo-
calities primarily from arid farmland used for grazing of domestic live-
5 mm
stock; assessed as Least Concern (LC) because of the large range across
a little-transformed region; however, essentially Data Deficient (DD).
Conservation measures: A survey across the full extent of Karoo regions
is required to ascertain the full EOO and AOO. Assessment of conser-
vation status would also be improved by quantitative data on ecological
associations; protected in Kgalagadi Transfrontier Park (South Africa)
and Namib-Naukluft Park (Namibia).

SCARABAEINI
606 SURICATA 6 (2020)

Scarabaeolus kochi
(Ferreira, 1952b)

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus kochi: ‘Kuke Pan’ [waterhole in Botswana].

Taxonomy: Accepted species.
Distribution: Centred on the arid SW Kalahari in Botswana and the
Northern Cape, South Africa, but possibly also with a continuous dis-
tribution further SW wherever deep sands occur.
Locality data (mean ± SD, range): Altitude: 1 094 ± 128, 876–1 485 m;
annual rainfall: 274 ± 68, 100–479 mm; annual temperature (max. +
min. /2): 19.1 ± 1.4, 15.1–22.1°C (N=53).
Habitat: No adequate quantitative assessment; in Northern Cape: higher
frequency on deep, loamy or stony sands in grassland or open shrubland
(n=1.0/trap at 37 sites) than on sandy loam (n=0.7/trap at 13 sites).
Food types: In Botswana: strongest bias to carrion (chicken livers: 148)
and omnivore dung (pig: 74); less frequent on herbivore dung (ele-
phant: 26, cattle: 28, sheep: 19); carrion bias supported by DBRU
collection records on goat meat (1) and dead millipede (1).
Temporal activity: Diurnal flight activity in the late summer rainy season
(Jan. to Mar.).
Ecoregions Namibia, Botswana: SE Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), Eastern
Kalahari Bushveld (SVk) (Savanna Biome); also deep sands bordering
saline pans in N Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 430 000 km2; widespread in a little-
transformed arid area that is either conserved or used for grazing do-
5 mm mestic livestock; probably a sand specialist, but support from quanti-
tative data required; assessed as Least Concern (LC) on the basis that
deep sand habitat, carrion and various dung types would be widely
available across its large EOO.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations; ± 15% of
the known range protected in the Central Kalahari Game Reserve and
Kgalagadi Transfrontier Park (Botswana and South Africa).

SCARABAEINI
SURICATA 6 (2020) 607

Scarabaeolus lucidulus
(Boheman, 1860)

= Scarabaeus vansoni Ferreira, 1958c

Global: DD

J A S O N D J F M A M J

Type localities: As Ateuchus lucidulus: ‘juxta Lacum N’gami et fluvium

Kuisip’ [both near Lake Ngami, Botswana, and Kuiseb River, Namib-
ia]; S. vansoni: ‘Mangetti’ [quarantine station, Ovamboland, Namibia].
Taxonomy: Accepted species; S. vansoni Ferreira, 1958c is a recent junior
synonym.
Distribution: Scattered records in savanna on deep sands of Namibia,
Botswana, Zimbabwe, South Africa.
Locality data (mean ± SD, range): Altitude: 1 020 ± 139, 707–1189 m;
annual rainfall: 559 ± 75, 414–647 mm; annual temperature (max. +
min. /2): 20.9 ± 1.5, 18.9–23.1°C (N=12).
Habitat: No quantitative assessment; all Namibia, Botswana and Zimbab­
we localities map as deep sand; Nylsvlei localities recorded as sand;
probably a deep sand specialist, possibly in open woodland.
Food types: No adequate quantitative assessment; at Nylsvlei: recorded to
banana (16), carrion (7), human dung (4) and unspecified dung (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Mar.).
Ecoregions Namibia, Botswana, Zimbabwe: Kalahari Acacia-Baikiaea
Woodlands (AT0709), Zambezian Baikiaea Woodlands (AT0726).
Bioregions South Africa: Vegetation unit: Central Sandy Bushveld (SVcb
12) (Central Bushveld (SVcb), Savanna Biome).
Assessment rationale: EOO = 382 950 km2 (dual type localities in Bot­
swana and W Namibia suggest the EOO is underestimated); AOO
presumed to be smaller according to occurrence of deep sand covered in
woodland, which is, nevertheless, widespread across the range although
SVcb 12 is 24% transformed; not well known, possibly owing to bias
to rotting plant matter and animal carrion rather than dung; owing to
limited information assessed as Data Deficient (DD).
Conservation measures: A quantitative survey is required to assess con- 2 mm
servation status following the recent revision of the species taxonomy;
the EOO, AOO and ecological associations have been hypothesised
on the basis of limited observations; however, they are poorly known
and require better support; protected in Nylsvlei Nature Reserve (South
Africa).

SCARABAEINI
608 SURICATA 6 (2020)

Scarabaeolus megaparvulus
(Davis & Deschodt, 2015b)

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) megaparvulus: ‘Farm Rooiput’

[Northern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Restricted to the arid late summer rainfall region; arid N
foothills of the Swartberg, plateau of Upper Karoo in South Africa, and
upper edge of the W escarpment in SW Namibia.
Locality data (mean ± SD, range): Altitude: 1 083 ± 241, 773–1 762 m;
annual rainfall: 177 ± 65, 71–275 mm; annual temperature (max. +
min. /2): 17.1 ± 1.8, 13.0–19.3°C (N=40).
Habitat: No adequate quantitative assessment; in Northern Cape: sam-
pled from Upper Karoo and uplands S of Orange River, on both sand
(1.3/trap, n=17 localities) and sandy loam or sandy clay loam (0.99/
trap, n=14); DBRU collection records entirely from deep sand (6) in
arid grassland (3), scrub (1), open woodland (1).
Food types: No quantitative assessment; sampled to mixed cattle/sheep
dung baits; DBRU collection records from dung of cattle (1), donkey
(1).
Temporal activity: Diurnal flight activity primarily during early/late
summer in extreme S part of range (bimodal rainfall region) and late
summer to the N in the arid late summer rainfall region (overall: Nov.
to Mar.).
Ecoregions Namibia: SW margin of Namibian Savanna Woodlands
(AT1316) overlooking the Namib Desert.
Bioregions South Africa: Higher-lying parts of N Bushmanland (NKb),
Upper Karoo (NKu), Lower Karoo (NKl) (Nama Karoo Biome).
2 mm Assessment rationale: EOO = 59 650 km2; a sizeable upland range across
farmland in a little-transformed arid region used primarily for the
grazing of domestic livestock to which this species is readily attracted;
currently assessed as Least Concern (LC) owing to widespread records
from cattle/sheep dung.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
hilly areas of the Namib-Naukluft National Park, Namibia; not so far
recorded from any protected area in South Africa, but not currently
considered to be threatened.

SCARABAEINI
SURICATA 6 (2020) 609

Scarabaeolus namibensis
(Zídek & Pokorný, 2018)

No synonyms
Global: LC
Endemic: ?Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) namibensis: ‘S.W. Afr. [Namib-

ia], 60 km NE Gobabeb’.
Taxonomy: Accepted species.
Distribution: Currently known only from south of the Cunene River in
two disjunct, arid, lowland population centres of the Kaokoveld in NW
Namibia and the Namib Desert in W central Namibia.
Locality data (mean ± SD, range): Altitude: 577 ± 170, 309–910 m;
annual rainfall: 60 ± 50, 27–228 mm; annual temperature (max. + min.
/2): 16.7 ± 0.1, 16.5–16.9°C (N=15).
Habitat: No quantitative assessment; population in W central Namibia
occurs on sparely grassed gravel plains of the Namib Desert.
Food types: No quantitative assessment; attracted to camel and pig dung.
Temporal activity: Diurnal flight activity; limited records in Sept., Jan. to
Mar. and June insufficient to determine if activity is aseasonal like some
other coastal desert Scarabaeolus species, e.g. S. rubripennis (Boheman,
1860).
Ecoregions Namibia: Kaokoveld Desert (AT1310), Namib Desert
(AT1315).
Assessment rationale: EOO = 27 000 km2; although known from only
a few widely separated areas, this is probably a collection artefact; after
light rain during an EIA study in June at Ongolo and Tumas (W cen-
tral Namibia), there was a high local frequency of capture, which may
suggest a continuous geographical distribution between population
centres; assessed as Least Concern (LC) on basis of wide EOO.
Conservation measures: Assessment of conservation status would be im-
2 mm
proved by quantitative data on ecological associations; also, a survey is
required to determine if there is continuous N–S distribution includ-
ing protected areas along the Skeleton Coast in the N Namib Desert;
although the Kaokoveld locality lies outside of reserves, the central
Namibian localities lie within the boundaries of the Namib-Naukluft
National Park; whilst the southernmost of these localities remain pro-
tected, those to the north (Ongolo, Tumas) lie within an area scheduled
for mining.

SCARABAEINI
610 SURICATA 6 (2020)

Scarabaeolus niemandi
(Deschodt & Davis, 2015b)

No synonyms (but see Taxonomy)

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) niemandi: ‘Soutpansberg, Lim-

popo, S22.87561° E29.73128°’ [mountain range, Limpopo Province,
South Africa].
Taxonomy: Accepted species; but see Preamble, notes with Scarabaeolus
rugosipennis (Zídek & Pokorný, 2018).
Distribution: Only known from dry savanna on the N faces of the Blou-
berg, Soutpansberg and Waterberg, W Limpopo Province, South Afri-
ca; unclear if the distribution is continuous or disjunct.
Locality data (mean ± SD, range): Altitude: 1 006 ± 269, 804–1 478 m;
annual rainfall: 521 ± 76, 460–651 mm; annual temperature (max. +
min. /2): 19.6 ± 1.4, 17.1–20.6°C (N=5).
Habitat: No quantitative assessment; possibly a deep sand specialist in
open woodland owing to records coinciding with sand at edge of
Central Sandy Bushveld (SVcb 12) and Musina Mopane Bushveld
(SVmp 2); also in Roodeberg Bushveld (SVcb 18) with mostly sandy
soils.
Food types: No quantitative assessment.
Temporal activity: Diel flight periodicity unknown, but presumed to be
diurnal on the basis of black pilosity; recorded during the summer rainy
season (Nov., Feb.).
Bioregions South Africa: Mopane (SVmp), Central Bushveld (SVcb)
(Savanna Biome).
Assessment rationale: EOO = 4 395 km2 (underestimated); relative-
2 mm ly poorly known species recently recorded from five localities with a
known EOO < 5 000 km2; AOO may be smaller according to occur-
rence of deep sand patches; transformation of range varies from 3%
(SVmp 1), 18% (SVcb 18) to 24 % (SVcb 12), with the type locality
scheduled for mining; although it would currently qualify for an IUCN
threat category, it must be assessed as Data Deficient (DD) owing to
the paucity of available information.
Conservation measures: To generate an assessment of conservation status,
a quantitative survey of arid savanna in Limpopo Province is required
to determine the full EOO; ecological associations also need to be de-
termined from quantitative data as the AOO could be restricted to
deep sand patches in open woodland; currently protected on the north
slopes of the Blouberg Nature Reserve and the private Goro Nature
Reserve.

SCARABAEINI
SURICATA 6 (2020) 611

Scarabaeolus obsoletepunctatus
(Balthasar, 1940)

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: As Scarabaeus obsoletepunctatus: ‘Angola, Benguella’ [Ben-

guela, Angola].
Taxonomy: Accepted species; a slightly larger-bodied, close relative of
S. bohemani Harold, 1868, with largely effaced elytral punctation; in-
dividual pictured, here, requires validation as S. obsoletepunctatus.
Distribution: Arid coastal region of NW Namibia, SW Angola.
Locality data (averages): Altitude: 559 m; annual rainfall: 228 mm; an-
nual temperature (max. + min. /2): 16.9°C (N=1).
Habitat: Not known.
Food types: Not known.
Temporal activity: Diurnal flight activity presumed on the basis of black
pilosity; recorded during the late summer rainy season (Feb.).
Ecoregions Namibia: One exact record from margin of Kaokoveld Desert
(AT1310) and N Namibian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 7 205 km2 (underestimated, but EOO
would be relatively restricted; probably mainly coastal SW Angola);
very poorly known taxonomically, biogeographically and ecologically;
assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative
survey of the Kaokoveld and SW Angola is required; this needs to de-
termine EOO, AOO and ecological associations; not currently known
from any protected region.

2 mm

SCARABAEINI
612 SURICATA 6 (2020)

Scarabaeolus pabulator
(Péringuey, 1908)

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Scarabaeus pabulator: ‘Cape Colony (Calvinia)’ [North-

ern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Centred on the S of the arid, late summer rainfall region,
primarily in climate types III1 and III2 (Walter & Lieth 1964): NW
South Africa, S Namibia; outlier occurrence at Nylsvlei requires valida-
tion (black square).
Locality data (mean ± SD, range): Altitude: 941 ± 166, 608–1 303 m;
annual rainfall: 168 ± 85, 43–585 mm; annual temperature (max. +
min. /2): 18.8 ± 1.2, 15.8–20.6°C (N=40).
Habitat: No adequate quantitative assessment; in Northern Cape: pri-
marily recorded in arid Bushmanland (n=30 sites) rather than moister
Karoo (n=7) or Kalahari (n=1); sampled from deep and stony sand
(1.8/trap) and sandy loam (0.6/trap) in scrub and open shrubland.
Food types: No quantitative assessment; sampled to dung of pig and
mixed cattle/sheep baits.
Temporal activity: Diurnal flight activity primarily in the late summer
rainy season (Jan. to Mar.); other records may reflect a distribution
extending to the edge of the winter rainfall region.
Ecoregions Namibia: S Namibian Savanna Woodlands (AT1316).
Bioregions South Africa: Centred on Bushmanland (NKb), margins of
Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 247 150 km2 (minus the Limpopo Prov-
ince record in a deep sand outlier at Nylsvlei – black square); although
few of the known records lie within reserves, S. pabulator is widespread
on farms in the Northern Cape, South Africa, that are used primarily
2 mm for the grazing of domestic livestock and have undergone very little
transformation; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by a quantitative survey to determine the full EOO, AOO and
ecological associations; it is necessary to determine if a bias to sandy
soils reduces the potential AOO; protected within the boundaries of
the Namib-Naukluft National Park, Namibia and the Riemvasmaak
Community Conservancy, South Africa.

SCARABAEINI
SURICATA 6 (2020) 613

Scarabaeolus parvulus
(Boheman, 1860)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Ateuchus parvulus: ‘Regione fluvii Svakop’ [Swakop

River region, W central Namibia].
Taxonomy: Accepted species.
Distribution: Centred on the arid late summer rainfall region from the
Northern Cape, South Africa, across extreme SW of Botswana plus S
and NW Namibia to SW Angola.
Locality data (mean ± SD, range): Altitude: 991 ± 279, 47–1 809 m;
annual rainfall: 196 ± 71, 27–367 mm; annual temperature (max. +
min. /2): 18.5 ± 1.6, 14.1–21.2°C (N=108).
Habitat: No adequate quantitative assessment; in the Northern Cape:
recorded at many study sites in SW Kalahari (26) and N Nama Karoo
(49) primarily on deep or stony sand (3.6/trap; n=55), less in sandy
loam (0.6/trap; n=19); DBRU collection records primarily from sand
(1), sandy loam (1) in open vegetation: grassland (3), scrub/shrubland
(8), open woodland (2).
Food types: No adequate quantitative assessment; in Botswana: sampled
to pig (2) and cattle (4) dung; limited DBRU collection records from
various dung types: cattle (3), wildebeest (1) donkey (1), horse (1),
springbok (1), and carrion: dead dog (1).
Temporal activity: Diurnal flight activity; recorded throughout the year
(Oct. to Aug.), possibly because of a range extending from a seasonal
rainfall system into the Namib Desert.
Ecoregions Namibia, Botswana: Nama Karoo (AT1314), Namib Desert
(AT1315), Kaokoveld Desert (AT1310), Namibian Savanna Wood-
lands (AT1316), SW Kalahari Xeric Savanna (AT1309).
2 mm
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), arid
deep sands that dominate W Eastern Kalahari Bushveld (SVk) (Savan-
na Biome), N Upper Karoo (NKu), N Bushmanland (NKb) (Nama
Karoo Biome).
Assessment rationale: EOO = 398 100 km2; widespread across a little-
transformed, arid region used primarily for the grazing of domestic
livestock with some areas under conservation; also recorded on a wide
variety of dung types; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations; in South
Africa, officially protected only across a very small part of its range in
Kgalagadi Transfrontier Park (South Africa, Botswana); also protected
in the Namib-Naukluft Park (Namibia).

SCARABAEINI
614 SURICATA 6 (2020)

Scarabaeolus planipennis
(Davis & Deschodt, 2015b)

No synonyms (but see Taxonomy)

Global: LC

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) planipennis: ‘Maputo Elephant

Reserve, Mozambique’.
Taxonomy: Accepted species; new species, Scarabaeolus krugeri (Zídek &
Pokorný, 2018), is considered an unpublished synonym of S. planipen-
nis (see Preamble).
Distribution: Known from coastal sands at the S tip of the Mozambique
Coastal Plain; also inland sand outliers up the valley of the Limpopo
River: South Africa, Mozambique.
Locality data (mean ± SD, range): Altitude: 92 ± 161, 0–475 m; annual
rainfall: 685 ± 99, 482–773 mm; annual temperature (max. + min. /2):
22.5 ± 0.3, 22.4–23.4°C (N=12).
Habitat: On deep sand in Maputo Special Reserve, Mozambique: associ-
ated with grassland of saline pans (17) rather than forest patches (small
(1), medium (0), large (0)); in Tembe and Sileza game reserves: asso-
ciated with mixed woodland (19) rather than sand forest (3); Kruger
National Park records all from sandveld.
Food types: No published quantitative data; cited as showing a bias to
omnivore rather than herbivore dung.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.).
Ecoregions Mozambique: Grassy sand patches in Maputaland Coastal
Forest Mosaic (AT0119).
Bioregions South Africa: Vegetation units on deep sand: Nwambyia–
Pumbe Sandy Bushveld (SVl 2), Tembe Sandy Bushveld (SVl 18),
2 mm
Cathedral Mopane Bushveld (SVmp 3) (Lowveld (SVl) and Mopane
(SVmp), Savanna Biome); Maputaland Wooded Grassland (CB 2) (In-
dian Ocean Coastal Belt Biome (CB)).
Assessment rationale: EOO = 4 900 km2 (may be underestimated); AOO
would be even smaller as the species is probably a deep sand specialist
in more open vegetation, but improved supporting data is required;
although the AOO is potentially < 5 000 km2 all known localities are
found in conserved areas; therefore, not currently considered to be
threatened; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by a quantitative survey of the S Mozambique Coastal Plain to
better define the EOO, AOO and ecological associations; all known
localities protected in the Kruger National Park plus Sileza and Tembe
game reserves (South Africa) or the Maputo Special Reserve (Mozam-
bique).

SCARABAEINI
SURICATA 6 (2020) 615

Scarabaeolus reichei
(Waterhouse, 1890) (but see Taxonomy)

No synonyms
Global: LC (see IUCN Red List – DD as Scarabaeus canaliculatus
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus reichei: ‘Cape of Good Hope’ [South Africa].

Taxonomy: Accepted species; recently revalidated from synonymy with
S. canaliculatus Fairmaire, 1888; more recently, again incorrectly syn-
onymised with S. canaliculatus (Fairmaire, 1888) by Zídek and Pokorný
(2018), who did not see the lost type of S. canaliculatus or material of
that species from southern Namibia (Namib Desert).
Distribution: Restricted to the arid West Coast in the winter rainfall re-
gion of South Africa with N range limits in Little Namaqualand; no
overlap with the ranges of S. fritschi Harold, 1868 (Northern Cape
interior, South Africa and S Namibia) or S. canaliculatus Fairmaire,
1888 (S Namib Desert, Namibia).
Locality data (mean ± SD, range): Altitude: 99 ± 84, 0–357 m; annual
rainfall: 170 ± 52, 87–323 mm; annual temperature (max. + min. /2):
17.7 ± 0.8, 15.2–19.1°C (N=63).
Habitat: No adequate quantitative assessment; in Western Cape: sampled
on deep sand in open shrubland (Modderrivier: 3) or very sandy loam
in sparse grass (Groote Post: 16); DBRU collection records entirely
from deep sand (6) in scrub (4) or crop fields (2).
Food types: No quantitative assessment; DBRU collection records from
carrion (1 – dead fox) and several dung types: cattle (5), sheep (1).
Temporal activity: Diurnal flight activity recorded in the moist spring to
dry early summer (Aug. to Dec.) and in moist autumn (Apr.).
Bioregions South Africa: Centred on Namaqualand Sandveld (SKs) (Suc-
5 mm
culent Karoo Biome); vegetation units in West Strandveld (FS), Sand
Fynbos (FFd), margins of Sandstone Fynbos (FFs) (Fynbos Biome).
Assessment rationale: EOO = 12 500 km2; relatively restricted range al-
though it frequents both low profile natural scrub as well as disturbed
vegetation, including fallow crop fields and dry grassland pastures on
farms cleared of natural shrubs in the S of its range where there is from
35 to 70% transformation (FFd 4, FS 2, FS 3); perhaps less threatened
than the many extreme, winter rainfall, shrubland specialists, although
immigration into disturbed patches from natural vegetation cannot be
ruled out; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; a small part of
the range is protected in the coastal portion of the Namaqua National
Park, which lies in the little-transformed (2%) Namaqualand Sandveld
Bioregion (South Africa).

SCARABAEINI
616 SURICATA 6 (2020)

Scarabaeolus rubripennis
(Boheman, 1860) (but see Taxonomy)

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Ateuchus rubripennis: ‘in regione fluvii Kuisip’ [Kuiseb

River region, W central Namibia].
Taxonomy: Accepted species, but southern populations in NW South
Africa now separated as Scarabaeolus lizleri (Zídek & Pokorný, 2018).
Distribution: Centred on the main Namib Desert Dunefield, SW Na-
mibia, and dunes straddling the Namibia/South Africa border.
Locality data (mean ± SD, range): Altitude: 594 ± 396, 1–1 655 m; an-
nual rainfall: 56 ± 30, 1–132 mm; annual temperature (max. + min.
/2): 16.0 ± 0.8, 13.5–17.5°C (N=35).
Habitat: No quantitative assessment; presumed to be specialised to deep
sands of sparsely vegetated dunes with which most known localities
coincide.
Food types: No quantitative assessment; attracted in abundance to pig
dung.
Temporal activity: Diel flight periodicity unknown, but black pilosity is
indicative of day-flying habits; recorded throughout the year (Apr. to
Feb.) suggesting relatively aseasonal habits although this assessment is
based on presence or absence rather than quantitative data.
Ecoregions Namibia: Namib Desert (AT1315), Succulent Karoo
(AT1322).
Bioregions South Africa: Vegetation unit: Richtersveld Coastal Dune­
veld (SKs 1) (Namaqualand Sandveld (SKs), Succulent Karoo Biome)
(S. lizleri).
Assessment rationale: EOO = 55 500 km2; widespread across an arid,
lowland region that is mostly under official protection; no protection
5 mm at the extreme southern margin of the species range in South Africa
(now S. lizleri), which coincides with a vegetation unit (SKs 1) that is
partially transformed (10%) by mining; nevertheless assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; predominantly
protected in Namibia where most known localities lie within the Namib-
Naukluft or Sperrgebiet national parks; protection within South Africa
would also be desirable, particularly as this population has now been
separated as the new species, S. lizleri.

SCARABAEINI
SURICATA 6 (2020) 617

Scarabaeolus soutpansbergensis
(Deschodt & Davis 2015b)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus (Scarabaeolus) soutpansbergensis: ‘Soutpans-

berg’ [mountain range, Limpopo Province, South Africa].
Taxonomy: Accepted species.
Distribution: Dry savanna just to the S of the Soutpansberg and to the N
in the Limpopo Valley, Limpopo Province, South Africa.
Locality data (mean ± SD, range): Altitude: 741 ± 220, 429–1 020 m;
annual rainfall: 434 ± 128, 306–653 mm; annual temperature (max. +
min. /2): 21.2 ± 1.6, 19.2–23.4°C (N=10).
Habitat: No quantitative assessment; holotype recorded on deep sand in
open mopane woodland.
Food types: No quantitative assessment.
Temporal activity: Diel flight periodicity unknown, but presumed to be
diurnal on the basis of black pilosity; recorded during the summer rainy
season (Dec., Feb.).
Bioregions South Africa: Vegetation units: Musina Mopane Bushveld
(SVmp 1), Limpopo Ridge Bushveld (SVmp 2), Roodeberg Bushveld
(SVcb 18), N Makhado Sweet Bushveld (SVcb 20) (Mopane (SVmp)
and Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 5 100 km2 (underestimated); relatively
poorly known species with a known EOO hardly more than 5 000 km2;
recorded from four vegetation units that are 18–27% transformed
(SVcb) or only 1–2% transformed (SVmp); little known concerning
species attributes; assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a more extensive
quantitative survey is required to determine the full EOO, AOO, and
ecological associations; protected within the Blouberg and Musina na-
ture reserves (South Africa). 2 mm

SCARABAEINI
618 SURICATA 6 (2020)

Genus Scarabaeus Linnaeus, 1758

Type species and designation: Scarabaeus sacer Linnaeus, 1758, by subsequent designation (Hope 1837).
Synonyms: = Heliocantharus Macleay, 1821 (pars): Scarabaeus sacer Linnaeus, 1758, by original
designation.
= Ateuchus auctorum (not Weber, 1801).
= Sebasteos Westwood, 1847: Sebasteos galenus Westwood, 1847, by original designa-
tion.
= Neateuchus Gillet, 1911: Ateuchus proboscideus Guérin-Meneville, 1844, by original
designation.
= Drepanopodus Janssens, 1940: Ateuchus costatus Wiedemann, 1823, by monotypy.
Last review: Entire genus reviewed by zur Strassen (1967) with updated checklist (Zídek 2004);
many of the species listed by Zídek (2004) transferred to Scarabaeolus (Balthasar,
1965b), after it was raised to genus level by Moretto (2016b); review of subgeneric af-
filiations (Zídek & Pokorný 2011), mainly Palaearctic species, now all raised to generic
status (Ziani & Gudenzi 2012).

The designation of Scarabaeus hercules Linnaeus, 1758 (Subfamily Dynastinae), as the type species of the genus (Latreille
1810) takes precedence over the subsequent designation of Scarabaeus sacer Linnaeus, 1758 (Subfamily Scarabaeinae) as
the type species (Hope 1837). However, on the basis of prevailing usage and to avoid confusion, it has been proposed that
S. sacer should be conserved as the type species of Scarabaeus. This currently forms part of Open Case 3590 submitted for
a ruling from the International Commission of Nomenclature (Krell et al. 2012).

In its strictest sense as a genus of the subfamily Scarabaeinae, Scarabaeus Linnaeus, 1758, is centred on the Afrotropical
and S Palaearctic regions extending into the W edge of the Oriental region. A recent revision divided the Palaearctic and
Oriental members of the genus into four subgenera (Zídek & Pokorný 2008) now all raised to generic level (Ziani &
Gudenzi 2012). One of these genera, Escarabaeus Zídek & Pokorný, 2011, also occurs S of 15°S in southern Africa, where
it is represented by at least two species. However, affiliations have not been formally assessed for the remaining 31 (or 33)
southern African species currently listed as Scarabaeus. A phylogeny (Barbero et al. 1998) suggests that at least some of
these species would belong to different genera to those described for the Palaearctic. However, pending further study, they
remain listed as Scarabaeus species (importantly, in the 2019 version of the Catalogue of Life, S. hottentorum Péringuey,
1901 and S. savignyi Macleay, 1821, are transferred to the genus, Ateuchetus Bedel, 1892).

Cited synonyms are for pterous genera and have been reduced from those listed by Zídek and Pokorný (2004) according to
the generic-level divisions proposed by Ziani and Gudenzi (2012). They may change, again, after revision of generic-level
affiliations shown by the 31 (or 33) southern African species. As three out of five flightless taxa have been revalidated at
generic level, listed synonyms do not include the two remaining flightless taxa from Madagascar that are still officially
synonymised with Scarabaeus: Neomnematium Janssens, 1938a, Madateuchus Paulian, 1953.

Patterns of species distribution in southern Africa have been divided into four centres. One was centred on the SW Ka-
lahari plus Namib Desert (1 sp.) plus W coast of South Africa (1), just the N Namib (1), S Namib plus Namaqualand
(1), Namaqualand (1), W coast of South Africa (2) or SW Western Cape (1). Another was centred on the drier Highveld
and N Namibia (2), Highveld to Upper Karoo (1), just N Highveld (2) or Karoo (1). The third was centred on moist NE
Highveld (2), Maputaland (1), NE Highveld to S Cape (1), or just the S Cape (1). The fourth was centred on savanna, all
regions (2), just E and W (1), W (1), or E (5), also, E and Kalahari (1) or just Kalahari (1).

The ranges of about half of the species wholly or partly coincide with drier regions in which there has been relatively little
habitat transformation. A few show ranges that are restricted to areas of high transformation. A total of 17 species were
assessed as Least Concern (LC), one was assessed as Near Threatened (NT) and the remainder (13) as Data Deficient
(DD). Several of the latter may deserve an IUCN threat category, particularly those occupying small known EOOs in
areas of high transformation.

SCARABAEINI
SURICATA 6 (2020) 619

Scarabaeus alienus
Péringuey, 1901

No synonyms
Global: DD
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: Cited as ‘not known’.

Taxonomy: Accepted species.
Distribution: Extremely arid range; records from NW South Africa occur
at the northern limits of climate type III1 (Walter & Lieth 1964); un-
known if there is a continuous distribution between N and S records
assuming that the Namibian record in climate type III2 is valid.
Locality data (mean ± SD, range): Altitude: 574 ± 178, 400–755 m;
annual rainfall: 65 ± 41, 18–90 mm; annual temperature (max. + min.
/2): 17.2 ± 0.8, 16.7–18.1°C (N=3).
Habitat: No quantitative assessment; DBRU collection records from sand
(1) and sandy loam (1) in scrub (2).
Food types: No quantitative assessment; DBRU collection records from
dung of horse (1) and donkey (1).
Temporal activity: Diurnal flight activity observed during daytime under
dry, sunny conditions; shows spring activity (Sept.) in the winter rain-
fall region of South Africa.
Ecoregions Namibia: Namib Desert (AT1315).
Bioregions South Africa: Richtersveld (SKr) (Succulent Karoo Biome).
Assessment rationale: EOO = 18 000 km2; known from only three wide-
ly separated localities; rarity possibly an artefact of the dry conditions
under which the collections were made; two South African records
occur outside of protected areas; assessed as Data Deficient (DD).
Conservation measures: Conservation status cannot be assessed from
current data; a quantitative survey is required to ascertain the full
EOO, AOO and ecological associations; in South Africa, not currently 5 mm

recorded in Richtersveld National Park; protected in Namibia within

the Namib-Naukluft Park assuming that the record from Gobabeb is
valid.

SCARABAEINI
620 SURICATA 6 (2020)

Scarabaeus ambiguus
(Boheman, 1857)

= Actinophorus leei Shipp, 1896b

= Scarabaeus viator deceptor Péringuey, 1901 (pars)
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Ateuchus ambiguus: ‘Caffraria meridionali’ [southern

Africa]; A. leei: ‘Swan River, West Australia’ [locality labelling error];
S. viator deceptor: ‘(Cape Town), Transvaal (Potchefstroom), Bechua-
naland (Kanye)’ [Potchefstroom, North-West Province, South Africa;
Kanye, Botswana], [Cape Town type locality would be an error = S.
viator Péringuey, 1901].
Taxonomy: Accepted species.
Distribution: Drier W and NW highveld, South Africa, and drier N
upland plateau, Namibia; rare occurrence at Gobabeb in Namib Des-
ert; also rare in the Botswana Kalahari, which essentially represents a
disjunction in the range; SW edge of range approaches that of the very
close relative, S. viator Péringuey, 1901.
Locality data (mean ± SD, range): Altitude: 1 323 ± 247, 424–2 074 m;
annual rainfall: 509 ± 156, 22–833 mm; annual temperature (max. +
min. /2): 18.0 ± 1.7, 12.2–22.6°C (N=106).
Habitat: In Gauteng bushveld: strong bias to sandy clay loam (38)
compared to deep sand (5), and to grassland (38) compared to open
woodland (5) and shaded thicket (0); similar vegetation bias in Ezem-
velo Nature Reserve (now Telperion): grass (105), open woodland
(10); conflicting soil type results in open woodland at Leeuwfontein
Game Reserve: loamy sand (1 595), sandy loam (145), stony sandy
loam (857); DBRU collection records from sand (18), sandy loam
5 mm (22), sandy clay loam (7) in grassland/pasture (27), scrub/shrubland
(9), open woodland (8).
Food types: No quantitative assessment; DBRU collection records mainly
from dung of cattle (40), also horse (2), zebra (3).
Temporal activity: Diurnal flight activity during the summer rainy sea-
son (Oct. to May); in Gauteng bushveld: primarily active in the early
morning after substantial rainfall.
Ecoregions Namibia, Botswana: Angolan Mopane Woodlands (AT0702),
NW Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea
Woodlands (AT0709), margins of two other ecoregions.
Bioregions South Africa: Eastern Kalahari Bushveld (SVk), higher SE and
E Central Bushveld (SVcb) (Savanna Biome); Dry Highveld Grassland
(Gh), W margins Mesic Highveld Grassland (Gm) (Grassland Biome);
scattered records at NE margins of Upper Nama Karoo (NKu) (Nama
Karoo Biome).
Assessment rationale: EOO = 371 530 km2 (main range; excludes three
outliers); widespread on various soil types with a bias to open grassland
habitat that is dominant over much of its upland range; on basis of
wide occurrence and abundance in several reserves, assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data to resolve conflicts between existing
results for ecological associations and lack of data on effects of habitat
transformation; protected in various national parks and reserves includ-
ing Telperion, Leeuwfontein (South Africa), Etosha (Namibia).

SCARABAEINI
SURICATA 6 (2020) 621

Scarabaeus basuto
zur Strassen, 1962

= Scarabaeus natalensis zur Strassen, 1962

= Scarabaeus ambiguus (Boheman, 1857) sensu Péringuey, 1901
Global: LC
Endemic: RSA, Lesotho

J A S O N D J F M A M J

Type localities: S. basuto: ‘Molokoolu’ (= Mohlokulu), Mohales Hoek

Distrikt, Basutoland’ [Mohlokulu, Lesotho]; S. natalensis: ‘Newcastle,
Natal’ [KwaZulu-Natal, South Africa]; S. ambiguus sensu Péringuey:
‘Cape Colony (Cape Town), Transvaal (Potchefstroom), Natal (Est-
court)’ [South Africa: Western Cape, North-West Province, KwaZulu-
Natal; see Taxonomy section].
Taxonomy: S. natalensis was recently synonymised with S. basuto since
a principal character forms a NE–SW cline whereas other characters
occur asymmetrically at end points on the cline; synonymy of S. am-
biguus sensu Péringuey with S. natalensis requires further investigation
on the basis of cited localities, particularly Cape Town.
Distribution: Ranging along the SE Highveld: South Africa, Lesotho;
recorded from Grassland Biome in the NE to Upper Nama Karoo in
the SW; map marks the type localities of the two species recognised by
zur Strassen (S. basuto: black; S. natalensis: blue).
Locality data (mean ± SD, range): Altitude: 1 460 ± 268, 1 065–2 562 m;
annual rainfall: 455 ± 214, 221–958 mm; annual temperature (max. +
min. /2): 14.8 ± 1.5, 8.4–17.3°C (N=63).
Habitat: No quantitative assessment; in the Northern Cape: primarily
from Upper Karoo (39 out of 68 study sites) on sandy loam (8.8 per
sample), sand/loamy sand (3.3 per sample); DBRU collection records
biased to E localities: sand (4), sandy loam (4), sandy clay loam (3) in
grassland/pasture (8), shrubland (2). 5 mm

Food types: No quantitative assessment; DBRU collection records from

dung of cattle (12), sheep (2).
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to Mar.) with a bias to late summer (Feb., Mar.) in Upper Karoo.
Bioregions South Africa: Highland edges Mesic Highveld Grassland
(Gm), Drakensberg Grassland (Gd), Sub-Escarpment Grassland (Gs)
(Grassland Biome); N Upper Karoo (NKu) (Nama Karoo Biome).
Assessment rationale: EOO = 134 360 km2; molecular analysis would
be useful to support synonymy of S. natalensis with S. basuto; whether
considered as one or two species, currently assessed as Least Concern
(LC) on basis of limited habitat transformation over most of the large,
combined, highland range.
Conservation measures: Assessment of conservation status would be
improved by resolution of taxonomic issues and quantitative data on
ecological associations, particularly, possible effects of habitat transfor-
mation in E of range; not currently known from any protected area.

SCARABAEINI
622 SURICATA 6 (2020)

Scarabaeus bornemisszai
zur Strassen, 1980

No synonyms
Global: NT (see IUCN Red List – NT)

J A S O N D J F M A M J

Type locality: ‘St Lucia Estuary, coastal dune forest’ [KwaZulu-Natal,

South Africa].
Taxonomy: Accepted species.
Distribution: Restricted to the Maputaland Centre of Endemism in NE
KwaZulu-Natal (South Africa) and SE Mozambique.
Locality data (mean ± SD, range): Altitude: 12 ± 24, 0–85 m; annual
rainfall: 875 ± 98, 743–1 011 mm; annual temperature (max. + min.
/2): 22.0 ± 0.3, 21.4–22.4°C (N=23).
Habitat: In Maputo Special Reserve: mostly in coastal-fringing dune forest
(16.8/trap); uncommon in warmer, inland sand forest (0.2); virtually
absent from grassland on fossil lagoons (0.04); across a post-mining
vegetation succession at Richards Bay: present only in dense woodland
(9–20 yr) (15) and natural dune forest (71), absent from unshaded
shrub/grassland (0–8 yr) and open woodland (21–33 yr); DBRU col-
lection records all from sand (2) in forest (9).
Food types: No quantitative assessment; DBRU collection records from
human (1) and a mixture of human and cattle dung (1); sampled to pig
dung and a mixture of pig and cattle dung.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to May) with some activity in spring (Aug.).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Vegetation unit: Centred on deep sands, North-
ern Coastal Forest (FOz 7) (Forest Biome (FO) encompassed by Indian
Ocean Coastal Belt Biome (CB)).
Assessment rationale: EOO = 3 250 km2 comprising a narrow range
along 325 km of coastline; AOO much smaller, primarily restricted
5 mm
to coastal dune forest patches; in South Africa, 50 km of coastline is
mined for titanium-bearing sands; 185 km is protected as a World Her-
itage Site, which includes 68% of the vegetation unit (FOz 7); assessed
as Near Threatened (NT) although currently well protected.
Conservation measures: Conservation status is dependent on continuing
protection of existing reserves containing dune forest patches, including
iSimangaliso Wetland Park (World Heritage Site) (South Africa) and
Maputo Special Reserve (Mozambique); it would be useful to monitor
status of the species in the mined area N of Richards Bay, of which a
third is being permitted to regenerate as secondary forest; monitoring
of illegal clearance and use of forest patches would also be useful.

SCARABAEINI
SURICATA 6 (2020) 623

Scarabaeus caffer
(Boheman, 1857)

No synonyms
Global: LC (see IUCN Red List – DD)
Endemic: RSA, Eswatini

J A S O N D J F M A M J

Type locality: As Ateuchus caffer: ‘Caffraria tota passim’ [everywhere, all

SE South Africa].
Taxonomy: Accepted species.
Distribution: Centred in moist highlands along the upper edge of the
E escarpment, NE South Africa, with a distribution southwards that
descends to the coastline in the NE Eastern Cape; also reported from
Eswatini; Zimbabwe report requires validation.
Locality data (mean ± SD, range): Altitude: 1 214 ± 469, 0–2 005 m;
annual rainfall: 891 ± 130, 594–1 157 mm; annual temperature (max.
+ min. /2): 16.7 ± 2.3, 12.5–21.4°C (N=29).
Habitat: On finer-grained soils near Carolina: present in natural Themeda-
dominated grassland (5), absent from fallow crop fields and improved
Kikuyu pasture; at Wolkberg (1 400 m): much less abundant in forest
(9) than adjacent grassland (299); one DBRU collection record from
sandy clay loam in grassland.
Food types: At Wolkberg (1 400 m): equally abundant on dung of pig
(154) and cattle (154); one DBRU collection record from cattle dung.
Temporal activity: Diurnal flight activity, primarily in the summer rainy
season (Oct. to Apr.).
Bioregions South Africa: Montane or mistbelt vegetation units: E Mesic
Highveld Grassland (Gm), Sub-Escarpment Grassland (Gs) (Grassland
Biome); S Indian Ocean Coastal Belt Biome (CB); vegetation unit:
moist S edge, Soutpansberg Mountain Bushveld (SVcb 21) (Central
5 mm
Bushveld (SVcb), Savanna Biome).
Assessment rationale: EOO = 70 500 km2; probably threatened by pas-
ture improvement, cultivation and creation of forest plantations in
parts of its range; transformation: 0–70% in the N (E Gm), 3–50%
in the S (Gs); However, re-assessed as Least Concern (LC) due to the
large EOO, abundance in one reserve, and ready attraction to dung of
domestic livestock.
Conservation measures: Assessment of conservation status would be
improved by a quantitative survey of farmland and reserves across the
known range; an investigation of soil associations and its abundance in
transformed vs natural grassland would be particularly useful; protect-
ed and abundant in Lekgalameetse Nature Reserve, Wolkberg.

SCARABAEINI
624 SURICATA 6 (2020)

Scarabaeus cognatus
Péringuey, 1901

No synonyms
Global: DD
Endemic: Namibia

J A S O N D J F M A M J

Type locality: ‘Damaraland (Salem)’ [Namibia].

Taxonomy: Accepted species; one of three elongate species, until recently,
classified in the genus Drepanopodus Janssens, 1940.
Distribution: Currently known from two disjunct, arid, lowland, pop-
ulation centres below the W escarpment, one near the coastline in N
Namibia; one in the central Namib Desert in Namibia; record from the
Cape, South Africa, would represent confusion with the closely related
S. proximus Péringuey, 1901.
Locality data (mean ± SD, range): Altitude: 531 ± 244, 283–594 m;
annual rainfall: 77 ± 50, 29–157 mm; annual temperature (max. + min.
/2): 17.7 ± 0.6, 16.8–18.3°C (N=5).
Habitat: No quantitative assessment; collected from the gravel plains of
the Namib Desert where there is very sparse scrub cover.
Food types: No quantitative assessment; sampled to pig dung near Soli-
taire, central Namibia.
Temporal activity: Diurnal flight activity in the late summer rainy season
(Feb. to Apr.); also recorded after light rainfall in June.
Ecoregions Namibia: Below escarpment: W Namibian Savanna
Woodlands (AT1316), Namib Desert (AT1315), Kaokoveld Desert
(AT1310).
Assessment rationale: EOO = 18 345 km2; known only from two ultra-
arid desert centres; some sampled localities lie within an area scheduled
for mining; however, most of potential EOO used for grazing of live-
stock or under conservation; a poorly known species; therefore, assessed
as Data Deficient (DD).
Conservation measures: A quantitative survey of the full EOO, AOO
and ecological associations is required before an assessment of conser-
5 mm
vation status may be made; this should investigate the possibility of
continuous distribution from the S range limits in the Namib-Naukluft
Park along the protected Skeleton Coast to and beyond the northern-
most known locality on the Angolan border.

SCARABAEINI
SURICATA 6 (2020) 625

Scarabaeus convexus
(Hausmann, 1807)

= Ateuchus laevis Thunberg, 1818

Scarabaeus spretus
zur Strassen, 1962

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Ateuchus convexus: Not stated, but collector (Friedrich

Hesse) was based in Cape Town [Western Cape, South Africa]; A. lae-
vis: ‘majorem horum partem in capite bonae spei collegi’ [amongst
material mostly collected in the Cape of Good Hope, South Africa];
S. spretus: ‘Umgebung Kapstadt’ [surroundings of Cape Town].
Taxonomy: Currently, two accepted species, presumably, sharing type lo-
calities in the vicinity of Cape Town (S. convexus is described togeth-
er with Western Cape endemics: S. suri, S. rugosus (both Hausmann,
1807)); apparently no consistent differences between individuals iden-
tified as S. convexus (S Cape to Mpumalanga) or S. spretus (W and S
Cape); further study required.
Distribution: Coastal localities from Western Cape (winter rainfall) to
KwaZulu-Natal and highlands of E escarpment of Mpumalanga in the
NE (both summer rainfall).
Locality data (mean ± SD, range): N localities (8): Altitude: 1 424 ±
412, 864–2 274 m; annual rainfall: 828 ± 113, 684–1 022 mm; an-
nual temperature (max. + min. /2): 15.6 ± 2.2, 10.6–17.4°C (N=9). S
localities (14): Altitude: 234 ± 439, 0–1 542 m; annual rainfall: 510 ± 5 mm

204, 201–955 mm; annual temperature (max. + min. /2): 16.5 ± 2.2,
11.7–19.5°C (N=15).
Habitat: On Cape Peninsula (S. spretus): sampled on deep sands in dense,
low cover of Restio-dominated fynbos shrubland (2–3 yr since burn- S. convexus
ing) (183 individuals), also, open cover but higher profile fynbos (9–10
yr since burning) (84), few on nearby transformed habitat comprising
low, dense cover of Kikuyu grass (3); also recorded from sandy loam
and sandy clay loam in forest and highland grassland.
Food types: No quantitative assessment; sampled to dung of cattle (S. spre-
tus, S. convexus) and humans (S. convexus).
Temporal activity: Diurnal flight activity, mostly at similar levels of abun-
dance from spring till autumn in winter rainfall region on Cape Pen-
insula (Aug.–Apr.); active during summer in the bimodal and summer
rainfall regions (Oct.–Mar.).
Bioregions South Africa: In S: Southwest Fynbos (FO 2), Karoo Renos-
terveld (FO 9) (Fynbos Biome); Albany Thicket (AT), Indian Ocean
Coastal Belt (CB) biomes; In N: Sub-Escarpment Grassland (Gs), Me-
sic Highveld Grassland (Gm) (Grassland Biome).
Assessment rationale: EOO = 45 980 km2; both names used interchange-
ably for populations in S Cape; S. convexus used for NE populations
and S. spretus for those in extreme W Cape, even though both names
5 mm
are represented by types from the same general locality around Cape
Town; may comprise a single species; does not show marked season-
al adaptation to winter rainfall on Cape Peninsula where it is active
throughout summer; not currently threatened in SW or NE; assessed as
Data Deficient (DD) owing to nomenclatural problems.
S. spretus

SCARABAEINI
626 SURICATA 6 (2020)

Scarabaeus convexus & S. spretus (continued)

Conservation measures: Before an assessment may be made of conser-

vation status, nomenclatural problems need to be resolved; quantita-
tive data on ecological associations are also required; SW populations
protected in Cape of Good Hope Nature Reserve, NE populations in
uKhahlamba Drakensberg Park (World Heritage Site).

SCARABAEINI
SURICATA 6 (2020) 627

Scarabaeus costatus
(Weidemann, 1823)

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type localities: As Ateuchus costatus: ‘Prom. bon. sp.’ [Cape of Good

Hope, South Africa]; some references cite type locality as Orange
[River] whose lower reaches are within the known range.
Taxonomy: Accepted species; one of three elongate species, until recently,
included in the genus Drepanopodus Janssens, 1940.
Distribution: Hot, arid, deep sands centred on the SW Kalahari and areas
bordering the Namib Desert: NW South Africa, SW Botswana, W
Namibia.
Locality data (mean ± SD, range): Altitude: 1 023 ± 195, 283–1 356 m;
annual rainfall: 225 ± 78, 9–555 mm; annual temperature (max. + min.
/2): 18.9 ± 1.2, 14.2–21.2°C (N=73).
Habitat: No adequate quantitative assessment; in Northern Cape: mostly
restricted to SW Kalahari (5.34/trap), virtually absent from Bush-
manland and Upper Karoo; in SW Kalahari: strong bias to deep sand
(6.7, n=69), less on sandy loam (2.8, n=25), sandy clay loam (0, n=3);
DBRU collection records from sand (9) in open vegetation: grassland
(2), scrub (5), shrubland (2).
Food types: In Botswana: bias to dung of omnivores (pig: 42), less so to
ruminant herbivores (cattle: 19, sheep: 4), absent from monogastric
herbivore dung (elephant) and carrion (chicken livers); DBRU collec-
tion records on various dung types: human (1), horse (1), donkey (1),
cattle (3).
Temporal activity: Diurnal flight activity in the late summer rainy season
(Dec. to Apr.); sensitive to movement, fly away if disturbed.
Ecoregions Namibia, Botswana: Nama Karoo (AT1314), S Kalahari
Xeric Savanna (AT1309), Namibian Savanna Woodlands (AT1316),
Namib Desert (AT1315), Kaokoveld Desert (AT1310).
Bioregions South Africa: Centred on Kalahari Duneveld (SVkd), W
Eastern Kalahari Bushveld (SVk) (Savanna Biome); marginal occur-
rence in four adjoining bioregions.
Assessment rationale: EOO = 298 810 km2; widespread in an arid, little- 5 mm

transformed region used primarily for conservation or grazing of

domestic livestock on farm rangeland; AOO probably influenced by
specialisation to sandy soils, but quantitative support required; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil associations; protected in Kga-
lagadi Transfrontier Park (South Africa/Botswana), Sperrgebiet and
Namib-Naukluft national parks (Namibia).

SCARABAEINI
628 SURICATA 6 (2020)

Scarabaeus deludens
zur Strassen, 1961a

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Roodeplaat, 24 km nö. Pretoria, Transvaal, ca. 1 300 m’

[Roodeplaat, near Pretoria, Gauteng, South Africa].
Taxonomy: Accepted species; habitus close in appearance to Escarabaeus
satyrus (Boheman, 1860), but aedeagus differs radically.
Distribution: Finer-grained soils in open shrub/woodland savanna; N
South Africa, NE Botswana, N Namibia; unclear if distribution is truly
disjunct to E and W of Kalahari deep sands due to association with finer-
grained soils; citations from Zimbabwe and, particularly, Tanzania re-
quire validation.
Locality data (mean ± SD, range): Altitude: 795 ± 432, 55–1 588 m;
annual rainfall: 610 ± 136, 314–900 mm; annual temperature (max. +
min. /2): 20.4 ± 2.0, 16.5–23.6°C (N=40).
Habitat: No quantitative assessment; limited DBRU collection records
from finer-grained soils, sandy clay loam (2), clay (2), in mopane shrub-
land (2).
Food types: No quantitative assessment; limited DBRU collection records
from dung of elephant (3), cattle (1).
Temporal activity: Flight activity during darkness in the summer rainy
season (Oct. to Apr.); attracted to light.
Ecoregions Namibia: S Angolan Mopane Woodlands (AT0702), margins
of two other ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); marginal in three other bioregions.
Assessment rationale: EOO = 186 900 km2 (possibly underestimated);
AOO probably smaller due to association with finer-grained soils al-
though quantitative support is required; assessed as Least Concern (LC)
5 mm owing to widespread occurrence in farmland and reserves, although
essentially Data Deficient (DD) owing to paucity of ecological asso-
ciation data.
Conservation measures: Assessment of conservation status would be
much improved by quantitative data on ecological associations, partic-
ularly testing for soil type specialisation; protected in Kruger National
Park (South Africa), Etosha National Park (Namibia).

SCARABAEINI
SURICATA 6 (2020) 629

Scarabaeus ebenus
(Klug, 1855)

= Scarabaeus politifrons Fairmaire, 1887

= Scarabaeus sericeipennis Fairmaire, 1887
= Scarabaeus glabratus Kolbe, 1895

Scarabaeus interstitialis
(Boheman, 1857)

No synonyms
Global: DD (for S. ebenus, see IUCN Red List – DD)

J A S O N D J F M A M J

Type localities: As Ateuchus ebenus: ‘Inhambane’ [central coast of Mo-

zambique]; S. politifrons: ‘Ouebbi’ [Somalia]; S. sericeipennis: ‘Kibanga’
[Democratic Republic of the Congo (DRC)]; S. glabratus: ‘Witu’ [NE
Kenya coast]. As Ateuchus interstitialis: ‘Caffraria Interiore’ [South Af-
rican interior].
Taxonomy: Both accepted species, but treated together as aedeagi are
identical and morphological characteristics represent a cline from
the SE coastline onto the NW uplands of South Africa; synonyms of
S. ebenus and records of both species require re-assessment.
Distribution: Typical S. ebenus are characterised by scattered fine pits on
prothoracic disc and elytra: NE to S coast (Mozambique), Zambezi
Valley (Zimbabwe, Botswana) (black squares); validation required for
Tanzania, Namibia records and Somalia, Kenya synonyms of S. ebenus;
Democratic Republic of the Congo (DRC) synonym and Ethiopia re-
cord are probably errors; on Pumbe Sands, NE lowlands, South Africa 2 mm
(arrowed): pits on the disc are largely similar to S. ebenus; however,
to the S: pits on the disc are often larger and grade into typical S. in-
terstitialis on the NW highlands, South Africa (red squares), showing
dense, large pits on the disc and scattered pits on the elytra; Tanzania, S. ebenus
Mozambique records of S. interstitialis require validation.
Locality data (mean ± SD, range): NE lowlands: Altitude: 301 ± 239,
0–1 000 m; annual rainfall: 672 ± 159, 383–971 mm; annual tempera-
ture (max. + min. /2): 22.5 ± 1.2, 20.9–25.2°C (N=20) (black squares).
NW uplands: Altitude: 1 151 ± 315, 323–1 510 m; annual rainfall:
584 ± 93, 403–765 mm; annual temperature (max. + min. /2): 18.8 ±
1.9, 16.2–23.0°C (N=20) (red squares).
Habitat: No quantitative assessment; ‘S. ebenus’ recorded from deep sand
in open woodland, S. interstitialis from sandy loam in grassland and
open woodland.
Food types: No quantitative assessment; ‘S. ebenus’ sampled to omnivore
dung (pig) at Phalaborwa (4) and Chobe National Park (3).
Temporal activity: Diurnal flight activity in the summer rainy season
(Nov. to Feb.).
Ecoregions Botswana, Zimbabwe: S. ebenus: Zambezian and Mopane
Woodlands (AT0725).
Bioregions South Africa: ‘S. ebenus’: centred on Lowveld (SVl) (Savanna
Biome); S. interstitialis: centred on Central Bushveld (SVcb) (Savanna
Biome), N Dry Highveld Grassland (Gh) (Grassland biome). 2 mm
Assessment rationale: EOO = 119 975 km2 (S. interstitialis: NW up-
lands, white), EOO = 422 685 km2 (S. ebenus: NE lowlands, black);
identity and distribution uncertain; possible associations of ‘S. ebenus’
S. interstitialis

SCARABAEINI
630 SURICATA 6 (2020)

Scarabaeus ebenus & S. interstitialis (continued)

with E coastal sands and ‘S. interstitialis’ with sandy loam, but quan-
titative support required; probably no serious threats despite apparent
low population density over much of their ranges; however, taxonomic
uncertainty (one or two species) and poor support for ecological differ-
ences demands an assessment as Data Deficient (DD).
Conservation measures: Taxonomic revision is required before conserva-
tion status can be assessed; quantitative data on ecological associations
is also required; in South Africa, protected in Kruger National Park
(‘S. ebenus’), Abe Bailey, Borakalalo and Tswaing nature reserves (‘S.
interstitialis’).

SCARABAEINI
SURICATA 6 (2020) 631

Scarabaeus funebris
(Boheman, 1857)

= Scarabaeus funebris pretoriensis Janssens, 1940b

Global: DD
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type localities: As Ateuchus funebris: ‘Caffraria interiore’ [interior of SE

South Africa]; S. funebris pretoriensis: Not explicitly stated.
Taxonomy: Accepted species.
Distribution: Validated records from one coastal and two upland centres:
coastal KwaZulu-Natal and drier W Highveld, South Africa, N plateau,
Namibia; also reported from Zimbabwe, Botswana.
Locality data (mean ± SD, range): Altitude: 1 135 ± 444, 55–1 559 m;
annual rainfall: 538 ± 146, 304–743 mm; annual temperature (max. +
min. /2): 18.9 ± 2.6, 15.5–23.0°C (N=22).
Habitat: No quantitative assessment; DBRU records mainly from finer-
grained soils: sand (2), sandy loam (5), sandy clay loam (6) in grassland
(3), scrub/shrubland (3), open woodland (7).
Food types: No quantitative assessment; DBRU records from dung of
cattle (6), wildebeest (1), elephant (1), zebra (3), springbok (1).
Temporal activity: Diurnal flight activity during the summer and late
summer rainy seasons (Oct. to Apr.).
Ecoregions Namibia: Angolan Mopane Woodlands (AT0702).
Bioregions South Africa: Dry Highveld Grassland (Gh) (Grassland
Biome); NE Upper Karoo (NKu) (Nama Karoo Biome); E Eastern
Kalahari Bushveld (SVk), S Central Bushveld (SVcb), S Lowveld (SVl)
(Savanna Biome).
Assessment rationale: EOO = 104 390 km2; wide range, but infrequent-
ly recorded; biogeographically and ecologically poorly known; AOO
might be limited by soil type association; most DBRU collection re-
cords (63%, n=10/16) are from two game reserves; assessed as Data 2 mm
Deficient (DD).
Conservation measures: A quantitative survey needs to determine if the
AOO is truly disjunct as is likely, or, merely a collection artefact; the
ecological associations also need to be quantified; the frequency of
game reserve records may be related to association with pellet-dropping
herbivores and this needs to be tested; protected in uMkhuze Game
Reserve, Abe Bailey Nature Reserve (South Africa), Etosha National
Park (Namibia).

SCARABAEINI
632 SURICATA 6 (2020)

Scarabaeus galenus
Westwood, 1847

= Ateuchus paradoxus Boheman, 1857

= Sebasteos adelphus Kolbe, 1895
Global: LC (see IUCN Red List – LC)*

J A S O N D J F M A M J

Type localities: As Scarabaeus (Sebasteos) galenus: ‘Africa meridionali.....in

the hilly country lying between 25° and 26° south lat. and 27° and 28°
east long.’ [Gauteng region, South Africa]; A. paradoxus: ‘prope fluvium
Limpopo’ [near River Limpopo, southern Africa]; S. adelphus: ‘Mkatta
fluss in Usagara’ [Mkatta River, Tanzania].
Taxonomy: Accepted species.
Distribution: Widespread in mostly drier, lower-altitude, sandy savannas
of SE and S Central Africa: South Africa, N Namibia, N Botswana,
Zimbabwe, Tanzania; also reported from Zambia.
Locality data (mean ± SD, range): Altitude: 750 ± 438, 55–1 495 m;
annual rainfall: 630 ± 90, 436–850 mm; annual temperature (max. +
min. /2): 20.6 ± 1.7, 16.9–23.6°C (N=43).
Habitat: No quantitative assessment; DBRU collection records biased to
sandier soils: sand (6), sandy loam (5), sandy clay loam (1), and partial-
ly shaded vegetation: grassland (2), open woodland (9).
Food types: At Phalaborwa: strong bias to dung of ruminant herbivores
(cattle: 21) rather than monogastric herbivores (elephant: 1) and omni-
vores (pig: 4); supported by DBRU collection records from cattle (8),
wildebeest (1), buck (1), rhinoceros (2); but, prominence of two central
clypeal teeth and observations of association with impala middens may
indicate ready use of dung dropped as pellets.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.); at Phalaborwa: primarily active on a warm day soon
after heavy rainfall (24) than on a hot dry (1) or warm cloudy day (1)
after light rainfall.
Ecoregions Namibia, Botswana, Zimbabwe: Angolan Mopane Wood-
lands (AT0702), Zambezian and Mopane Woodlands (AT0725),
Southern Miombo Woodlands (AT0719).
5 mm
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 699 090 km2; AOO would likely be small-
er owing to bias to woody vegetation on sandy soils although quantita-
tive support is required; widely recorded in both farmland and reserves
although clearance of woodland would probably be detrimental to
population density; in South Africa, transformation 0–58% (SVl),
2–49% (SVcb), 0–20% (SVmp); assessed as Least Concern (LC) on
basis of wide range. *IUCN Red List for S. galenus also includes data
for S. vicinus Janssens, 1940b.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly
relative bias to woodland on sandy soils and relative use of antelope
pellets compared to other dung types; protected in Kruger National
Park (South Africa).

SCARABAEINI
SURICATA 6 (2020) 633

Scarabaeus geminogalenus
Davis & Deschodt, 2017

= Sebasteos westwoodi (Harold, 1869a) sensu Péringuey, 1908

Global: LC

J A S O N D J F M A M J

Type localities: S. geminogalenus: ‘ST. LUCIA ESTUARY, Natal.’

[KwaZulu-Natal, South Africa]; S. westwoodi sensu Péringuey: cited
from ‘Transvaal (Soutpansberg)’ [Limpopo Province, South Africa].
Taxonomy: Accepted species; recently re-described as new after misidenti-
fication as Scarabaeus westwoodi Harold, 1869a, and synonymy with S.
galenus Westwood, 1847.
Distribution: Primarily Limpopo River Valley and coastal deep sands: SE
Mozambique, NE KwaZulu-Natal in South Africa.
Locality data (mean ± SD, range): Altitude: 380 ± 416, 0–1 331 m;
annual rainfall: 652 ± 176, 455–971 mm; annual temperature (max. +
min. /2): 21.8 ± 1.6, 17.1–23.4°C (N=16).
Habitat: No quantitative assessment of soil type, but many records from
sand patches or areas of deep sands; on deep sands in Maputo Elephant
Reserve: mostly in grassland (15) as opposed to dune forest (1); sup-
ported by DBRU collection records in grassland (4) and thicket (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (1), rhinoceros (1).
Temporal activity: Diurnal flight activity with long seasonal activity due
to occurrence in the warm coastal lowlands (Aug. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); N Indian Ocean Coastal Belt Biome
(CB).
Assessment rationale: EOO = 40 770 km2; limited data suggest the AOO
would be much smaller, centred in open vegetation on deep sand al-
though quantitative support for soil associations is lacking; currently 5 mm
assessed as Least Concern (LC) on the basis of fairly wide range in open
vegetation at a number of locations, of which the E records are entirely
within reserves.
Conservation measures: Further quantitative data on ecological associa-
tions is required to support the assessment of conservation status, par-
ticularly soil and food association data; protected in Kruger National
Park, iSimangaliso Wetland Park (World Heritage Site) (South Africa),
Maputo Special Reserve (Mozambique).

SCARABAEINI
634 SURICATA 6 (2020)

Scarabaeus goryi
(Castelnau, 1840)

= Ateuchus profanus Boheman, 1857

= Scarabaeus nepos Fairmaire, 1884b
= Scarabaeus sacer var. laevigatus Kolbe, 1887
= Scarabaeus pacatus Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As Ateuchus goryi: ‘Sénégal’; A. profanus: ‘Portum Natal-

ense’ [Durban, South Africa]; S. nepos: ‘Zanzibar’; S. sacer var. laevigatus:
‘Chinchoxo’ [Angola]; S. pacatus: ‘Southern Rhodesia’ [Zimbabwe].
Taxonomy: Accepted species; a very close Palaearctic/Oriental relative is
classified under Scarabaeus s. str.
Distribution: Widespread in sandy, moist, open savanna regions from S,
E and W Africa: Senegal, Côte d’Ivoire, Nigeria, Democratic Republic
of the Congo (DRC), Kenya, Tanzania, Angola, Zimbabwe, Namibia,
Botswana, Mozambique, South Africa; also reported from Gambia,
Guinea, Togo, Gabon, Uganda, Malawi; Sudan and Somalia records
may be Scarabaeus isidis (Castelnau, 1840), a close, arid-region relative.
Locality data (mean ± SD, range): Altitude: 807 ± 480, 0–1 503 m; an-
nual rainfall: 593 ± 191, 171–1 010 mm; annual temperature (max. +
min. /2): 20.5 ± 1.7, 15.6–23.4°C (N=136).
Habitat: In open woodland at Leeuwfontein Nature Reserve: strong bias
to sand (43) rather than sandy loam (11) or stony sandy loam (4);
on deep sand in Maputo Special Reserve: abundant in both grassland
(27) and sand forest patches (small: 29, medium: 59, large: 40); DBRU
collection records also indicate a deep sand specialist and vegetation
generalist: sand (21), sandy loam (2) in grassland (13), shrubland (4),
open woodland (7), forest (3).
5 mm
Food types: In Botswana: bias to omnivore (pig: 14) rather than herbivore
dung (cattle: 5, sheep: 1, elephant: 1); DBRU collection records reflect
commonly sampled dung types: human/cattle mixture (2), cattle (21),
wildebeest (1), warthog (1).
Temporal activity: Flight activity in darkness primarily during the sum-
mer rainy season (Oct. to May); also Aug. at the warmer coastline;
attracted to light.
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: Deep
sands: Namibian Savanna Woodlands (AT1316), Kalahari Xeric
Savanna (1 309), Angolan Mopane Woodlands (AT0702), Kalaha-
ri Acacia-Baikiaea Woodlands (AT0709), Zambezian and Mopane
Woodlands (AT0725), Southern Miombo Woodlands (AT0719),
Southern African Bushveld (AT0717), Maputaland Coastal Forest
Mosaic (AT0119).
Bioregions South Africa: Centred on Eastern Kalahari Bushveld (SVk),
Central Bushveld (SVcb), Lowveld (SVl) (Savanna Biome), Indian
Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 2 990 240 km2; AOO rather more restrict-
ed due to specialisation to deep sands, but still widespread in open or
shaded vegetation from drier SW Africa to moist tropical savanna where
it may show a bias to omnivore dung although it is also readily attract-
ed to the dung of domestic livestock; assessed as Least Concern (LC).
Conservation measures: None recommended; protected in various re-
serves including Tembe Elephant Park (South Africa), Bicuar National
Park (Angola).

SCARABAEINI
SURICATA 6 (2020) 635

Scarabaeus heqvisti
zur Strassen, 1962

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Magalies-berge, Transvaal’ [South Africa].

Taxonomy: Accepted species.
Distribution: To date, recorded only along the E–W trending Magalies-
berg range at the N edge of the South African Highveld on the transi-
tion between grassland and bushveld.
Locality data (mean ± SD, range): Altitude: 1 173 ± 202, 1 043–1 405 m;
annual rainfall: 652 ± 42, 625–701 mm; annual temperature (max. +
min. /2): 18.8 ± 1.0, 17.6–19.4°C (N=3).
Habitat: No adequate quantitative assessment; in Ezemvelo (now Telperi-
on) Nature Reserve, Gauteng: recorded mainly in upland grassland on
a patch of very sandy soils (1 419 m); rare in grassland on sandy soils at
1 338 m; not recorded on sandy loam or in open woodland on koppies.
Food types: No quantitative assessment; sampled to composite baits of
pig and cattle dung.
Temporal activity: Diurnal flight activity in the summer rainy season
(Oct. to May); absence in mid-summer undoubtedly a collecting ar-
tefact.
Bioregions South Africa: Vegetation unit: Rand Highveld Grassland
(Gm 11) (Mesic Highveld Grassland (Gm), Grassland Biome); Vegeta-
tion unit: Moot Plains Bushveld (SVcb 8) (Central Bushveld (SVcb),
Savanna Biome).
Assessment rationale: EOO = 305 km2; although under protection in at
least one reserve where it may show highly localised abundance after
rainfall, the extremely small known EOO in few locations on a single
mountain system would qualify this species for a IUCN threat category
of at least Near Threatened (NT) or Vulnerable (VU), particularly as it 5 mm
is centred in an area where vegetation units are 28–50% transformed
by cultivation, urbanisation or mining; as the full EOO and AOO are
currently unknown, assessed as Data Deficient (DD).
Conservation measures: To assess the conservation status of this species,
a quantitative survey is required to determine its full EOO and the
ecological associations that restrict its AOO; the survey should focus on
patches of sandy and other soil types in the Magaliesberg and nearby
mountain blocks, and also determine possible effects of pasture mod-
ification; currently protected in Telperion Nature Reserve where it is,
nevertheless, abundant on sandy soils after rainfall.

SCARABAEINI
636 SURICATA 6 (2020)

Scarabaeus hottentorum
Péringuey, 1901

= Sebasteos procles Kolbe, 1908

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: S. hottentorum: ‘Cape Colony (Namaqualand, Barkly

West)’ [South Africa]; S. procles: ‘Klein-Namaland’ [Namaqualand].
Taxonomy: Accepted species; of the type localities: Namaqualand in NW
South Africa is considered correct, Barkly West in central South Africa
is considered an error; classified in the genus, Ateuchetus Bedel, 1892,
in the 2019 version of Catalogue of Life.
Distribution: Namaqualand, South Africa; recorded along the upper edge
of the W escarpment in the N as well as coastal regions in the S.
Locality data (mean ± SD, range): Altitude: 564 ± 323, 99–1 116 m;
annual rainfall: 126 ± 21, 93–154 mm; annual temperature (max. +
min. /2): 17.6 ± 0.7, 16.2–18.3°C (N=7).
Habitat: No quantitative assessment; limited DBRU collection records
from sand (1), sandy loam (1) in scrub/succulent Karoo (2).
Food types: No quantitative assessment; limited DBRU collection records
from human (1) and cattle dung (1).
Temporal activity: Diurnal flight activity recorded in spring (Aug., Sept.)
towards the end of the winter rainy season.
Bioregions South Africa: Centred on Namaqualand Hardeveld (SKn)
(Succulent Karoo Biome).
Assessment rationale: EOO = 3 335 km2; qualifies for an IUCN threat
category of at least Vulnerable (VU) on the basis of small range
(< 5 000 km2) and few recorded localities; however, its arid EOO prob-
ably largely equates to its AOO, which is little transformed (0–6%) as
it is used primarily for grazing of farm livestock; the effects of trans-
formation for cultivation of winter wheat are unclear; currently poorly
known and assessed as Data Deficient (DD).
Conservation measures: Before conservation status may be assessed, a
5 mm
quantitative survey is required to better determine the EOO, AOO
and ecological associations; not currently known from any reserves, but
probably not threatened owing to limited regional transformation.

SCARABAEINI
SURICATA 6 (2020) 637

Scarabaeus karae
Davis & Deschodt, 2017

No synonyms (but see Taxonomy)

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Abe Bailey NR, Gauteng, S26°17’–21’ E027°16’–18’,

1 496 m–1 544 m’ [South Africa].
Taxonomy: Accepted species; newly described, smaller-bodied, close rela-
tive of Scarabaeus funebris (Boheman, 1857); aedeagus quite different;
further new species, Scarabaeolus orientalis (Zídek & Pokorný, 2018),
is considered an unpublished synonym of Scarabaeus karae (see Pre-
amble).
Distribution: Known from a few points at the type locality (Abe Bailey
Nature Reserve) and recently recorded as a rarity at two other localities,
respectively, on a farm and near a rehabilitating mined area along the N
edge of the Highveld, South Africa.
Locality data (mean ± SD, range): Altitude: 1 447 ± 110, 1 284–1 521 m;
annual rainfall: 676 ± 20, 661–705 mm; annual temperature (max. +
min. /2): 16.9 ± 0.8, 16.0–17.8°C (N=4).
Habitat: No quantitative assessment; sampled on sandy loam in natural
grassland (Abe Bailey) or on sand, either in degraded natural grassland
(farm) or a natural grassland patch surrounded by areas of mining and
rehabilitation.
Food types: No quantitative assessment; sampled to composite baits of
pig and cattle dung.
Temporal activity: Diel flight periodicity unknown, but presumably di-
urnal based on black pilosity; recorded in the summer rainy season
(Dec., Mar.).
Bioregions South Africa: Vegetation units: Carletonville Dolomite Grass-
land (Gh 15) (Dry Highveld Grassland (Gh)); N edge of Eastern High-
veld Grassland (Gm 12) (Mesic Highveld Grasslnd (Gm)) (Grassland
Biome). 2 mm
Assessment rationale: EOO = 3 080 km2 (presumably underestimated);
only known from natural grassland on sand and sandy loam in vege-
tation units that are 25–44% transformed by cultivation, urbanisation
or mining; otherwise ecologically poorly known; S. karae would qualify
for an IUCN threat category of at least Endangered (EN) on the basis
of records from < 5 locations within an EOO < 5 000 km2; however, the
EOO is presumably underestimated and a collection artefact; therefore,
S. karae is currently assessed as Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative sur-
vey is required in drier grasslands of the NW Highveld; this needs to
determine the full EOO, AOO and ecological associations, particularly
if habitat transformation is a detrimental influence on the AOO (min-
ing, cultivation and pasture improvement); protected in Abe Bailey
Nature Reserve (South Africa).

SCARABAEINI
638 SURICATA 6 (2020)

Scarabaeus piliventris
zur Strassen, 1962

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Saldanha Bay, Hopefield Distrikt’ [Western Cape, South

Africa].
Taxonomy: Accepted species.
Distribution: SW coastal lowlands and upper edge of SW escarpment,
Western Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 295 ± 299, 2–705 m; annual
rainfall: 181 ± 44, 145–269 mm; annual temperature (max. + min. /2):
17.8 ± 1.3, 15.5–19.2°C (N=6).
Habitat: No quantitative assessment; one DBRU collection record from
sandy loam in ‘herbaceous pasture’ cleared of natural shrubland; also
recorded in coastal bioregions characterised by deep sands.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Diurnal flight activity recorded in late winter (July)
and spring (Sept.) towards the end of the winter rainy season.
Bioregions South Africa: Primarily bioregions either side of the transition
between the Fynbos (F) and Succulent Karoo Biomes (SK): Western
Strandveld (F 11), Northwest Fynbos (FO 1), Namaqualand Sandveld
(SKs), Knersvlakte (SKk), W Trans-Escarpment Succulent Karoo (SKt).
Assessment rationale: EOO = 3 305 km2; the small known range and low
number of localities in an area subject to 10–70% transformation by
cultivation, urbanisation and mining would qualify this species for an
IUCN threat category of at least Vulnerable (VU); however, represent-
ed by relatively few museum specimens and poorly known; assessed as
2 mm Data Deficient (DD).
Conservation measures: A quantitative survey is required to determine
the full EOO, AOO and ecological associations before an assessment
may be made of conservation status; the effect of habitat transforma-
tion needs to be studied as one of six known localities (Nieuwoudtville)
was in an area widely transformed into pasture; not currently known
from any reserve.

SCARABAEINI
SURICATA 6 (2020) 639

Scarabaeus plausibilis
Péringuey, 1892

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: Not stated.

Taxonomy: Accepted species, but status requires validation in comparison
to close, previously described relatives, namely: Scarabaeus sulcipennis
(Quedenfeldt, 1888) and Scarabaeus poggei (Waterhouse, 1890), both
cited from inexact type localities in S Democratic Republic of the
Congo (DRC).
Distribution: S. plausibilis cited from Angola plus Ovamboland in Na-
mibia; overlapping citations for S. poggei (Katanga in Democratic
Republic of the Congo (DRC), Lunda in Angola) and S. sulcipennis
(Kassai in Democratic Republic of the Congo (DRC), Lunda in An-
gola, Namibia).
Locality data: No precise data on map other than a suspect locality ’30 m
W of Windhoek’ [central Namibian uplands] (black square); grey
shaded panel estimates S edge of EOO with respect to citation from
Ovamboland, inexact locality along ‘Okavango R’, and record from
Liuwa Plains, SW Zambia (S14.6433 E22.6264) (to N of E end of
grey panel).
Habitat: No quantitative assessment; citation from Ovamboland and
records from Okavango River and Liuwa Plains may indicate an associ-
ation with deep sands on floodplain grassland.
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Diel flight periodicity unknown, but probably diur-
nal based on black pilosity; recorded during the summer rainy season
(Nov.).
Ecoregions Zambia: A single exact record on Liuwa Plains: Western Zam-
bezian Grasslands (AT0724).
Assessment rationale: EOO unclear but at least from Ovamboland to
Liuwa Plains (grey panel); AOO would be much smaller if specialised
to moist flood plain grasslands; assessed as Data Deficient (DD) owing
to taxonomic uncertainty and poor ecological knowledge.
5 mm
Conservation measures: A survey of N Namibia, E Angola, S Democratic
Republic of the Congo (DRC) and SW Zambia is required to obtain
further material and resolve questions on validity of nomenclature: how
many species, three, two or only one; survey should determine the full
EOO, AOO and provide quantitative data on ecological associations;
no current records known from reserves.

SCARABAEINI
640 SURICATA 6 (2020)

Scarabaeus proboscideus
(Guérin-Meneville, 1844)

= Ateuchus modestus Boheman, 1860

= Scarabaeus rostratus Péringuey, 1888
Global: LC

J A S O N D J F M A M J

Type localities: As Ateuchus proboscideus: ‘le Cap de Bonne-Espérance’

[Cape of Good Hope, South Africa]; A. modestus: ‘prope lacum Ngami’
[near Lake Ngami, Botswana]; S. rostratus: ‘Spektakel, Namaqualand...
Kenhardt and Bushmanland’ [Northern Cape, South Africa].
Taxonomy: Accepted species; one of two species, until recently, included
in the genus Neateuchus Gillet, 1911.
Distribution: Widespread on deep sands from the arid SW Kalahari to
the SW coast and N along inland edge of Namib Desert to the Kao­
koveld; South Africa, Botswana, Namibia, Angola.
Locality data (mean ± SD, range): Altitude: 808 ± 424, 0–1 782 m; an-
nual rainfall: 219 ± 95, 29–525 mm; annual temperature (max. + min.
/2): 18.8 ± 1.6, 14.5–22.3°C (N=222).
Habitat: No adequate quantitative assessment; in Northern Cape: re-
corded primarily in Kalahari (n=91 sites) and N Bushmanland (n=23);
in Western Cape: sampled primarily from shrubland on deep sand
(Modderrivier: 10, Pampoenvlei sand: 52, Pampoenvlei sandy loam:
2); DBRU collection records exclusively from deep sands (14) in open
woodland, shrubland or grassland (3), scrub (3), shrubland (7), open
woodland (1).
Food types: On deep sands in Botswana: most strongly attracted to dung
of pellet-dropping sheep (1 182), also sampled in abundance to other
dung types: pig (252), elephant (112), cattle (163), rare on carrion
(chicken livers: 1); DBRU collection records from dung of cattle (11)
and donkey (1).
Temporal activity: Flight activity in darkness; observed flying up to 23:00
in the Kalahari at 26°C; attracted to light; in SW winter rainfall re-
gion: active in warmer, dry, early summer (Oct., Nov.) tailing off into
5 mm
mid-summer (Dec. to Feb.); in NE: primarily active during late sum-
mer rains.
Ecoregions Namibia, Botswana: Centred on deep sands of Succulent
Karoo (AT1322), Namibian Savanna Woodlands (AT1316), Kalaha-
ri Xeric Savanna (AT1309), marginal occurrence in three adjoining
ecoregions.
Bioregions South Africa: Centred on deep sands of Eastern Kalahari
Bushveld (SVk), Kalahari Duneveld (SVkd), (Savanna Biome); N
Bushmanland (NKb) (Nama Karoo Biome); Namaqualand Sandveld
(SKs) (Succulent Karoo Biome); Western Strandveld (F 11) (Fynbos
Biome).
Assessment rationale: EOO = 502 100 km2; observed to be a sand spe-
cialist although poorly supported by available data; AOO would be
similar to EOO owing to large extent of deep sands across its wide
range that is little transformed and used primarily for the grazing of
domestic livestock or conservation; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data to support observed specialisation to deep
sands in open vegetation; protected in Central Kalahari Game Reserve
(Botswana), Kgalagadi Transfrontier Park, coastal portion of Namaqua
National Park (South Africa) and Sperrgebiet National Park (Namibia).

SCARABAEINI
SURICATA 6 (2020) 641

Scarabaeus proximus
Péringuey, 1901

No synonyms
Global: LC
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: ‘Cape Colony (Namaqualand), Port Nolloth’ [South Af-

rica].
Taxonomy: Accepted species; one of three species, until recently, included
in the genus Drepanopodus Janssens, 1940.
Distribution: Centred on arid sands to the S and N of the Orange River:
South Africa, Namibia.
Locality data (mean ± SD, range): Altitude: 274 ± 243, 1–940 m; annual
rainfall: 62 ± 22, 32–115 mm; annual temperature (max. + min. /2):
15.9 ± 0.8, 14.5–17.6°C (N=24).
Habitat: No quantitative assessment; all South African records (9) record-
ed from areas of deep sands in Succulent Karoo.
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Presumably diurnal flight activity on the basis of black
pilosity; recorded in autumn (May) and from late winter to early sum-
mer (July to Dec.) at the end of the winter rainy season and early dry
season.
Ecoregions Namibia: Primarily Succulent Karoo (AT1322), one record at
extreme S margins of Namib Desert (AT1315).
Bioregions South Africa: Only in arid N Namaqualand Sandveld (SKs)
(Succulent Karoo Biome).
Assessment rationale: EOO = 16 030 km2; observed to be a soil specialist,
therefore, AOO would be restricted to areas of arid deep sands that are
extensive within its limited lowland EOO; little protection in South
Africa although transformation of SKs is only 0–10% primarily by
mining along the N coastline; however, as the 50% of the EOO that 5 mm
occurs in Namibia is officially conserved, S. proximus is assessed as Least
Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations; in South Africa,
only the extreme S range limits are protected in Namaqua National
Park; however, most of its range in Namibia is protected in Sperrgebiet
National Park.

SCARABAEINI
642 SURICATA 6 (2020)

Scarabaeus rixosus
Péringuey, 1901

No synonyms
Global: DD (see IUCN Red List – DD)

J A S O N D J F M A M J

Type locality: ‘Southern Rhodesia (Middle Limpopo)’ [Zimbabwe].

Taxonomy: Accepted species; known only by the type specimen recently
rediscovered in the reference collection of the National Collection of
Insects, South Africa (Deschodt et al. 2017); not previously marked as
a type, but clearly different to any other southern African Scarabaeus
spp.; identified as S. rixosus from original description, with monofid
spur on mentum as the defining diagnostic character; one of two spe-
cies, until recently, included in the genus Neateuchus Gillet, 1911.
Distribution: Essentially unknown; middle (cited type locality) to upper
Limpopo Valley (etiquette label on putative type specimen in Pé-
ringuey’s handwriting); cited from Zimbabwe along border with South
Africa; unclear if it would occur in South Africa.
Locality data (range): Estimated approximations; altitude: 560–830 m;
annual rainfall: 290–330 mm; annual temperature (max. + min. /2):
23.0–24.5°C.
Habitat: No quantitative assessment; unknown; middle Limpopo Valley is
an isolated island of low rainfall with expanses of loam and loamy sands
on Zimbabwean side of border; South African side of border more het-
erogeneous although small patches of deep sands occur; putative close
relative, S. proboscideus Guérin-Meneville, 1844, is a sand specialist
(relationship based on shared character of spur on mentum).
Food types: No quantitative assessment; unknown.
Temporal activity: Unknown.
Ecoregions Zimbabwe: Speculative; Limpopo Valley extension of Zambe-
zian and Mopane Woodlands (AT0725).
Bioregions South Africa: Speculative; Mopane (SVmp) (Savanna Biome).
Assessment rationale: EOO and AOO unknown; no ecological data
5 mm
whatsoever; only known by the single type specimen described over a
century ago and with no subsequent records; could be truly rare and
endangered or a collection artefact; currently assessed as Data Deficient
(DD).
Conservation measures: A quantitative survey is required to determine
EOO, AOO and ecological associations before an assessment of conser-
vation status may be made; this should be conducted across the range
of habitats found from the middle to the upper Limpopo Valley, partic-
ularly loamy sands (Zimbabwe) and deep sand patches (South Africa);
not known to be protected in any reserve.

SCARABAEINI
SURICATA 6 (2020) 643

Scarabaeus rugosus
(Hausmann, 1807)

= Ateuchus convalescens Wiedemann, 1823

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Ateuchus rugosus: Not stated; A. convalescens: ‘Prom. bon.

sp.’ [Cape of Good Hope, South Africa].
Taxonomy: Accepted species.
Distribution: Except for a few inland localities that require validation,
restricted to dry west coastal sands in the winter rainfall region of South
Africa.
Locality data (mean ± SD, range): Altitude: 98 ± 100, 0–636 m; annual
rainfall: 186 ± 78, 48–393 mm; annual temperature (max. + min. /2):
17.2 ± 1.0, 15.2–18.7°C (N=86).
Habitat: In Western Cape: sampled primarily from shrubland on deep
sand (Modderrivier: 307, Geelbek: 397, Pampoenvlei: 71); uncommon
at adjacent sites: shrubland on sandy loam at Pampoenvlei (2), pasture
created by clearance of shrubs on deep sand at Geelbek (12); DBRU
collection records exclusively from deep sand (14) in scrub/shubland
(9); single records in grassland and a fallow crop field.
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (15) and horse (1).
Temporal activity: Diurnal flight activity in sunshine, especially after
rainfall as long as temperatures remain sufficiently high; on the SW
coast: flight activity commences at about 21°C with a midday peak
in the cooler early spring or a 09:00–10:00 peak in the warmer later
spring; seasonal peaks of sclerotised adults in spring (July to Oct.) and
both sclerotised and teneral adults in autumn (Mar. to May).
Bioregions South Africa: Centred on Namaqualand Sandveld (SKs) (Suc-
culent Karoo Biome), West Strandveld (F 11) (Fynbos Biome); few
localities in vegetation units of Sand Fynbos (FFd), (Northwest Fynbos 5 mm

(FO 1), Fynbos Biome).

Assessment rationale: EOO = 11 000 km2; N part of range has been
little transformed (3–10%); S part of range in Western Strandveld is
from 25–50% transformed by cultivation and urban development; this
would be detrimental considering the strong shrubland association;
however, due to low transformation in the N and some protection in
the S and N, currently assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on food associations; protected in the
coastal portion of Namaqua National Park and in strandveld of West
Coast National Park where patches of sparse grassland resulting from
clearance of shrubs on the Farm Geelbek have now been replaced by
regenerated shrubland.

SCARABAEINI
644 SURICATA 6 (2020)

Scarabaeus rusticus
(Boheman, 1857)

No synonyms
Global: LC
Endemic: RSA, Botswana, ?Namibia

J A S O N D J F M A M J

Type locality: As Ateuchus rusticus: ‘Caffraria tota’ [inexact, may be inter-

preted as anything between all SE South Africa or all southern Africa].
Taxonomy: Accepted species.
Distribution: Drier upland grasslands and woodland patches on the N
Highveld; also wooded rocky koppies in bushveld regions with a few
outliers at the edge of the Kalahari; unclear if there is truly a disjunction
between N Highveld and the Blouberg/Soutpansberg; mainly South
Africa, marginal in Botswana; suspect outliers from uMkhuze Game
Reserve (coastal KwaZulu-Natal) and Fish River Canyon (S Namibia)
not included on map.
Locality data (mean ± SD, range): Average altitude: 1 228 ± 275,
563–2 005 m; average annual rainfall: 681 ± 122, 338–1 271 mm;
average annual temperature (max. + min. /2): 18.3 ± 1.7, 12.5–22.4°C
(N=156).
Habitat: No adequate quantitative assessment; in Telperion Nature Re-
serve: mainly on upland wooded koppies at 1 379–1 452 m (29.9 per
sample), fewer in adjoining grassland at 1 419 m (12.0) or at distance
from wooded areas in grassland at 1 338–1 389 m (1.1); limited DBRU
collection records on sand (1), sandy loam (2), clay (1) in open wood-
land (4).
Food types: No quantitative assessment; limited DBRU collection records
on dung of baboon (1), cattle (2).
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Aug. to Apr.).
Ecoregions Botswana: Marginal in Zambezian and Mopane Woodlands
5 mm (AT0725) (Okavango Swamps).
Bioregions South Africa: Centred on Central Bushveld (SVcb), also, E
margin Lowveld (SVl) (Savanna Biome); N margin Mesic Highveld
Grassland (Gm), N extremity Sub-Escarpment Grassland (Gs) (Grass-
land Biome); scattered records in two other bioregions.
Assessment rationale: EOO = 105 335 km2 (main range in South Africa;
does not include outliers); widespread in N South Africa where it is
centred on drier, cooler upland areas in bushveld and at the edge of
warmer N Highveld Grassland; AOO apparently centred on wooded
patches in an often transformed region (SVcb: 1–49%), if so, clearance
of trees would be detrimental to population density; however, locally
abundant and found in various nature reserves; therefore, assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, particularly,
vegetation associations; protected in Telperion, Abe Bailey, Leeuwfon-
tein nature reserves, also Ithala Game Reserve.

SCARABAEINI
SURICATA 6 (2020) 645

Scarabaeus savignyi
Macleay, 1821

= Ateuchus transversus Castelnau, 1840

Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Scarabaeus savignii: Not stated; A. transversus: ‘Cap de

Bonne-Espérance’ [Cape of Good Hope, South Africa].
Taxonomy: Accepted species; occurs in a phylogenetic clade together with
Palaearctic species classified in the genus Ateuchetus Bedel, 1892; duly
classified as an Ateuchetus species in the 2019 version of Catalogue of
Life.
Distribution: S coastal lowlands, Western and Eastern Cape, South Afri-
ca; records from elsewhere in South Africa require validation; reports
from Namibia, Zimbabwe, presumed as errors.
Locality data (mean ± SD, range): Altitude: 388 ± 264, 99–822 m; an-
nual rainfall: 445 ± 112, 270–595 mm; annual temperature (max. +
min. /2): 16.8 ± 1.0, 15.6–18.3°C (N=7).
Habitat: No quantitative assessment; two DBRU collection records from
sandy clay loam, one from a fallow crop field.
Food types: No quantitative assessment; one DBRU collection record
from cattle dung.
Temporal activity: Diurnal flight activity late in the Western Cape, win-
ter rainy season (Oct.), or, the Eastern Cape, bimodal rainy season in
spring / early summer (Oct. to Dec.) and autumn (Mar., Apr.).
Bioregions South Africa: Vegetation units: Sandstone Fynbos (FFs),
Granite Fynbos (FFg), Limestone Renosterveld (FRl), Shale Renoster-
veld (FRs) (Fynbos Biome); Albany Thicket Biome (AT); Eastern Cape
margin of Sub-Escarpment Savanna (SVs) (Savanna Biome).
Assessment rationale: EOO = 31 775 km2; widespread along S coastal re-
5 mm
gions but very poorly known; AOO possibly influenced by finer-grained
soils, but requires confirmation; natural shrubland vegetation widely
transformed across range, but unclear if this influences current EOO
and/or AOO; assessed as Data Deficient (DD).
Conservation measures: A quantitative survey of the full EOO, AOO
and ecological associations is necessary before conservation status may
be assessed; any possible effect of natural shrubland conversion to pas-
ture grass should also be determined; protected in the Baviaanskloof
Nature Reserve (Eastern Cape).

SCARABAEINI
646 SURICATA 6 (2020)

Scarabaeus schulzeae
Ferreira, 1969

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘República da África do Sul – Transval: Great Saltpan’ [salt-

pan at W end of Soutpansberg, Limpopo Province, South Africa].
Taxonomy: Accepted species.
Distribution: Limpopo Province, South Africa; centred on and around
the W–E trending Soutpansberg; one Waterberg locality requires vali-
dation (black square).
Locality data (mean ± SD, range): Altitude: 1 015 ± 326, 461–1 626 m;
annual rainfall: 574 ± 99, 456–747 mm; annual temperature (max. +
min. /2): 19.4 ± 2.1, 15.5–22.9°C (N=13).
Habitat: No quantitative assessment; a single DBRU collection record
from sand in open woodland; one record from forest.
Food types: No quantitative assessment; a single DBRU collection record
from cattle dung.
Temporal activity: Presumably diurnal flight activity on the basis of black
pilosity; primarily active during the summer rainy season (Nov. to
Mar.).
Bioregions South Africa: Vegetation units: Limpopo Sweet Bushveld
(SVcb 19), Makhado Sweet Bushveld (SVcb 20), Soutpansberg
Mountain Bushveld (SVcb 21), Musina Mopane Bushveld (SVmp 1),
Makuleke Sandy Bushveld (SVl 1) (Central Bushveld (SVcb), Mopane
(SVmp), Lowveld (SVl) in Savanna Biome).
Assessment rationale: Smaller EOO (red squares) = 2 500 km2; larger
5 mm EOO (includes Potgietersrus [Mokopane] locality, black square) =
5 500 km2; very restricted, validated EOO would qualify S. schulzae
for an IUCN threat category of at least Vulnerable (VU); AOO could
be even more restricted if it shows specialised soil and vegetation type
associations; range centred around SVcb 21 (21% transformed) in an
area that is poorly conserved; currently not well known and assessed as
Data Deficient (DD).
Conservation measures: A quantitative survey of EOO, AOO and
ecological associations is required before conservation status may be
assessed; one record from the N Kruger National Park (South Africa),
but most of the range and most current records are from unprotected
areas, including an area scheduled for mining.

SCARABAEINI
SURICATA 6 (2020) 647

Scarabaeus suri
(Hausmann, 1807)

= Ateuchus caffer, Le Peletier & Serville, 1828

= Scarabaeus hottentotus Macleay, 1821
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Ateuchus suri: Not stated; A. caffer: ‘Cap de Bonne-

Espérance’; S. hottentotus: ‘Cap. Bon. Spei.’ [both Cape of Good Hope,
South Africa].
Taxonomy: Accepted species.
Distribution: Primarily W and SW coastal regions, Western Cape Prov-
ince, South Africa; outlier occurrence in Namaqualand (black square)
requires validation.
Locality data (mean ± SD, range): Altitude: 214 ± 384, 0–1 924 m; an-
nual rainfall: 276 ± 98, 99–466 mm; annual temperature (max. + min.
/2): 16.7 ± 1.6, 8.7–18.9°C (N=38).
Habitat: No adequate quantitative assessment; in Western Cape: sampled
in abundance from natural shrubland of cooler coastal deep sands
(Cape of Good Hope Nature Reserve: open cover: 28, dense low
cover of Restio spp.: 53, Modderrrivier: 31); rare in Kikuyu pasture
(Bonne Attente: 1) adjoining Cape of Good Hope Nature Reserve, but
super-abundant in dry sparse pasture grass on sandy loam in Darling
Hills (Groote Post: 664); DBRU collection records biased to coarse-
grained soils in open vegetation: sand (9), sandy loam (1), sandy clay
loam (1) in pasture/grassland (5), scrub (3) and crop fields (2).
Food types: No quantitative assessment; DBRU collection records on
dung: cattle (11), buffalo (1), horse (1).
Temporal activity: Diurnal flight activity from late winter till following
autumn (July to Apr.); primarily active in spring and summer (Sept. to 5 mm
Feb.); peak activity in early summer (Nov., Dec. – fully sclerotised) or
mid- to late summer (Dec. to Feb. – tenerals).
Bioregions South Africa: Centred on SW vegetation units of Sand Fyn-
bos (FFd), Sandstone Fynbos (FFs), Limestone Fynbos (FFl), Western
Strandveld (F 11), Granite Renosterveld (FRg) (Fynbos Biome).
Assessment rationale: EOO = 23 250 km2; widespread in open vegeta-
tion on sandy soils in drier parts of the Western Cape; highly trans-
formed in many places (F 11: 25–70%; FFl: 14%; FRg: 80%; FFs:
15–75%; FFd: 40–80%); in Western Cape: results are conflicting with
highest abundance in one pasture cleared of natural shrubs; otherwise
primarily in natural shrubland vegetation on sand; currently assessed as
Least Concern (LC) pending further study.
Conservation measures: Due to conflicting results, conservation status
needs to be re-assessed after conducting a survey of association with
soil type and natural shrubland versus transformed pasture habitats,
including effects of surface cover density and irrigation; protected in
Cape of Good Hope Nature Reserve (South Africa).

SCARABAEINI
648 SURICATA 6 (2020)

Scarabaeus viator
Péringuey, 1901

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: ‘Rare in the western part of the Colony (Worcester, Beau-
fort West, Knysna), and more abundant in the eastern (Graham’s Town,
Port Elizabeth, Uitenhage)’ [SW South Africa].
Taxonomy: Accepted species, but further work on Eastern Cape and
Northern Cape populations would be useful to resolve past questions
on their identity as S. viator or two different taxa.
Distribution: Arid SW South Africa; E range edge approaches that of the
extremely closely related, Scarabaeus ambiguus (Boheman, 1857).
Locality data (mean ± SD, range): Altitude: 1 135 ± 223, 249–1 797 m;
annual rainfall: 252 ± 63, 122–531 mm; annual temperature (max. +
min. /2): 17.0 ± 1.7, 12.1–19.8°C (N=169).
Habitat: No adequate quantitative assessment; in the Northern Cape:
present at almost every study site in floral regions of Upper Karoo
(99.2% of sites, N=121) and an upland area to S of Orange River strad-
dling the Upper Karoo/Bushmanland boundary (100%, N=68), rare
in SW Kalahari (11.2%, N=98) and lower-lying areas of Bushmanland
(3.0%, N=68); soil generalist in first two Northern Cape regions: sand
(21.1/site), sandy loam (21.5), sandy clay loam (30.9); DBRU collec-
tion records on sand (4), sandy loam (6), sandy clay loam (4), clay (1)
in grassland/pasture (3), scrub/shrubland (9).
Food types: No quantitative assessment; DBRU collection records on
dung: cattle (11), buffalo (1), sheep (2), elephant (1), donkey (1).
Temporal activity: Diurnal flight activity during spring and summer
across its range in winter, bimodal and late summer rainfall regions
5 mm (Oct. to May).
Bioregions South Africa: Upper Karoo (NKu), E Bushmanland (NKb),
Lower Karoo (NKl) (Nama Karoo Biome); Albany Thicket Biome
(AT); uncommon in Trans-Escarpment Succulent Karoo (SKt), Rain-
shadow Valley Karoo (SKv) (Succulent Karoo Biome); marginal NE
records from two other bioregions/biomes.
Assessment rationale: EOO = 288 635 km2; AOO would be about
half of the EOO as S. viator is rare in the hot, extremely arid, low-
altitude parts of Bushmanland lying between rare extreme SW records
at the edge of the winter rainfall region and more frequent NE records
in the higher altitude Upper Karoo; nevertheless, AOO is large in a
little-transformed area where it is readily attracted to dung of farm live-
stock; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by balanced data on ecological associations; currently only
known to be protected in Addo Elephant National Park (Eastern Cape,
South Africa).

SCARABAEINI
SURICATA 6 (2020) 649

Scarabaeus vicinus
Janssens, 1940b

No synonyms
Global: LC
Endemic: RSA, Botswana, Namibia

J A S O N D J F M A M J

Type locality: ‘Afrique australe intérieure’ [interior, southern Africa].

Taxonomy: Accepted species; very close to S. galenus Westwood, 1847.
Distribution: Centred on drier areas: NE Nama Karoo, southern Kalaha-
ri; South Africa, Botswana, Namibia.
Locality data (mean ± SD, range): Altitude: 1 180 ± 205, 870–1 998 m;
annual rainfall: 309 ± 117, 139–660 mm; annual temperature (max. +
min. /2): 17.9 ± 2.0, 14.3–22.4°C (N=74).
Habitat: No adequate quantitative assessment; in Northern Cape survey:
recorded at study sites on the continuous deep or loamy sands of SW
Kalahari or sandy pans in the Upper Karoo and Karoo areas to S of Or-
ange River (27), also sandy loam (21), but few on sandy clay loam (6);
DBRU collection records primarily from sand (10), also sandy loam
(2), sandy clay loam (1), in open vegetation that predominates in arid
EOO, Grassland (5), scrub/shrubland (7), open woodland (1).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (12).
Temporal activity: Diurnal flight activity recorded primarily in the late
summer rainy season that predominates in the SW of the EOO (Oct.
and Dec. to Mar.).
Ecoregions Namibia, Botswana: Kalahari Xeric Savanna (AT1309), mar-
gins of two adjoining ecoregions.
Bioregions South Africa: Bushmanland Nama Karoo (NKb), Upper
Karoo (NKu) (Nama Karoo Biome); Eastern Kalahari Bushveld (SVk),
Central Bushveld (SVcb) (Savanna Biome); marginal in two other
bioregions.
Assessment rationale: EOO = 791 785 km2; stronger supporting data is 5 mm
required to determine if AOO is limited by specialisation to coarser-
grained soils, which are nevertheless widespread across the large EOO;
may have been associated with dung of pellet-dropping mammals in
past when large pads of ruminant herbivores were absent from SW part
of range; now supported by farm livestock in a little-transformed area
used primarily for grazing; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by better quantitative data on ecological associations; protected
in Central Kalahari Game Reserve (Botswana).

SCARABAEINI
650 SURICATA 6 (2020)

Scarabaeus westwoodi
Harold, 1869a

= Scarabaeus laticeps Péringuey, 1901

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: S. westwoodi: ‘Africa austr. inter.’ [interior of southern

Africa]; S. laticeps: ‘Natal (Frere, Durban)’ [KwaZulu-Natal, South
Africa].
Taxonomy: Accepted species; synonymy of S. laticeps should be re-
assessed owing to lower-altitude occurrence of type localities at Frere
(blue square, 1 065 m) and Durban (inexact coastal locality) within the
range occupied by a close relative, S. caffer (Boheman).
Distribution: Primarily known from high altitude natural grasslands on
the Drakensberg escarpment along the E Lesotho border; wider distri-
bution on the N escarpment edge (Mmafefe – black square) requires
validation.
Locality data (mean ± SD, range): Altitude: 2 155 ± 695, 1 007–3 218 m;
annual rainfall: 830 ± 85, 722–1 001 mm; annual temperature (max.
+ min. /2): 11.1 ± 4.0, 4.8–18.3°C (N=8). Frere: Average altitude:
1 065 m; average annual rainfall: 730 mm; average annual temperature
(max. + min. /2): 17.2°C.
Habitat: No adequate quantitative assessment; across a gradsect up the
Drakensberg escarpment: recorded in natural grassland on sandy clay
loam at 1 900 m, 2 400 m and 2 800 m, but not at 1 500 m, 1 000 m
or 500 m; but one DBRU collection record from sandy clay loam in
pasture at ± 1 450 m (Gs 10 – see Bioregions).
Food types: No quantitative assessment; sampled to cattle dung; also one
DBRU collection record from cattle dung; abundant at 1 900 m on the
Sani Pass where no cattle were present.
5 mm Temporal activity: Diurnal flight activity (observed in Loteni Nature Re-
serve) during the summer rainy season (Nov. to Apr.).
Bioregions South Africa: Vegetation units: validated records on Sani
Pass: Southern Drakensberg Highland Grassland (Gd 4), uKhahlamba
Basalt Grassland (Gd 7), Lesotho Highland Basalt Grassland (Gd 8);
also Northern Drakensberg Highland Grassland (Gd 5), Drakensberg
Foothill Moist Grassland (Gs 10) (Sub-Escarpment Grassland (Gs),
Drakensberg Grassland (Gd)), (Grassland Biome).
Assessment rationale: EOO = 18 000 km2; (includes N record, black
square); validated EOO much smaller along the Drakensberg of the
Lesotho/South Africa border; however, validated range in Drakensberg
grasslands mostly protected; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by taxonomic revision and a quantitative survey to ascertain
the true EOO, AOO and ecological associations; most of the validated
range is protected in the uKhahlamba Drakensberg Park (World Her-
itage Site).

SCARABAEINI
SURICATA 6 (2020) 651

Scarabaeus zambesianus
Péringuey, 1901

= Scarabaeus arrowi Gillet, 1911

Global: LC

J A S O N D J F M A M J

Type localities: S. zambesianus: ‘Southern Rhodesia (Victoria Falls)’

[Zimbabwe]; S. arrowi: ‘Lac Ngami, P. B. Spei’ [Lake Ngami, Botswa-
na; Northern Cape, South Africa].
Taxonomy: Accepted species.
Distribution: Deep sand areas in dry to moist, southern African, wood-
land savanna: N South Africa, NE Namibia, Botswana, Zimbabwe;
also reported from Mozambique, Angola; report from Somalia is an
error and would probably equate to the recently described S. karlwer-
neri Nicolas & Moretto, 2002.
Locality data (mean ± SD, range): Altitude: 947 ± 313, 55–1 958 m;
annual rainfall: 445 ± 146, 216–781 mm; annual temperature (max. +
min. /2): 20.7 ± 1.7, 14.3–23.7°C (N=100).
Habitat: In uMkhuze Game Reserve: extreme bias to wooded deep sand
(deep sand: 43.0, sand over clay: 1.3, sandy clay loam: 0, clay: 0);
supported by DBRU collection records: sand (7), sandy loam (2) in
shrubland (4) or open woodland (4).
Food types: In Botswana: primarily sampled to dung (pig: 340, elephant:
255, cattle: 137, sheep: 445), few to carrion (chicken livers: 7); DBRU
collection records on various dung types: human/cattle mix (1), ele-
phant (2), rhinoceros (1), horse (2), cattle (6).
Temporal activity: Flight activity in darkness during the summer rainy
season (Oct. to Apr.); attracted to light; uses polarised light around the
moon to orientate during ball-rolling in darkness.
Ecoregions Namibia, Botswana, Zimbabwe: Moister parts of Kalaha-
ri Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian and Mopane Woodlands (AT0725), Zambezian
Baikiaea Woodlands (AT0726), Southern African Bushveld (AT0717). 5 mm

Bioregions South Africa: W Eastern Kalahari Bushveld (SVk), Central

Bushveld (SVcb), Mopane (SVmp), Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 579 415 km2; the AOO would be smaller
due to soil and possible vegetation specialisation; however, shows wide-
spread occurrence in farm rangeland and reserves; clearance of wood-
land might be detrimental, but no quantitative support; in the SW:
sand patches occur in drier areas used primarily for grazing of livestock
where transformation is low (0–2 %, W SVk); in the NE: sand patches
coincide with more threatened vegetation units (0–58% (SVl), 2–49%
(SVcb), 0–20% (SVmp)); however, wide EOO with generalist dung
associations currently warrants an assessment as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on vegetation associations, particularly
data on possible effects of woodland clearance on sand; protected in
uMkhuze Game Reserve, Kruger National Park (South Africa), Chobe
National Park (Botswana).

SCARABAEINI
652 SURICATA 6 (2020)

Genus Sceliages Westwood, 1837b

Type species and designation: Ateuchus adamastor Le Peletier de Saint Fargeau & Serville, 1828, by monotypy.
Synonyms: = Parascarabaeus Balthasar, 1961b: Type species: Scarabaeus (Parascarabaeus) tonkineus
Balthasar, 1961b, by original designation.
Last review: Entire genus reviewed by Forgie et al. (2002).

The long-established genus, Sceliages Westwood, 1837b, was recently reduced to subgeneric status by Forgie et al. (2005).
However, as it forms a distinct clade in a molecular/morphological phylogeny (Forgie et al. 2006) and shows specialist
behaviour in the use of millipede body contents for breeding (Forgie et al. 2002), there is a strong case for revalidation at
generic level (Deschodt et al. 2015b). Accordingly, it is treated as a full genus here.

The genus currently comprises seven closely related species that are restricted to southern Africa. Six of these species are
found S of 15°S. They are centred on E savanna (2), Maputaland (1), S Cape and Highveld (1), SW Kalahari (1), and
W Cape coast (1). Four species are characteristic of deep sands. All but one are currently assessed as Least Concern (LC)
although all are only sporadically collected, probably due to their specialised habits. Further ecological data would be
desirable for all species to assist assessment of conservation status.

SCARABAEINI
SURICATA 6 (2020) 653

Sceliages adamastor
(Le Peletier & Serville, 1828)

= Sceliages iopas Westwood, 1837b

= Sceliages curvipes Gillet, 1911
= Sceliages jopas Westwood, 1837b, sensu zur Strassen, 1965
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Ateuchus adamastor: ‘Cap de Bonne-Espérance’ [Cape

of Good Hope, South Africa], neotype: ‘De Hoop Nature Res.’ [South
Africa]; S. iopas: ‘Africa Austral’ [southern Africa]; S. curvipes: ‘Prom.
bon. Spei’ [Cape of Good Hope]; S. jopas sensu zur Strassen: Misspell-
ing of S. iopas.
Taxonomy: Accepted species.
Distribution: Primarily, S coast and inland areas of winter and bimodal
rainfall regions, South Africa (red squares); doubtful localities in sum-
mer rainfall region (black squares).
Locality data (mean ± SD, range): Altitude: 339 ± 453, 3–1 226 m;
annual rainfall: 337 ± 122, 132–438 mm; annual temperature (max. +
min. /2): 17.3 ± 0.9, 15.3–18.2°C (N=8, red squares only).
Habitat: No quantitative assessment; no recorded data, but collecting lo-
calities map onto shrubland vegetation units on both sandy and finer-
grained soils.
Food types: No quantitative assessment; neotype sampled using a millipede-
baited pitfall trap; observed to relocate and bury portions of dead mil-
lipedes.
Temporal activity: Diurnal flight activity during spring rainfall peaks
with activity extending into summer (Sept. to Feb.).
5 mm
Bioregions South Africa: Occurrence scattered across vegetation units
in Fynbos Biome: De Hoop Limestone Fynbos (FFl 2); Canca Lime-
stone Fynbos (FFl 3); others in Nama Karoo Biome: Eastern Upper
Karoo (NKu 4); Gamka Karoo (NKl 1); still others in Succulent Karoo
Biome: Namaqualand Klipkoppe Shrubland (SKn 1); also in Algoa
Dune Strandveld (AZs 1).
Assessment rationale: EOO = 174 095 km2 (range defined by red squares
only); impossible to adequately assess conservation status as validation
of EOO and AOO is required; locality data suggest a widely ranging
soil generalist in arid to dry areas supporting natural shrubland plant
assemblages, but quantitative support is lacking; sampling of neotype
to millipede bait supports specialist food association; currently, assessed
as Data Deficient (DD).
Conservation measures: To adequately assess conservation status, a
quantitative survey is required to validate the EOO and determine the
AOO plus ecological associations; protected along with millipedes in
De Hoop Nature Reserve, Western Cape and Addo Elephant National
Park, Eastern Cape (South Africa).

SCARABAEINI
654 SURICATA 6 (2020)

Sceliages brittoni
zur Strassen, 1965

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Süfafrika, westl. Kap-Provinz, Umgebung von Leipoldt-

ville, Clanwilliam Distr.’ [environs of Leipoldtville, Western Cape,
South Africa].
Taxonomy: Accepted species.
Distribution: W coastal deep sands of the winter rainfall region, South
Africa.
Locality data (mean ± SD, range): Altitude: 100 ± 98, 0–368 m; annual
rainfall: 199 ± 88, 103–378 mm; annual temperature (max. + min. /2):
17.2 ± 1.1, 15.4–18.5°C (N=17).
Habitat: No adequate quantitative data; sampled in low numbers on deep
sand in natural shrubland at farms Modderrivier and Geelbek, not in
adjoining sparse grassland cleared of shrubs on Geelbek (now part of
West Coast National Park where grassland has been permitted to regen-
erate to natural shrubland).
Food types: No adequate quantitative data; at Farm Geelbek: only sampled
by a single cattle dung-baited pitfall trap that also contained millipedes.
Temporal activity: Diurnal flight activity during spring of the winter
rainy season (Aug., Sept.) and in early summer (Dec.); possibly shows
a strategy of activity by mature adults in spring and emergence of the
teneral F1 generation in early summer like some other winter rainfall
endemics.
Bioregions South Africa: Vegetation units mostly on deep sand: Nama­
qualand Strandveld (SKs 7), Namaqualand Coastal Duneveld (SKs
8) (Succulent Karoo Biome); Namaqualand Sand Fynbos (FFd 1),
5 mm
Lamberts Bay Strandveld (FS 1), Langebaan Dune Strandveld (FS 5),
Leipoldtville Sand Fynbos (FFd 2), Swartland Granite Renosterveld
(FRg 2) (Fynbos Biome).
Assessment rationale: EOO = 6 180 km2; limited data suggest a soil,
vegetation and food type specialist whose AOO might be negatively
influenced by habitat transformation on deep coastal sands, which is
extensive in the S of its range (25–55%: FS 1, FS 5, FFd 2), but low
in the N (8–10%: SKs 7, SKs 8); currently assessed as Least Concern
(LC) although small range and exploitation pressures might warrant an
IUCN threat category of Near Threatened (NT) in the future.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data to support ecological associations;
protected in West Coast National Park and the private Grotto Bay
Nature Reserve, Western Cape; also in the coastal portion of Namaqua
National Park, Northern Cape (South Africa).

SCARABAEINI
SURICATA 6 (2020) 655

Sceliages difficilis
zur Strassen, 1965

No synonyms
Global: LC (see IUCN Red List – LC as Scarabaeus)

J A S O N D J F M A M J

Type locality: ‘Südafrika, östl. Kap-Provinz, Grahamstown’ [Grahams­

town, Eastern Cape, South Africa].
Taxonomy: Accepted species, but extremely close to S. gagates Shipp,
1895.
Distribution: Mainly moderate-altitude, moist savannas of SE Africa:
South Africa, Zimbabwe.
Locality data (mean ± SD, range): Altitude: 1 206 ± 387, 1–2 231 m;
annual rainfall: 734 ± 168, 519–1 171 mm; annual temperature (max.
+ min. /2): 18.0 ± 2.2, 11.1–22.5°C (N=41).
Habitat: No quantitative assessment; a single DBRU collection record
from grassland within bushveld on deep sand.
Food types: No quantitative assessment; currently, no published records
of the use of millipedes as food and breeding resource.
Temporal activity: Diurnal flight activity in the summer rainy season and
early dry season (Sept. to June).
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719), mar-
gins of two other ecoregions.
Bioregions South Africa: Primarily Central Bushveld (SVcb) (Savan-
na Biome) to extreme margins of Mesic Highveld Grassland (Gm)
(Grassland Biome); also margins of Sub-Escarpment Savanna (SVs) to
Sub-Escarpment Grassland (Gs).
Assessment rationale: EOO = 214 130 km2; currently assessed as Least
Concern (LC) on basis of large range; however, nothing known about
5 mm
habitat associations; a range centred on bushveld and miombo wood-
land suggests association with woody savanna within which millipedes
have been observed to retreat into thickets during unfavourable condi-
tions; thus, clearance of woody vegetation could be a threat although
quantitative support is currently lacking.
Conservation measures: Assessment of conservation status would be
much improved by quantitative data on ecological associations, par-
ticularly if woodland clearance influences AOO or population density
of both S. dificilis and millipedes; protected in various nature reserves
including: Blouberg, Abe Bailey, Nylsvlei (South Africa); Lake Mutirik-
we Game Reserve (Zimbabwe).

SCARABAEINI
656 SURICATA 6 (2020)

Sceliages gagates
Shipp, 1895a

No synonyms
Global: LC (see IUCN Red List – LC as Scarabaeus)

J A S O N D J F M A M J

Type localities: ‘Limpopo River’ [southern Africa], neotype: ‘Sodwana

Bay, Natal (3 km from camp)’ [South Africa].
Taxonomy: Accepted species, but extremely close to S. difficilis zur Stras-
sen, 1965.
Distribution: Deep E coastal sands: NE South Africa, SE Mozambique.
Locality data (mean ± SD, range): Altitude: 31 ± 25, 0–64 m; annual
rainfall: 778 ± 98, 654–971 mm; annual temperature (max. + min. /2):
22.4 ± 0.6, 21.8–23.8°C (N=11).
Habitat: No quantitative assessment; most localities coincide with coastal
deep sands; on deep sands in Maputo Special Reserve, Mozambique: a
single individual recorded in grassland, none recorded in forest patches.
Food types: No quantitative assessment: currently, no published records
of use of millipedes as food and breeding resource.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119),
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128) (not
shown on map).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 13 335 km2; known EOO comparatively
small; ecological data insufficient to determine if AOO would be ap-
preciably smaller; assessed as Least Concern (LC) in line with IUCN
Red List, but essentially Data Deficient (DD).
Conservation measures: Assessment of conservation status would be
5 mm much improved by quantitative data on ecological associations; pro-
tected in Maputo Special Reserve (Mozambique) and iSimangaliso
Wetland Park (World Heritage Site) (South Africa).

SCARABAEINI
SURICATA 6 (2020) 657

Sceliages granulatus
Forgie & Scholtz, 2002

No synonyms
Global: LC
Endemic: RSA, Botswana

J A S O N D J F M A M J

Type locality: ‘Botswana: Kang, 35 km SE’.

Taxonomy: Accepted species.
Distribution: Arid to dry, deep S Kalahari sands: South Africa, Botswana.
Locality data (mean ± SD, range): Altitude: 1 167 ± 80, 1 102–1 331 m;
annual rainfall: 352 ± 75, 223–455 mm; annual temperature (max. +
min. /2): 18.9 ± 0.9, 18.0–20.1°C (N=7).
Habitat: No quantitative assessment; limited DBRU collection records all
from deep sand (3) in grassland/pasture (2) or shrubland (1).
Food types: No quantitative assessment; limited DBRU collection records
all from dead millipedes (3) either from field collection or baited pitfall
traps.
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Feb.).
Ecoregions Botswana: SE Kalahari Xeric Savanna (AT1309).
Bioregions South Africa: Eastern Kalahari Bushveld (SVk) (Savanna
Biome); NE Bushmanland (NKb) (Nama Karoo Biome).
Assessment rationale: EOO = 109 200 km2; soil and food type specialist
in an area that is dominated by the preferred deep sands although that
association has not been tested quantitatively; range used, mostly, for
grazing of domestic livestock; little transformation, except in the SE
(18%, SVk 4); assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
2 mm
proved by quantitative data on habitat associations; no known localities
in protected areas, but no conservation measures recommended due to
sizeable, little-transformed range.

SCARABAEINI
658 SURICATA 6 (2020)

Sceliages hippias
Westwood, 1844

= Ateuchus microcephalus Boheman, 1857

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: S. hippias: ‘Africâ meridionali’ [southern Africa]; A. micro-

cephalus: ‘Caffraria interiore’ [interior of SE South Africa].
Taxonomy: Accepted species.
Distribution: Centred on moderate-altitude, bushveld savanna in N
South Africa.
Locality data (mean ± SD, range): Altitude: 1 302 ± 226, 600–1 702 m;
annual rainfall: 680 ± 60, 549–828 mm; annual temperature (max. +
min. /2): 17.9 ± 1.3, 15.1–20.4°C (N=41).
Habitat: No quantitative assessment; no data available.
Food types: No quantitative assessment; observed to relocate whole or
portions of dead millipedes (two species of Spirostreptidae) for burial
in Rustenburg Nature Reserve; attracted to traps baited with either live
millipedes or paper exposed to quinone-based secretions of millipedes.
Temporal activity: Diurnal flight activity primarily in the summer rainy
season (Sept. to May).
Bioregions South Africa: Primarily Central Bushveld (SVcb), also N
Sub-Escarpment Savanna (SVs) (Savanna Biome); extreme margins of
three adjoining bioregions in Grassland Biome.
Assessment rationale: EOO = 103 400 km2; currently assessed as Least
Concern (LC) on basis of fairly large range; however, nothing known
about habitat associations although centring of range on bushveld
suggests association with woody savanna within which millipedes have
been observed to retreat into thickets during unfavourable conditions;
thus, clearance of woody vegetation could be a threat although quanti-
tative support is required, particularly in SVcb vegetation units where
5 mm transformation is extensive (up to 49%).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on habitat associations of both millipedes
and S. hippias; protected in Telperion, Loskop Dam, Rustenburg and
Tswaing nature reserves (South Africa).

SCARABAEINI
660 SURICATA 6 (2020)

TRIBE SISYPHINI
Mulsant, 1842

As Sisyphaires; Type genus: As Sisyphe Latreille, 1807, but prevailng usage, Si-
syphus Latreille, 1807.

The tribe Sisyphini is probably monophyletic with an old history. It shows quite different morphology
and behaviour to other ball-rolling taxa. However, molecular phylogenies suggest close affinities to the
ball-rolling canthonine genus, Epirinus Dejean, 1833 (Tarasov & Dimitrov 2016). Therefore, they removed
Epirinus from the polyphyletic tribe, Deltochilini/Canthonini and incorporated it into the tribe Sisyphini
Mulsant, 1842, as the tribe Sisyphini sensu novo. However, based on evidence offered in a subsequent study,
Daniel et al. (2020) removed Epirinus to its own tribe, Epirinini van Lansberge, 1874a.

As previously defined, the Sisyphini s. str. comprise only three genera showing combined distributions
centred primarily on Afro-Eurasia with outliers in central America and on the island of Mauritius. More
precisely, Nesosisyphus Vinson, 1946, comprises four species endemic to Mauritius. The other two genera
(Sisyphus Latreille, 1807; Neosisyphus Müller, 1942) are widespread in forests, savannas, grasslands and win-
ter rainfall shrublands of the Afrotropical and Oriental regions with Sisyphus also present in the Palaearctic
and N Neotropical regions.

Daniel et al. (2020) have recently reduced the Sisyphini to two genera (Nesosisyphus Vinson, 1946 and
Sisyphus Latreille, 1807) with Sisyphus comprising three subgenera (Sisyphus Latreille, 1807; Parasisyphus
Barbero, Palestrini & Zunino, 1991; Neosisyphus Müller, 1942). However, as this revision was unpublished
at the time of writing, this book uses the pre-2019 classification system, but with the changes of Daniel et
al. (2020) noted in genus or species accounts.
SURICATA 6 (2020) 661

Both Sisyphus Latreille, 1807, and Neosisyphus Müller, 1942, are widespread in South Africa, with species
also found in N Botswana and N Namibia. They do not occur across most of the S Kalahari deep sands or
in most of the SW Arid region.
(1) Neosisyphus Müller, 1942: Represented by 11 species with distributions centred on winter and bi-
modal rainfall shrubland (1), Upper Karoo (1), upland grassland (2), E savanna (7), two in shade.
(2) Sisyphus Latreille, 1807: Represented by 15 currently validated species with distributions centred
on the bimodal rainfall region (1), upland grassland (3), all savanna (2), W savanna (1), E savanna
(7), E forest (2), seven in shade. No accounts provided for three new species from South Africa or
Eswatini (Daniel et al. 2018) (see Preamble).

Most species are widespread and abundant and do not face serious threats. However, those centred in E
coast forest are less secure and one has been assessed as Near Threatened (NT).

SISYPHINI
662 SURICATA 6 (2020)

Genus Neosisyphus Müller, 1942

(recent status change: subgenus of Sisyphus Latreille, 1807)

Type species and designation: Sisyphus atratus Klug, 1855, by original designation.
Synonyms: None.
Last review: Afrotropical species revised by Montreuil (2015a) with the addition of eight new spe-
cies; new southern African species added by Daniel et al. (2018); Neosisyphus has be-
come a subgenus of Sisyphus Latreille, 1807 (Daniel et al. 2020).

Neosisyphus Müller, 1942, was formally raised to full genus only recently (Montreuil 2015a) having been described as
a subgenus of Sisyphus Latreille, 1807. Although Neosisyphus retained its generic status when the book was completed
at the end of 2017, the phylogeny of Daniel et al. (2020) again reduces it to a subgenus of Sisyphus Latreille (1807). In
Montreuil’s review (2015a), species revision and cited distributions of the N. barbarossa species group differ to those of a
previous unpublished review of southern African Neosisyphus (Paschalidis 1974a). Using more detailed southern African
data, the species identities and distributions cited here for the barbarossa group follow those of Paschalidis.

Neosisyphus currently comprises 33 species widely distributed in the savannas of Africa (31) with limited representation in
the Oriental region (2). In southern Africa, Neosisyphus is represented by 11 species distributed between five groups with
different behavioural patterns. They mostly show widespread distributions centred on moister E and S regions.

Many Neosisyphus species are widespread in farm pastures and rangeland on dung of domestic livestock. They would face
few threats at present. Therefore, nine have been assessed as Least Concern (LC) although two are assessed as Data Defi-
cient (DD).

Group patterns in southern Africa:

Group designations are based on Montreuil (2015a).
Group barbarossa: Three species centred in E highlands, moist E savanna or dry E savanna in which the
egg is laid before burial of the brood ovoid.
Group quadricollis: Two species centred dry E highland grasslands or Upper Karoo and SW coastal
shrubland in which the egg is laid before burial of the brood ovoid.
Group rubrus: Two species centred on E savanna and moist upland grassland or on dry upland
grassland in which the egg is laid after burial of the ball, which has a protuberance
above the egg chamber.
Group spinipes: Three E savanna-centred species with shade, open woodland or grassland bias, two
distributed into the tropics, brood balls have a large protuberance above the egg
chamber, they are attached to grass stalks or twigs, or abandoned on the soil surface.
Group tibialis: One E coastal shade species in which the egg is laid after burial of the ball, which has
a protuberance above the egg chamber.

SISYPHINI
SURICATA 6 (2020) 663

Neosisyphus barbarossa
(Wiedemann, 1823) (but see Taxonomy)

= Sisyphus rugosus Gory, 1833

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Sisyphus barbarossa: ‘Prom. bon. sp.’ [Cape of Good

Hope, South Africa]; S. rugosus: ‘Cap B.-Esp.’ [Cape of Good Hope,
South Africa], lectotype: No locality stated.
Taxonomy: Accepted species (recent status change: genus Sisyphus La-
treille, 1807; subgenus Neosisyphus Müller, 1942), but interpretation
differs to both the unpublished revision of southern African Sisyphini ♂
(Paschalidis 1974a) and the published revision of Afrotropical Neosi-
syphus (Montreuil 2015a); both authors cite S. rugosus as a synonym;
further study is required to determine if the lectotype created for S. ru-
gosus is truly a synonym of N. barbarossa (Wiedemann), which was not
seen by Montreuil; limited sexual dimorphism (legs).
Distribution: South Africa; in the N: cool moist highlands along NE edge
of E escarpment encompassing the Lesotho highlands (1 677 ± 246 m,
N=19); in the S: descending to lowlands along the S Cape coastline
(441 ± 321 m, N=5); this distribution is consistent with that cited
by Pascahalidis (1974), but differs radically to that cited by Montreuil
(2015a).
Locality data (mean ± SD, range): Altitude: 1 408 ± 581, 110–2 199 m;
annual rainfall: 682 ± 158, 360–958 mm; annual temperature (max. +
min. /2): 14.7 ± 1.6, 11.2–17.4°C (N=23).
Habitat: No quantitative assessment; DBRU collection records from sand 2 mm
(3), sandy loam (3), sandy clay loam (6) in grassland/pasture (8) or a
fallow crop field (1).
Food types: No quantitative assessment; DBRU collection records entire-
ly from cattle dung (13).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season in the N (Oct. to Apr.); recorded in Apr. in the bimodal
rainfall region.
Bioregions South Africa: In the N: higher parts of E Mesic Highveld
Grassland (Gm), W Sub-Escarpment Grassland (Gs) (Grassland
Biome); on the S coast: cleared patches in Albany Thicket Biome (AT)
and vegetation units in Renosterveld (Fynbos Biome).
Assessment rationale: EOO = 27 150 km2; widely distributed and readily
attracted to dung of domestic livestock in farmland; however, vegeta-
tion units in N highland range threatened by transformation, particu-
larly cultivation or forest plantations (transformation in Mpumalanga
range: 22–23%, Gm 18, Gm 21; Lesotho border range: 33–50%,
Gm 1, Gm 4); unclear if S Cape pasture records reflect natural occur-
rence or are due to range expansion in response to clearance of natural
shrubland; currently assessed as Least Concern (LC) on basis of rela-
tively large range.
Conservation measures: Correct assessment of conservation status is
dependent on revision and validation of taxonomic identity and geo-
graphical range; assessment would be improved by quantitative data on 2 mm
ecological associations, particularly effects of pasture improvement and
transformation of habitats; only a single known locality, Baviaanskloof
Nature Reserve (South Africa), occurs within officially protected areas.
♀

SISYPHINI
664 SURICATA 6 (2020)

Neosisyphus calcaratus
(Klug, 1855) (but see Taxonomy)

= Sisyphus rubripes Boheman, 1857

Global: LC

J A S O N D J F M A M J

Type localities: As Sisyphus calcaratus: ‘Sena’ [central Mozambique]; S. ru-

bripes: ‘Caffraria interiore’ [interior of SE South Africa].
Taxonomy: Accepted species (but recent status change: genus Sisyphus La-
treille, 1807; subgenus Neosisyphus Müller, 1942); formerly erroneously
synonymised with Neosisyphus rugosus (Gory, 1833); valid species status
supported by revisions of southern African (Paschalidis 1974a, unpub-
♂ lished) and African Sisyphini (Montreuil 2015a, published); limited
sexual dimorphism (legs).
Distribution: Widespread in hot, dry, southern African savanna wood-
land: South Africa, Eswatini, Botswana, Zimbabwe, Mozambique,
Zambia; other citations associated with N. rugosus presumably errors:
Angola, Democratic Republic of the Congo (DRC), Rwanda, Kenya,
Ethiopia.
Locality data (mean ± SD, range): Altitude: 744 ± 422, 41–1 847 m;
annual rainfall: 612 ± 152, 306–1 184 mm; annual temperature (max.
+ min. /2): 21.0 ± 2.1, 14.7–25.7°C (N=86).
Habitat: No adequate quantitative data; in uMkhuze Game Reserve:
inconclusive soil data: sand (3.8), duplex soil (sand over clay) (24.3),
sandy clay loam (12.3), clay (14.3); DBRU collection records on sand
(7), sandy loam (6), sandy clay loam (1), clay (1) with a bias to wooded
habits: grassland/pasture (2), shrubland (4), open woodland (8), forest
(2).
2 mm
Food types: At Phalaborwa: readily attracted to dung with bias to cattle
(450) as opposed to pig (230) or elephant (130); DBRU collection
records from dung of cattle (15), buffalo (2), wildebeest (1), waterbuck
(1), elephant (6), rhinoceros (1), horse (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.); at Phalaborwa: more abundant on a warm
sunny day soon after heavy rainfall (554) than on a hot sunny day (225)
or a warm cloudy day following light rainfall (21).
Ecoregions Botswana, Zimbabwe: Drier areas: Zambezian and Mopane
Woodlands (AT0725), Southern African Bushveld (AT0717), margins
of Kalahari Xeric Savanna (AT1309), Kalahari Acacia-Baikiaea Wood-
lands (AT0709), margins of two other ecoregions.
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl), Mo-
pane (SVmp) (Savanna Biome).
Assessment rationale: EOO = 712 200 km2; AOO would be smaller if
associated with woody habitats; proportion of tree clearance unknown,
but transformation across South African range varies from 0 to 58%
(SVl), 2–49% (SVcb), 0–20% (SVmp); however, widespread and read-
ily attracted to various dung types in both reserves and farmland, so
assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations, particularly
those with vegetation; protected in Kruger National Park and uMkhuze
Game Reserve (South Africa).

♀
2 mm

SISYPHINI
SURICATA 6 (2020) 665

Neosisyphus confrater
(Kolbe, 1914) (but see taxonomy)

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: As Sisyphus confrater: ‘Bei Bukoba’ [near Bukoba, Tanzania].

Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); currently synony-
mised with Neosisyphus setiger (Roth, 1851) described from ‘Abyssinie’
[Ethiopia]; interpreted here according to an unpublished revision of
southern African Sisyphini (Paschalidis 1974a); however, species iden-
tity of southern African populations needs to be validated by further ♂
comparison with types (Tanzania (lectotype male), Ethiopia (holotype
female)) and other E African material; formerly synonymised in error
with Neosisyphus armatus (Gory, 1833), a species now considered to
show a W African distribution; limited sexual dimorphism (legs).
Distribution: Moist savanna on SE coast and lower edges of SE uplands
of Africa: South Africa, Eswatini, Mozambique, Zimbabwe; putative
type locality in Tanzania.
Locality data (mean ± SD, range): Altitude: 687 ± 631, 0–1 890 m; an-
nual rainfall: 892 ± 159, 627–1 247 mm; annual temperature (max. +
min. /2): 19.7 ± 3.0, 13.1–25.2°C (N=50).
Habitat: In Ithala Game Reserve: only recorded on sandy loam in grass-
land (8) at upper edge of escarpment (1 142 m); DBRU collection
records biased to finer-grained soils, sand (7), sandy loam (8), sandy
clay loam (7), clay (2) in unshaded vegetation, pasture/grassland (25), 2 mm
open woodland (3) forest (1).
Food types: In Ithala Game Reserve: attracted to dung of pig (4), horse
(2), cattle (2); DBRU collection records primarily on cattle dung (27)
with few on horse (1), elephant (1) or human dung (3).
Temporal activity: Diurnal flight activity primarily during the summer
rainy season (Sept. to May).
Ecoregions Zimbabwe, Mozambique: Southern Miombo Woodland
(AT0719), Eastern Zimbabwe Montane Forest Mosaic (AT1006),
Maputaland Coastal Forest Mosaic (AT0119), Southern Zanzibar-
Inhambane Coastal Forest Mosaic (AT0128).
Bioregions South Africa: Indian Ocean Coastal Belt Biome (CB) and
warmer areas on edge of escarpment in Mesic Highveld Grassland
(Gm) (Grassland Biome) bordering Central Bushveld (SVcb) and
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 118 955 km2 (range in southern Africa
only; putative Bukoba type locality also marked); apparent bias to
grassland association may indicate a different species to that described
from Bukoba; currently assessed as Least Concern (LC) on the basis of
extensive EOO in open vegetation.
Conservation measures: Correct assessment of conservation status is
dependent on revision and validation of taxonomic identity and geo-
graphical range; assessment would also be improved by further quanti-
tative data on ecological associations; protected in Ithala Game Reserve
2 mm
(South Africa).

SISYPHINI
666 SURICATA 6 (2020)

Neosisyphus fortuitus
(Péringuey, 1901) (but see Taxonomy)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Sisyphus fortuitus: ‘Natal (Durban)’ [Durban, KwaZulu-

Natal, South Africa].
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); limited sexual
dimorphism (legs).
Distribution: Moist, coastal and upland, woodland savanna of SE Africa:
♂ N South Africa, E Zimbabwe, Eswatini, S Mozambique; citations from
Cameroon and Guinea (Conakry) are errors.
Locality data (mean ± SD, range): Altitude: 748 ± 560, 0–1 613 m; an-
nual rainfall: 746 ± 155, 491–1 188 mm; annual temperature (max. +
min. /2): 19.8 ± 2.0, 15.6–22.6°C (N=56).
Habitat: In Gauteng bushveld: moderate bias to deep sand (264) com-
pared to sandy clay loam (97) with strong bias to shaded thicket (252)
and open woodland (98) compared to grassland (11); in uMkhuze
Game Reserve: 66% of 279 recorded in shade with weak bias to sandy
soils, sand (10.8), sand over clay (3.8), sandy clay loam (8.5), clay (3.1);
largely supported by DBRU collection records from sand (6), sandy
loam (4), sandy clay loam (2) in forest/thicket (4), shrub/woodland
(6), grassland (4).
Food types: No quantitative assessment; DBRU collection records on
dung of cattle (14), wildebeest (2), rhinoceros (2), elephant (1), horse
(1), human (1), human/cattle mix (5).
2 mm Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to June).
Ecoregions Zimbabwe, Mozambique:, Eastern Zimbabwean Montane
Forest Grassland Mosaic (AT1006), Maputaland Coastal Forest Mosaic
(AT0119) ), margins of one other ecoregion.
Bioregions South Africa: Centred on Central Bushveld (SVcb), Lowveld
(SVl) (Savanna Biome); Indian Ocean Coastal Belt Biome (CB); also
extreme margins of adjoining Grassland Biome.
Assessment rationale: EOO = 154 350 km2; AOO centred on woodland
savanna owing to habit of abandoning brood balls in the litter under
shade offered by shrubs and trees; therefore EOO and AOO would be
threatened by clearance of woody vegetation; transformation in vegeta-
tion units is 2–49% in SVcb and 0–58% in SVl; however, widespread
in N South Africa in both reserves and on farms; currently assessed as
Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on dung type associations and an appraisal
of the occurrence of shaded vegetation across its EOO; protected in
uMkhuze, Hluhluwe–iMfolozi and Ithala game reserves (South Africa).

♀
2 mm

SISYPHINI
SURICATA 6 (2020) 667

Neosisyphus infuscatus
(Klug, 1855) (but see Taxonomy)

= Sisyphus atratus Klug, 1855, sensu Péringuey, 1901

= Neosisyphus bornemisszai Ferreira, 1972
Global: LC

J A S O N D J F M A M J

Type localities: As Sisyphus infuscatus: ‘Sena’, lectotype: ‘Sena’ [central

Mozambique]; S. atratus sensu Péringuey: ‘Cape Colony (Queenstown)’
[Eastern Cape, South Africa]; N. bornemisszai: ‘Manhoca (Moçam-
bique)’ [Mozambique].
Taxonomy: Accepted species (but recent status change: genus Sisyphus La-
treille, 1807; subgenus Neosisyphus Müller, 1942); formerly erroneously ♂
synonymised with Neosisyphus spinipes (Thunberg, 1818); valid species
status supported by revisions of southern African (Paschalidis 1974a,
unpublished) and African Sisyphini (Montreuil 2015a, published);
limited sexual dimorphism (legs).
Distribution: Dry, hot lowland savanna on the E seaboard from S to E
Africa: South Africa, Zimbabwe, Zambia, Mozambique, Tanzania,
Kenya.
Locality data (mean ± SD, range): Altitude: 431 ± 392, 0–1 432 m; an-
nual rainfall: 687 ± 221, 306–1 275 mm; annual temperature (max. +
min. /2): 21.2 ± 2.4, 15.0–25.7°C (N=56).
Habitat: No adequate quantitative assessment; in uMkhuze Game Re-
serve: recorded on sand (2.0), duplex soil (sand over clay) (9.3), sandy
clay loam (7.5) and clay (4.5); DBRU collection records on sand (9),
sandy loam (2), sandy clay loam (8) and clay (4) in grassland (5), shrub/
woodland (9) and forest (3).
Food types: At Phalaborwa: stronger association with cattle dung (49)
compared to pig (15) or elephant dung (14); DBRU collection records
from dung of cattle (19), buffalo (4), wildebeest (1), waterbuck (1),
elephant (4), human/cattle dung mix (2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Apr.); also in cooler drier months (Aug., May) in the warmer 2 mm
coastal region of KwaZulu-Natal; at Phalaborwa: abundant both in
warm conditions soon after heavy rainfall (42) and in hot, dry condi-
tions (28), but not in warm cloudy conditions soon after light rainfall
(2).
Ecoregions Zimbabwe: Zambezian and Mopane Woodlands (AT0725).
Bioregions South Africa: Lowveld (SVl), Mopane (SVmp) (Savanna
Biome); drier parts, Indian Ocean Coastal Belt (CB) and Albany
Thicket (AT) biomes; marginal in two other bioregions.
Assessment rationale: EOO = 900 785 km2 (gross estimate); apparently
a soil type generalist, possibly, with a bias to woodland savanna; if so
might be influenced by clearance of trees; in the main part of its range
in South African savanna, transformation is 0–58% (SVl), 0–20%
(SVmp); however, assessed as Least Concern (LC) owing to wide range
in dry savanna.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on habitat associations, particularly
vegetation associations; protected in the Kruger National Park and
uMkhuze Game Reserve (South Africa).

2 mm

SISYPHINI
668 SURICATA 6 (2020)

Neosisyphus kuehni
(Haaf, 1955) (but see Taxonomy)

No synonyms
Global: DD (see IUCN Red List – DD)
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Sisyphus kühni: ‘Südafrika (Natal, Griqualand)’ [KwaZulu-

Natal, South Africa].
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); limited sexual
dimorphism (legs).
♂ Distribution: Moist upland to highland grassland along middle to upper
edge of the E escarpment, summer rainfall region, South Africa.
Locality data (mean ± SD, range): Altitude: 1 646 ± 239, 1 225–1 890 m;
annual rainfall: 783 ± 143, 556–996 mm; annual temperature (max. +
min. /2): 14.1 ± 1.4, 12.6–17.0°C (N=8).
Habitat: No adequate quantitative assessment; on Farm Boschberg in
natural grassland on sandy clay loam: only recorded on Highveld (7,
1 675–1 688 m) not in grassland at base of hill (0, 1 265–1 302 m) lim-
ited DBRU collection records from various soil types, sand (1), sandy
loam (2), sandy clay loam (1) in grassland/pasture (3).
Food types: No quantitative assessment; limited DBRU collection records
all from cattle dung (4).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Nov. to May).
2 mm Bioregions South Africa: In the N: Mesic Highveld Grassland (Gm),
Sub-Escarpment Grassland (Gs); in the S: S Dry Highveld Grassland
(Gh), S patch Drakensberg Grassland (Gd) on Amatola Mts (Grassland
Biome).
Assessment rationale: EOO = 30 620 km2; widespread, but poorly
known; probably restricted to highland grassland, but quantitative
support required; could probably be assessed as Least Concern (LC) on
basis of widespread occurrence, but currently assessed as Data Deficient
(DD); low numbers in collections could equally be a collection artefact
or due to low population density; further study required, particularly
considering there is 23–52% habitat transformation around known
localities in the N of its range (Gm 18, Gm 22, Gs 8).
Conservation measures: Before conservation status may be adequately
assessed, a quantitative survey of highland grassland is required to
determine ecological associations and possible effects of habitat distur-
bance, particularly conversion to arable land, pasture improvement or
commercial forestry; not currently known to occur in any protected
area.

2 mm

SISYPHINI
SURICATA 6 (2020) 669

Neosisyphus macrorubrus
(Paschalidis, 1974b) (but see Taxonomy)

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA, Namibia

J A S O N D J F M A M J

Type locality: As Sisyphus macrorubrus: ‘Vanstadensrus, O.F.S.’ [Free

State, South Africa].
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); limited sexual
dimorphism (legs).
Distribution: Disjunct pattern: South Africa, Namibia; cool, dry, SW ♂
Highveld and arid NE Nama Karoo uplands, South Africa; uncommon
outlier presence on uplands of N Namibian plateau.
Locality data (mean ± SD, range): Altitude: 1 295 ± 235, 814–2 392 m;
annual rainfall: 366 ± 148, 221–815 mm; annual temperature (max. +
min. /2): 16.8 ± 2.2, 9.9–22.2°C (N=129).
Habitat: No adequate quantitative assessment; in Northern Cape: greatest
frequency in cooler Upper Karoo (80% of study sites), less in lower-
altitude Karoo (14%) and Kalahari (34%), more frequent on sand
(1.5/trap, N=75), but similar abundance on sandy loam (2.2, N=29);
DBRU collection records from sand (11), sandy loam (12), sandy clay
loam (10), clay (2) in grassland/pasture (15), scrub/shrubland (30),
open woodland (5).
Food types: No quantitative assessment; most DBRU collection records
from cattle dung (41), also zebra (1)
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
Ecoregions Namibia: S Angolan Mopane Woodlands (AT0702).
Bioregions South Africa: Cooler S Mesic Highveld Grassland (Gm), Dry
Highveld Grassland (Gh) (Grassland Biome); Eastern Kalahari Bush- 2 mm
veld (SVk) (Savanna Biome); moister NE Upper Karoo (NKu) (Nama
Karoo Biome).
Assessment rationale: EOO = 239 965 km2; in South Africa: SW limits
of distribution centred in an area used primarily for grazing of domestic
livestock to whose dung it is readily attracted; E limits more trans-
formed (4–50%, Gh, Gm); although poorly protected in South Africa,
little threatened over most of range; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further quantitative ecological data; protected in Etosha
National Park (Namibia) and the private Tswalu Kalahari Reserve
(South Africa).

♀
2 mm

SISYPHINI
670 SURICATA 6 (2020)

Neosisyphus mirabilis
(Arrow, 1927) (but see Taxonomy)

= Sisyphus spinipes Gory, 1833

Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type localities: As Sisyphus mirabilis: nom nov. for Sisyphus spinipes Gory:
‘Cap B.-Esp.’ [Cape of Good Hope, South Africa], lectotype: Not stat-
ed.
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); limited sexual
dimorphism (legs).
♂ Distribution: Possible disjunction between two centres in coastal low-
lands of Maputaland Centre of Endemism and Eastern Cape; South
Africa, Mozambique; more northerly distribution on Mozambique
coast requires investigation.
Locality data (mean ± SD, range): Altitude: 82 ± 170, 0–762 m; annual
rainfall: 825 ± 154, 360–1 010 mm; annual temperature (max. + min.
/2): 21.7 ± 1.2, 17.4–22.6°C (N=35).
Habitat: In uMkhuze Game Reserve: bias to deep sand (116.2), but abun-
dant on duplex soil (sand over clay) (29.4), sandy clay loam (11.9) and
clay (35.3); on deep sands in Maputo Special Reserve: more abundant
in large and medium patches of inland sand forest (4.4, 4.0/trap) than
5 mm in small patches (0.7) or cooler coastal dune forest (0.5), virtually ab-
sent from grassland (0.1); on deep sands at Richards Bay: more abun-
dant in regenerating woodland on mined coastal dunes (12–33 yr)
(6.8/trap), rare in deeper shade of natural dune forest (0.02); limited
DBRU collection records from sandy clay loam (1) in open woodland
(1) and forest (5).
Food types: No quantitative assessment; limited DBRU collection records
from horse (1) and a mixture of human and cattle dung (2).
Temporal activity: Diurnal flight activity throughout much of the year
in warmer N coastal range (July to May); probably only in warmer
summer in Eastern Cape.
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: In KwaZulu-Natal: N Indian Ocean Coastal
Belt Biome (CB); also adjoining Lowveld (SVl) (Savanna Biome); in
Eastern Cape: Albany Thicket Biome (AT).
Assessment rationale: EOO = 14 150 km2; AOO more limited due to
specialised shade association; protected over about 50% of the known
range, but would be negatively influenced by woodland or forest clear-
ance; transformation in Zululand SVl: 22–58%, but assessed as Least
Concern (LC) owing to extent of protection.
Conservation measures: Assessment of conservation status would be
improved by data on food associations; protection offered over much
of the range, particularly in Baviaanskloof Nature Reserve, Hluhluwe–
iMfolozi and uMkhuze game reserves (South Africa); Maputo Special
Reserve (Mozambique).

SISYPHINI
SURICATA 6 (2020) 671

Neosisyphus quadricollis
(Gory, 1833) (but see Taxonomy)

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J West

J A S O N D J F M A M J East

Type localities: As Sisyphus quadricollis: ‘Cap B.-Esp.’ [Cape of Good

Hope, South Africa], lectotype: Not stated.
Taxonomy: Accepted species (but recent status change: genus Sisyphus
♀
Latreille, 1807; subgenus Neosisyphus Müller, 1942); despite ecological
differences between W and E populations, now established as a single
species (Daniel et al. 2018); limited sexual dimorphism (legs).
Distribution: Possibly disjunct occurrence between dry SW coastal deep
sands in Western Cape (red squares) and dry fynbos and Karoo of East-
ern Cape (black squares), South Africa.
Locality data (mean ± SD, range): West: average altitude: 32 ± 13,
15–45 m; average annual rainfall: 282 ± 63, 191–326 mm; average
annual temperature (max. + min. /2): 16.5 ± 1.0, 15.7–17.9°C (N=4).
East: average altitude: 1 072 ± 235, 868–1 392 m; average annual rain-
fall: 314 ± 38, 270–363 mm; average annual temperature (max. + min.
/2): 14.9 ± 0.2, 14.7–15.1°C (N=4).
Habitat: No adequate quantitative assessment; W populations on W
coast deep sands in natural shrubland (23); in the W, not recorded
on finer-grained soils or in pastures cleared of natural shrubland; E
populations on sandy loam in Karoo grassland, also sand and clay soil 5 mm
areas in E Fynbos-Renosterveld Bioregion.
Food types: No quantitative assessment; sampled to cattle dung and cat-
tle/sheep mixture.
Temporal activity: Diurnal flight activity; in the winter rainfall region:
West Coast populations on farms Modderrivier and Pampoenvlei show
spring activity declining into early summer (Oct., 13; Nov., 8; Dec.,
2); in the bimodal rainfall region: records are only from late summer
(Jan. to March).
Bioregions South Africa: Vegetation units in the W: West Strandveld (FS
1, FS 5), Atlantis Sand Fynbos (FFd 4); in the E: Uniondale Shale
Renosterveld (FRs 16), South Rooiberg Sandstone Fynbos (FFs 22)
(Fynbos Biome); one E record in Upper Karoo (NKu) (Nama Karoo
Biome).
Assessment rationale: EOO = 2 750 km2 (W), 23 050 km2 (E); W and E
populations apparently differ in seasonal activity, altitude and tempera-
ture range in similarly dry areas; most records from areas of Fynbos in
winter and bimodal rainfall regions with one record from Nama Karoo
in the late summer rainfall region; W populations face threats as they
occupy a small range in areas that are 25–40% transformed; however,
many contradictions in existing data; currently assessed as Data Defi-
cient (DD).
Conservation measures: Assessment of conservation status would benefit
from a survey to establish the full EOO and AOO as well as quanti-
tative data on ecological associations in both W and E populations;
no current records known from officially protected areas although the
coastal part of Modderrivier is now in the private Grotto Bay Nature
Reserve.

SISYPHINI
672 SURICATA 6 (2020)

Neosisyphus rubrus
(Paschalidis, 1974b) (but see Taxonomy)

No synonyms
Global: LC (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: As Sisyphus rubrus: ‘Castle Gorge, Tvl. 42 mi W [of ] Preto-

ria’ [North-West Province, South Africa].
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); misidentified
by Péringuey (1902), as S. rubripes Boheman, 1857; re-described by
Haaf (1955), as S. rubripes Péringuey, a pre-occupied name, thus,
♂ invalidating the lectotype designated from Péringuey’s material; limited
sexual dimorphism (legs).
Distribution: Warmer areas of upland grassland; also moist, SE African
open savanna woodland: South Africa, E Botswana, Zimbabwe, Zam-
bia, Mozambique.
Locality data (mean ± SD, range): Altitude: 1 067 ± 461, 8–3 218 m;
annual rainfall: 704 ± 148, 336–1 203 mm; annual temperature (max.
+ min. /2): 18.3 ± 2.4, 4.8–25.1°C (N=299).
Habitat: In Gauteng bushveld: strong bias to sandy clay loam (2 759)
compared to deep sand (638) primarily in open vegetation: grassland
(2 197), open woodland (1 171), shaded thickets (29); supported in
uMkhuze Game Reserve: sand (0.1), sand over clay (0.8), sandy clay
loam (6.5), clay (1.0); also supported by DBRU collection records from
sand (40), sandy loam (52), sandy clay loam (53), clay (5) in grassland/
pasture (85), open shrub/woodland (44), forest/thicket (3).
Food types: At Phalaborwa: attracted more abundantly to dung of cattle
(287) than that of elephant (90) or pig (93); supported in Gauteng
bushveld: cattle (48), horse (24), pig (9), carrion (0); DBRU collection
records from dung of cattle (167), buffalo (3), wildebeest (1), impala
(1), elephant (3), horse (2), human (2).
Temporal activity: Diurnal flight activity; in Gauteng: midday peak
2 mm
(12:00–14:00) in Nov. to Feb., mid-afternoon peak (14:00–16:00) in
Mar.; active mainly throughout summer rains (Oct. to Apr.), declining
into dry season (May); low numbers active at end of dry season (Aug.,
Sept.); at Phalaborwa: greater abundance on a warm sunny day soon
after heavy rainfall (322) than on a hot sunny (124) or a warm cloudy
day following light rainfall (3); after colonising and shredding cattle
pads, make and roll ball only on the following day.
Ecoregions Zimbabwe: Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
vanna Biome); Sub-Escarpment Grassland (Gs) (Grassland Biome);
Albany Thicket (AT) and Indian Ocean Coastal Belt (CB) biomes;
marginal in E Eastern Kalahari Bushveld (SVkb) (Savanna Biome) plus
N Mesic Highveld Grassland (Gm), Dry Highveld Grassland (Gh)
(Grassland Biome).
Assessment rationale: EOO = 720 705 km2 (? underestimated); wide-
spread across both reserves and farmland where it shows a bias to the
dung of pad-dropping ruminant herbivores in open vegetation includ-
ing farm pastures; thus little threatened in areas dominated by grazing
of farm livestock; assessed as Least Concern (LC).
2 mm Conservation measures: None recommended; protected in Kruger Na-
tional Park, uMkhuze Game Reserve (South Africa).
♀

SISYPHINI
SURICATA 6 (2020) 673

Neosisyphus spinipes
(Thunberg, 1818) (but see Taxonomy)

= Sisyphus atratus Klug, 1855

= Sisyphus appendiculatus Boheman, 1857
= Sisyphus hessei Gory, 1833
= Sisyphus quadricollis Gory, 1833, sensu Péringuey, 1901
Global: LC

J A S O N D J F M A M J

Type localities: As Ateuchus spinipes: ‘majorem horum partem in capite bo-

nae spei collegi’ [amongst material mostly collected in the Cape of Good
Hope, South Africa]; S. atratus: ‘Tette’ [Tete, central Mozambique]; S. ap-
pendiculatus: ‘Caffraria tota’ [inexact; ?all SE South Africa], lectotype: ♂
‘Caffraria’; S. hessei: ‘Cap B.-Esp.’ [Cape of Good Hope, South Africa],
lectotype: Not stated; S. quadricollis Gory sensu Péringuey: ‘Cape Col-
ony (Transkei, Port Elizabeth, Stockenstroom, Graham’s Town, Somer-
set East) Natal (Durban), Mozambique (Lourenço-Marquez and Beira,
Sena...)’ [South Africa: Eastern Cape, KwaZulu-Natal; Mozambique].
Taxonomy: Accepted species (but recent status change: genus Sisyphus
Latreille, 1807; subgenus Neosisyphus Müller, 1942); limited sexual di-
morphism (legs).
Distribution: Moist, warm savanna and grasslands on the E seaboard
from S to NE Africa: South Africa, Zimbabwe, Mozambique, Zambia,
Democratic Republic of the Congo (DRC), Rwanda, Burundi, Ugan-
da, Tanzania, Kenya, Ethiopia; also reported from Botswana; citation
from Senegal presumed an error.
Locality data (mean ± SD, range): Altitude: 484 ± 485, 0–1 706 m;
annual rainfall: 823 ± 200, 462–1 659 mm; annual temperature (max.
+ min. /2): 20.2 ± 2.3, 14.4–25.2°C (N=126).
Habitat: No adequate quantitative data; soil generalist in uMkhuze Game
Reserve: sand (2.8), duplex soil (sand over clay) (4.3), sandy clay loam
(4.6), clay (2.2); supported by DBRU collection records: sand (23),
sandy loam (7), sandy clay loam (21), clay (6) in grassland/pasture (37),
open shrub/woodland (18), forest/thickets (4). 2 mm
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (60), wildebeest (3), elephant (3), rhinoceros (1), horse
(1), human (1), mix of human and cattle or buffalo (3).
Temporal activity: Diurnal flight activity during the summer rainy sea-
son (Oct.–Apr.) and also cooler, drier months at the warmer coastline
(Aug., Sept., May).
Ecoregions Zimbabwe, Mozambique: Southern Miombo Woodlands
(AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Moist parts of Lowveld (SVl) (Savanna Biome),
Indian Ocean Coastal Belt Biome (CB); also moist, warm E margins of
Central Bushveld (SVcb) (Savanna Biome), Mesic Highveld Grassland
(Gm), Sub-Escarpment Grassland (Gs) (Grassland Biome); outlier in
transformed pasture habitats in Eastern Cape (Fynbos Biome).
Assessment rationale: EOO = 1 156 170 km2 (underestimated; NE Afri-
can range omitted); apparently a soil generalist centred on warm moist
grassland and savanna habitats, probably due to habit of cementing
brood balls to grass stalks, or other vegetation, rather than burying
them in soil; not susceptible to woodland clearance due to grassland
habitat bias; also widespread; therefore assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
Kruger National Park and Hluhluwe–iMfolozi Game Reserve. ♀ 2 mm

SISYPHINI
674 SURICATA 6 (2020)

Genus Sisyphus Latreille, 1807

(recent status change, three subgenera: Sisyphus, Neosisyphus Müller,
1942, and Parasisyphus Barbero, Palestrini & Zunino, 1991)

Type species and designation: Sisyphus schaefferi Linnaeus, 1758, by monotypy.

Synonyms: None.
Last review: All species reviewed by Haaf (1955); southern African species reviewed by Paschalid-
is (1974, unpublished); recent piecemeal revisions and addition of many new valid
Afrotropical species (Endrödi 1973; Walter 1982; Cambefort 1984; Schafer & Fischer
1992, 2001; Montreuil 2014, 2015b, 2015c, 2017; Daniel et al. 2016); recent status
change: Sisyphus divided into three subgenera (Sisyphus; Neosisyphus Müller, 1942; Pa-
rasisyphus Barbero, Palestrini & Zunino, 1991) by Daniel et al. (2020).

The long-established genus, Sisyphus Latreille, 1807, currently remains without a full recent revision. Subsequent to the
earlier revision of the entire genus as two subgenera, Neosisyphus and Sisyphus s. str. (Haaf 1955), revisions were made to
the southern African species by Paschalidis (1974). More recently, there has been piecemeal revision of some Sisyphus spe-
cies groups and the addition of many new Afrotropical species (references listed above). Because of incomplete revision,
southern African species listed here largely, though not entirely, follow Paschalidis (1974a) with the addition of modifica-
tions and new species added by Montreuil (2015b, 2015c) and Daniel et al. (2016). However, following the recent formal
raising of Neosisyphus Müller, 1942, from subgeneric to generic level (Montreuil 2015a) , Sisyphus has now been redefined
as three subgenera (Daniel et al. 2020) (Sisyphus Latreille, 1807; Parasisyphus Barbero, Palestrini & Zunino, 1991; Neosi-
syphus Müller, 1942). Although the old pre-2020 classification system is used, here, a note is included with group patterns
and species accounts to indicate whether they belong to the subgenera Sisyphus or Parasisyphus under the new system.

At the end of 2017, Sisyphus comprised a total of 50 species with distributions throughout the Afrotropical (37), S Ori-
ental (10) and S Palaearctic regions (1) with further species (2) isolated in central America at the N tip of the Neotropical
region. A few new Oriental species have also been described since the revision of Haaf (1955) whereas further new Afro-
tropical species were added by Daniel et al. (2018) for which there are no species accounts. Species accounts have, however,
been prepared for a total of 15 named species occurring in South Africa, Botswana and Namibia.

Group patterns in southern Africa: Group designations are based on relationships indicated by Paschalidis (1974) and
Montreuil (2015b, 2015c, 2017). Five groups occur in southern Africa.
Group costatus: Four species centred on SE upland grasslands (2), SE savanna (1) or widespread
in savanna with a distribution into the tropics (1) (recent status change: subgenus
Parasisyphus Barbero, Palestrini & Zunino, 1991).
Group impressipennis: One savanna species with bias to shade distributed into the E tropics (recent status
change: subgenus Parasisyphus Barbero, Palestrini & Zunino, 1991).
Group muricatus: Six species characterised by tufts of setae; three larger-bodied species centred on SE
Highveld Grassland plus cooler S coast (1), SE coastal forest (1) or SE shady savanna
plus NE escarpment forest (1); three smaller-bodied species centred on moist SE
upland grassland and moist SE savanna (1), S coast (1) or N Namibia distributed
into the E tropics (1, brachypterous) (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991).
Group ocellatus: Two species found in shady situations in E savanna (1) or SE coastal forest (1) (re-
cent status change: subgenus Sisyphus Lateille, 1807.
Group seminulum: Two species found in shady lowland savanna along the E seaboard of Africa (1) or in
N Namibia and Angola (1) (recent status change: subgenus Sisyphus Lateille, 1807.

SISYPHINI
SURICATA 6 (2020) 675

Sisyphus alveatus
Boucomont, 1935

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘Kenya Colony, Kibwezi, Wa-Kamba’, lectotype: Afrique

Orle anglaise, Kibwézi (Wa-Kamba) [Kibwezi, Kenya].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991) in a group comprising macropter-
ous and brachypterous taxa; in an unpublished revision of southern Af-
rican Sisyphus (Paschalidis 1974a), S. alveatus from South Africa is cited
in error for a species recently described as Sisyphus manni Montreuil,
2015c; citations from Zimbabwe (as Rhodesia) (Paschalidis, 1974a)
cannot be validated (material missing).
Distribution: Moist savanna of S central Africa to NE Africa: Namibia,
Angola, Zambia, Mozambique, Tanzania, Kenya, Ethiopia; in Na-
mibia, known only from Epupa Falls, which are bordered by riparian
vegetation.
Locality data: Altitude: 910 m; annual rainfall: 541 mm; annual tempera-
ture (max. + min. /2): 22.6°C (N=1).
Habitat: No quantitative assessment; no data.
Food types: No quantitative assessment; no data.
Temporal activity: Brachypterous; diurnal activity during the summer
rainy season (Nov.).
Ecoregions Namibia: Zambezian Baikiaea Woodlands (AT0726).
Assessment rationale: EOO = 1 801 180 km2 (gross estimate); AOO
presumably very much smaller than widespread EOO, but very poorly
known ecologically; may be susceptible to habitat modification as a
brachypterous species; in the absence of detailed data, assessed as Data 2 mm

Deficient (DD).
Conservation measures: To assess conservation status, a survey is required
to determine if AOO is particularly restricted within the extensive
EOO; quantitative data on ecological associations is also required from
several localities across its wide range; protected in Mkomazi Game
Reserve, Tanzania.

SISYPHINI
676 SURICATA 6 (2020)

Sisyphus caffer
Boheman, 1857

No synonyms
Global: LC
Endemic: RSA (see IUCN Red List – LC)

J A S O N D J F M A M J

Type locality: ‘Caffraria tota, passim’ [everywhere in all SE South Africa].

Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991).
Distribution: Cool, moist grasslands of central to S Highveld and SSE
coastal regions, South Africa with outlier occurrence in the N at higher
altitude on the Blouberg.
Locality data (mean ± SD, range): Altitude: 1 260 ± 383, 43–1 782 m;
annual rainfall: 674 ± 118, 339–948 mm; annual temperature (max. +
min. /2): 16.5 ± 1.7, 12.9–20.0°C (N=71).
Habitat: On sandy loam in Ithala Game Reserve: recorded primarily
in upland grassland (47, 1 330 m), few in mid-altitude grassland (1,
963 m), none in open shrub-woodland at lower altitude; on an altitu-
dinal transect at latitude 29–30°S on the wet E escarpment: recorded
in low numbers at 500 m (0.1 per trap) and 1 500 m (0.1), but not at
higher altitudes (1 900–2 800 m); DBRU collection records primarily
on finer-grained soils, sand (7), sandy loam (11), sandy clay loam (7),
clay (3) almost exclusively in grassland/pasture (22) with one in shrub­
land.
Food types: In Ithala Game Reserve: primarily on pig (27) and cattle dung
(19), few on horse dung (2); DBRU collection records exclusively from
cattle dung (35).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
Bioregions South Africa: Mesic Highveld Grassland (Gm), Sub-
Escarpment Grassland (Gs), E Dry Highveld Grassland (Gh) (Grass-
land Biome); Indian Ocean Coastal Belt Biome (CB); S upland margins
of Central Bushveld (SVcb) including high altitude outlier on Blouberg
2 mm
(Savanna Biome); marginal occurrence in four other bioregions.
Assessment rationale: EOO = 31 865 km2 (main range minus N outlier
on Blouberg); widespread and locally abundant on the South African
Highveld in both pastures and natural grassland, mainly in cattle dung
on farms, but also in reserves; therefore, assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be im-
proved by quantitative data on ecological associations, including effects
of loss of natural grassland through pasture improvement or commer-
cial forestry; protected in Ithala Game Reserve and Vernon Crookes
Nature Reserve (KwaZulu-Natal, South Africa).

SISYPHINI
SURICATA 6 (2020) 677

Sisyphus costatus
(Thunberg, 1818)

= Sisyphus rugosus Roth, 1851

Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type localities: As Ateuchus costatus: ‘majorem horum partem in capite

bonae spei collegi’ [amongst material mostly collected in the Cape of
Good Hope, South Africa]; S. rugosus Roth: ‘Tigré in N. Abyssinien’
[Tigray Province, N Ethiopia].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991); as S. costatus is considered to be
endemic to South Africa, the synonymy of S. rugosus Roth, 1851, is
probably an error as it would represent a different Ethiopian species
bearing a pre-occupied name.
Distribution: Only moist E Highveld and lowlands along the S coast of
South Africa; reports from Zimbabwe, Mozambique, Tanzania, Kenya
not supported by extensive DBRU collecting; also, further E and W
African records (Burundi, Democratic Republic of the Congo (DRC),
Ethiopia, Guinea, Côte d’Ivoire, Senegal) likely represent different spe-
cies (see Taxonomy).
Locality data (mean ± SD, range): Altitude: 1 326 ± 541, 17–2 274 m;
annual rainfall: 788 ± 134, 427–1 157 mm; annual temperature (max.
+ min. /2): 15.8 ± 2.0, 10.6–19.6°C (N=45).
Habitat: No adequate quantitative assessment; at Carolina: recorded in
natural Themeda grassland (22) and fallow crop fields (31), but rare in
improved Kikuyu pasture (2); DBRU collection records entirely from
2 mm
finer-grained soils: sandy loam (5), sandy clay loam (5), clay (2) in
grassland/pasture (10).
Food types: In Ithala Game Reserve: strong bias to pig dung (306) com-
pared to horse (24) and cattle dung (57); DBRU collection records all
from cattle dung (13).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Mar.).
Bioregions South Africa: Primarily, Mesic Highveld Grassland (Gm),
Sub-Escarpment Grassland (Gs) (Grassland Biome); also Indian Ocean
Coastal Belt Biome (CB) and pasture or crop fields in Eastern Fynbos-
Renosterveld (F0 6).
Assessment rationale: EOO = 142 520 km2; widespread with some toler-
ance of habitat transformation, which is often extensive across its main
Grassland Biome range (6–69%, Gm; 3–50%, Gs); marginal presence
in Fynbos Biome may be a response to habitat transformation; assessed
as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by quantitative data on soil and vegetation associations; pro-
tected in Vernon Crookes and Suikerbosrand nature reserves; also Itha-
la Game Reserve, but largely localised to degraded open Combretum
woodland at relatively low altitude.

SISYPHINI
678 SURICATA 6 (2020)

Sisyphus fasciculatus
Boheman, 1857

No synonyms
Global: LC (see IUCN Red List – LC)
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘Caffraria interiore rarius’ [at wide intervals, SE South Af-
rica].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991).
Distribution: Localised in patches of lower-altitude forest and dense
woodland; NE KwaZulu-Natal and inland along E escarpment to the
NW of South Africa.
Locality data (mean ± SD, range): Altitude: 755 ± 487, 48–2 005 m;
annual rainfall: 837 ± 78, 712–1 022 mm; annual temperature (max. +
min. /2): 19.1 ± 2.2, 12.5–22.0°C (N=31).
Habitat: No adequate quantitative study; in uMkhuze Game Reserve:
only on clay (1.2/trap), not sandy clay loam or sand; on finer-grained
soils in Ithala Game Reserve: sampled to pig dung more abundantly
in low-altitude shaded woodland (143, 758 m) than in mid-altitude
forest (100, 1 077 m) and low-altitude degraded Combretum woodland
(39, 884 m) or shrubland (32, 998 m), absent from lower (919 m)
and higher altitude grassland (1 184 m); at Wolkberg: only in lower
forest (16, 901 m) not higher forest (0, 1 355 m); on Mariepskop:
only in forest at 900–1 000 m (21), not 1 500 m or above (0); poor
DBRU collection records: sandy clay loam (2), clay (1) in forest (2) and
grassland/pasture (2).
Food types: In Ithala Game Reserve: abundant on pig (214) and cat-
tle dung (165), less so on horse (35); at Mariepskop and Wolkberg:
pig dung (17/6), cattle dung (4/10); DBRU collection records from
2 mm
human/cattle dung mixture (2) and cattle dung (1).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to Apr.).
Bioregions South Africa: Lower margins of Northern Mistbelt Forest
patches (FOz 4), Scarp Forest patches (FOz 5), (Forest Biome) and
dense woodland along the moist W edge of the Lowveld (SVl) (Savanna
Biome) where it abuts onto the Grassland and Indian Ocean Coastal
Belt Biomes.
Assessment rationale: EOO = 50 820 km2; AOO would be much smaller
due to possible finer-grained soil and observed shade association; thus,
highly susceptible to clearance of dense woody vegetation over entire
range; in forest patches (FOz 4, FOz 5), 20–25% is currently protected;
although abundant in dense woodland of vulnerable lowland savanna
units of Zululand, transformation is 22–26% (SVl 22, SVl 23) with
only 4–11% conserved; however, currently assessed as Least Concern
(LC) owing to protection in various reserves.
Conservation measures: Across the EOO, protection of forest and dense
woodland patches would be essential for the conservation of this spe-
cies; assessment of conservation status would be improved by further
quantitative data on habitat associations; protected in Ithala, uMkhuze,
Hluhluwe–iMfolozi game reserves plus Lekgalameetse, iSandlwana and
Vernon Crookes nature reserves.

SISYPHINI
SURICATA 6 (2020) 679

Sisyphus goryi
Harold, 1859

= Sisyphus hirtus Gory, 1833

Global: LC

J A S O N D J F M A M J

Type locality: S. goryi, nom nov. for S. hirtus Gory [pre-occupied name]:
‘Sénégal’ [Senegal].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991), but may represent a species com-
plex considering the geographical variation reported from across its
widespread distribution.
Distribution: Moist savanna from S to E to W Africa; Senegal, Côte d’Ivo-
ire, Nigeria, Kenya, Tanzania, Angola, Zimbabwe, Mozambique, Bot­
swana, Namibia, South Africa; also reported from Gambia, Ethiopia.
Locality data (mean ± SD, range): Altitude: 906 ± 419, 5–1 850 m; an-
nual rainfall: 666 ± 191, 246–1 415 mm; annual temperature (max. +
min. /2): 20.1 ± 2.2, 13.4–25.2°C (N=214).
Habitat: In Gauteng bushveld: relatively generalist soil associations (sand:
1 091; sandy clay loam: 797) with very strong bias to open woodland
(1 666) compared to grassland (130) and shaded thicket (92); DBRU
collection records from sand (21), sandy loam (35), sandy clay loam
(27), clay (5) in grassland/pasture (28), open shrub/woodland (45),
thicket/forest (3), crop field (1).
Food types: In Gauteng bushveld: extreme bias to pig dung (46) with
few on carrion (3); however, at Phalaborwa: equally strong bias to pig
(3 041) and cattle dung (2 770) with many also attracted to elephant
dung (584); DBRU collection records from dung of cattle (83), buffa-
lo (4), impala (3), elephant (9), rhinoceros (1), zebra (1), donkey (4),
horse (1), baboon (1), human (3).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to May); in Gauteng bushveld: late morning peak in flight activity
(10:00–12:00, Nov. to Feb.) tailing off to late afternoon (12:00–16:00)
moving to a midday peak in Mar. (12:00–14:00); seasonal peak pri-
2 mm
marily in early to mid-summer (Oct. to Jan.) tailing off steeply in late
summer (Feb. to Apr.); at Phalaborwa: more active in warm sunshine
after heavy rain (3 839) than in hot, sunny, but dry weather (1 904) or
warm, but cloudy weather after light rain (554).
Ecoregions Namibia, Botswana, Zimbabwe, Mozambique: N Namibian
Savanna Woodlands (AT1316), Kalahari Acacia-Baikiaea Woodlands
(AT0709), Zambezian and Mopane Woodlands (AT0725), Southern
African Bushveld (AT0717), Southern Miombo Woodland (AT0719);
Maputaland Coastal Forest Mosaic (AT1119).
Bioregions South Africa: Central Bushveld (SVcb), Mopane (SVmp),
Lowveld (SVl) (Savanna Biome); marginal in seven other bioregions.
Assessment rationale: EOO = 5 042 590 km2 (gross estimate); wide-
spread and locally abundant on various dung types; however, popula-
tion density would be strongly reduced by clearance of woodland (hab-
itat transformation: 0–58% (SVl); 2–49% (SVcb); 0–20% (SVmp));
nevertheless common with extremely wide range; assessed as Least
Concern (LC).
Conservation measures: Further taxonomic study is required to deter-
mine if S. goryi comprises a single widespread species or a complex of
species; protected in various national parks including: Kruger (South
Africa), Manyara (Tanzania), Tsavo (Kenya).

SISYPHINI
680 SURICATA 6 (2020)

Sisyphus impressipennis
van Lansberge, 1886

= Sisyphus transvaalensis Péringuey, 1908

= Sisyphus callosipes Arrow, 1909
Global: LC

J A S O N D J F M A M J

Type localities: S. impressipennis: ‘Humpata’ [Angola]; S. transvaalensis:

‘Transvaal (Zoutpansberg)’ [Limpopo Province, South Africa]; S. callo-
sipes: ‘German East Africa: Massailand; British Central Africa: Nyasa-
land; Katanga, 150–200 miles west of Kambove; Mashonaland: Chir-
inda’ [Tanzania, Malawi, Democratic Republic of the Congo (DRC),
Zimbabwe].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991); limited sexual dimorphism (legs).
Distribution: Localised in lowland forest and dense, shady woodland,
savanna patches from E to S central and southern Africa: N South
Africa, Botswana, Zimbabwe, Malawi, Mozambique, N Namibia, An-
gola, Tanzania; also reported from Democratic Republic of the Congo
(DRC).
Locality data (mean ± SD, range): Altitude: 987 ± 443, 13–2 005 m;
annual rainfall: 693 ± 227, 306–1 256 mm; annual temperature (max.
+ min. /2): 20.0 ± 2.8, 12.5–25.2°C (N=59).
Habitat: No adequate quantitative assessment; in dry savanna of Tswaing
Nature Reserve: mainly in shaded thicket on crater rim (22), rare in
pasture (1), absent from shrubland and open woodland; DBRU col-
lection records from sand (1), sandy loam (4) in shrubland (1), open
woodland (1), forest (18).
Food types: No quantitative assessment; DBRU collection records from
various dung types: mixture of human and cattle or buffalo (14), ba-
boon (1), elephant (4), rhinoceros (1), cattle (4), waterbuck (1).
Temporal activity: Diurnal flight activity during the summer rainy season
2 mm
(Nov. to Mar.).
Ecoregions Namibia, Zimbabwe: Dense shaded patches: N Namibian
Savanna Woodlands (AT1316) (on Kunene River), Southern Miombo
Woodlands (AT0719), Zambezian and Mopane Woodlands (AT0725),
margins of two other ecoregions.
Bioregions South Africa: Central Bushveld (SVcb) (Savanna Biome).
Assessment rationale: EOO = 2 175 705 km2; large range, but, as a shade
specialist, AOO would be restricted to thicket and forest patches often
associated with riverine, lacustrine or hill systems; would be susceptible
to thinning as well as clearance of often drier forest and woodland; in
South Africa transformation across range amounts to 2–49% (SVcb);
assessed as Least Concern (LC) since widespread and widely protected.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations; protected in
various nature reserves (Tswaing, Leeuwfontein, Roodeplaat) as well
as game reserves and national parks in South Africa (Kruger NP) and
Zimbabwe (Lake Mutirikwe GR, Victoria Falls NP).

SISYPHINI
SURICATA 6 (2020) 681

Sisyphus manni
Montreuil, 2015c

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘S. Africa: Limpopo, Kruger N.P.’ [Kruger National Park,
South Africa].
Taxonomy: Accepted macropterous species (recent status change: subge-
nus Parasisyphus Barbero, Palestrini & Zunino, 1991) in a species group
comprising macropterous and brachypterous taxa; in an unpublished
revision of southern African Sisyphus (Paschalidis 1974a), South Af-
rican material cited in error as Sisyphus alveatus Gory, 1833 whereas
cited Mozambique material is not S. manni; status of cited Zimbabwe
material uncertain (material unavailable).
Distribution: Centred in warm, moist upland grassland or moist savanna
around the N edge of the Highveld, along the Waterberg, and along the
NE escarpment; also along the moist E coast in savanna or forest: South
Africa; occurrence in Zimbabwe uncertain; occurrence in Mozambique
not supported.
Locality data (mean ± SD, range): Altitude: 1 126 ± 509, 3–1 862 m;
annual rainfall: 749 ± 118, 476–1 022 mm; annual temperature (max.
+ min. /2): 18.0 ± 2.3, 13.5–23.1°C (N=45).
Habitat: Data inconclusive; in Gauteng bushveld: recorded only on sandy
clay loam (27), not deep sand (0) in grassland (12) and open woodland
(15), not shaded thickets (0); in Leeuwfontein Nature Reserve: record-
ed in open woodland on both sand (13) and stony sandy loam (20); at
Carolina: recorded primarily in natural Themeda grassland (337); much
less common in fallow crop fields (37), rare in improved Kikuyu pas-
ture (1); limited DBRU collection records from sandy loam (2), sandy
clay loam (1) in pasture/grassland (3).
Food types: No quantitative assessment; limited DBRU collection records
from dung of cattle (3).
2 mm
Temporal activity: Macropterous; diurnal flight activity during the sum-
mer rainy season (Oct. to Mar.); on the Gauteng bushveld primarily
active in early (Oct. to Dec.) and late summer (Mar.).
Bioregions South Africa: Largely centred along the moist N and E up-
land edges of the Grassland (Mesic Highveld Grassland) (Gm); Sub-
Escarpment Grassland (Gs)) and Savanna Biomes (Central Bushveld
(SVcb); Lowveld (SVl); also Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 124 410 km2 (possibly underestimated);
apparently susceptible to disturbance of natural vegetation through
pasture improvement and cultivation; South African distribution co-
incides with often highly transformed vegetation units (e.g. 6–69%, N
Gm, N Gs; 15–58%, S SVcb, S SVl); however, widespread and protect-
ed in various reserves; assessed as Least Concern (LC).
Conservation measures: Assessment of conservation status would be
improved by further distributional and quantitative ecological data,
particularly as soil associations seem to vary geographically; protected
in Suikerbosrand, Abe Bailey and Leeuwfontein nature reserves as well
as Hluhluwe–iMfolozi Game Reserve.

SISYPHINI
682 SURICATA 6 (2020)

Sisyphus muricatus
(Olivier, 1789)

No synonyms
Global: LC
Endemic: RSA

J A S O N D J F M A M J

Type locality: As Scarabaeus muricatus: ‘l’Amérique méridionale’ [Central

America is undoubtedly an erroneous type locality for this South Afri-
can endemic].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991).
♂ Distribution: South Africa; in summer rainfall region: found in a narrow
band of cool, wet, montane conditions along E escarpment edge (av-
erage altitude: 1 556 m); in bimodal rainfall region of Eastern Cape:
found on S coastal belt (262 m).
Locality data (mean ± SD, range): Altitude: 1 094 ± 735, 16–2 274 m;
annual rainfall: 709 ± 228, 360–1 157 mm; annual temperature (max.
+ min. /2): 15.5 ± 2.4, 10.6–21.8°C (N=28).
Habitat: No adequate quantitative assessment; on a 500–2 800 m
KwaZulu-Natal gradsect (29–30°S): rare in natural grassland on sandy
clay loam at 1 500 m (0.2/trap), abundant at 1 900 m (28.7); at Car-
olina: recorded on sandy clay loam in natural grassland as a rarity (2),
absent from improved Kikuyu pasture (Kikuyu) and fallow crop fields;
recorded in forest areas but, currently, few records from forest; a single
DBRU collection record from pasture on clay.
Food types: No quantitative assessment; a few DBRU collection records
5 mm from cattle dung (3).
Temporal activity: Diurnal flight activity coinciding with summer (in the
N) or spring/autumn rainfall peaks (in the S) (Sept. to Apr.).
Bioregions South Africa: In the N and E: moist E montane vegetation
units, Mesic Highveld Grassland (Gm), Drakensberg Grassland (Gd),
also Sub-Escarpment Grassland (Gs) (Grassland Biome); in the S: E
vegetation units, Sandstone Fynbos (FFs), Shale Fynbos (FFh) (Fynbos
Biome).
Assessment rationale: EOO = 24 500 km2; extensive range in the N co-
incides with E edge of Grassland Biome, but unclear if it is a grassland
specialist; thus, unknown if Eastern Cape range represents range expan-
sion into pastures resulting from clearance of woody vegetation; range
often coincides with vulnerable or endangered vegetation units result-
ing from tree plantations, cultivation and urbanisation (Gd 3–16%;
montane Gm 23–52%, Gs 10–25%; Garden Route FFs 28–33%, FFh
50%); negative influence of pasture improvement and transformation
possible, but poorly supported; currently assessed as Least Concern
(LC) owing to widespread range.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on ecological associations, including
possible effects of pasture improvement and habitat transformation;
protected within the extensive uKhahlamba Drakensberg Park (World
Heritage Site) along the E and NE Lesotho border.

SISYPHINI
SURICATA 6 (2020) 683

Sisyphus nanniscus
Péringuey, 1901

= Sisyphus ocellatus Reiche subsp. nanniscus Péringuey, 1901, sensu Haaf,

1955
Global: LC

J A S O N D J F M A M J

Type locality: ‘Natal (Durban)’ [KwaZulu-Natal, South Africa].

Taxonomy: Recently removed from published synonymy with Sisyphus
costatus Thunberg, 1818, and Sisyphus seminulum Gerstaecker, 1871,
to be re-established as a valid species (Montreuil 2016) (recent status
change: subgenus Sisyphus Latreille, 1807); synonym (Haaf 1955) rep-
resents a misidentification of the species as a subspecies of S. ocellatus
Reiche, 1847.
Distribution: Widespread across hot, woodland savanna from S to E
Africa: South Africa, Botswana, Zimbabwe, Zambia, Malawi, Mozam-
bique, Tanzania, Kenya.
Locality data (mean ± SD, range): Altitude: 523 ± 479, 0–1 613 m; an-
nual rainfall: 811 ± 227, 383–2 097 mm; annual temperature (max. +
min. /2): 20.7 ± 1.9, 13.8–25.2°C (N=74).
Habitat: In uMkhuze Game Reserve: relative soil generalist with slight
bias to finer-grained soils, sand (36.8/trap), duplex soil (sand over clay)
(45.5), sandy clay loam (57.0), clay (73.4) with 97% (2 255 out of
2 325) sampled under shade; in Hluhluwe Game Reserve: recorded in
long grass on cloudy days; DBRU collection records from sand (15),
sandy loam (10), sandy clay loam (5), clay (1) in forest/thicket (10),
2 mm
woodland (10), open shrub/woodland (8), grassland/pasture (4).
Food types: In Ithala Game Reserve: bias to dung of pig (2 902) and
cattle (1 300) compared to horse (144); DBRU collection records from
various dung types: human (1), baboon (1), human/cattle mix (6), ele-
phant (1), rhinoceros (3), zebra (1), horse (1), warthog (1), cattle (20),
buffalo (2), wildebeest (2), waterbuck (1), impala (1).
Temporal activity: Diurnal flight activity throughout much of the year
due to warm lowland range, especially at the coast (Aug. to May).
Ecoregions Zimbabwe, Mozambique: Maputaland Coastal Forest Mosa-
ic (AT0119), edge of the dry lowland Zambezian and Mopane Wood-
lands (AT0725) where it abuts onto moister, higher altitude ecoregions,
particularly Southern Miombo Woodlands (AT0719).
Bioregions South Africa: Centred on Lowveld (SVl) (Savanna Biome),
Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 919 450 km2 (gross estimate); would be
threatened by clearance of dense woodland and forest due to shade spe-
cialisation; in South Africa: transformation in main range (SVl) varies
from 0 to 58%; however, widespread along E seaboard of Africa in hot,
shaded, savanna vegetation; therefore, assessed as Least Concern (LC).
Conservation measures: None recommended; protected in Ithala,
uMkhuze and Hluhluwe–iMfolozi game reserves as well as moister,
densely wooded parts of the Kruger National Park (South Africa); also
Meru Game Reserve (Kenya).

SISYPHINI
684 SURICATA 6 (2020)

Sisyphus neobornemisszanus
Daniel & Davis, 2016

= Sisyphus bornemisszanus Endrödi, 1983 (pars)

Global: NT

J A S O N D J F M A M J

Type localities: S. neobornemisszanus: ‘Zululand, St. Lucia, 28.22S–32.25E’;

S. bornemisszanus: ‘Zululand, St. Lucia, 28.22S–32.25E’ [KwaZulu-
Natal, South Africa].
Taxonomy: Recently re-described as S. neobornemisszanus since S. bor-
nemisszanus was represented by a type series comprising two differ-
ent species; the holotype and some paratypes were synonymous with
Sisyphus sordidus Boheman, 1857; only the allotype and remaining
paratypes represented the species re-described as S. neobornemiszanus
(recent status change: subgenus Parasisyphus Barbero, Palestrini & Zu-
nino, 1991).
Distribution: Coastal-fringing dune forest in the Maputaland Centre of
Endemism and further N along the Mozambique coast; South Africa,
Mozambique.
Locality data (mean ± SD, range): Altitude: 8 ± 18, 0–64 m; annual
rainfall: 844 ± 103, 738–1 006 mm; annual temperature (max. + min.
/2): 22.3 ± 0.5, 22.7–23.6°C (N=22).
Habitat: On deep sand in Maputo Special Reserve: primarily in cooler
coastal-fringing dune forest (34.8/trap) rather than warmer inland
sand forest patches (large: 0.4, medium: 3.6, small: 1.9), absent from
grassland patches (0); across a deep sand, post-mining chronosequence
at Richards Bay: fewer in deep shade of early succession woodland (7)
and natural dune forest (3) than in less-shaded later succession open
2 mm
understorey woodland (75); limited DBRU collection records in dune
forest (5).
Food types: No quantitative assessment; sampled to pig dung.
Temporal activity: Diurnal flight activity during most of the year due to
warm coastal range (Aug. to May).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119);
Southern Zanzibar-Inhambane Coastal Forest Mosaic (AT0128) (not
shown on map).
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB).
Assessment rationale: EOO = 4 500 km2; AOO smaller, restricted to
dense woodland and forest patches; occurs along a coastline that has,
in places, been transformed by mining; although it shows a larger range
than the Maputaland endemic, Scarabaeus bornemisszai zur Strassen,
which has been assessed as Near Threatened (NT), it is also assessed as
NT owing to threats of coastal development; it would even qualify for
an IUCN threat category approaching Vulnerable (VU) on the basis of
small range and vegetation specialisation.
Conservation measures: Although quantitative data on soil and dung
type associations would improve the assessment, conservation would
clearly be dependent on the presence of regenerating woodland and/
or protection of dune forest fringing the coast of NE KwaZulu-Natal
(South Africa) and SE Mozambique; currently protected in iSimanga-
liso Wetland Park (World Heritage Site) (South Africa) and Maputo
Special Reserve (Mozambique).

SISYPHINI
SURICATA 6 (2020) 685

Sisyphus oralensis
Daniel & Davis, 2016

No synonyms
Global: LC

J A S O N D J F M A M J

Type locality: ‘KZN, Richard’s Bay’ [Richards Bay, KwaZulu-Natal,

South Africa].
Taxonomy: Accepted species (recent status change: subgenus Sisyphus La-
treille, 1807) referenced as Sisyphus sp. y in the unpublished taxonomic
revision of Paschalidis (1974a) and various ecological papers on the
dung fauna of coastal NE KwaZulu-Natal.
Distribution: Moist to wet forest and shaded thicket on deep sands bor-
dering SE African coast: NE South Africa, SE Mozambique.
Locality data (mean ± SD, range): Altitude: 28 ± 56, 0–273 m; annual
rainfall: 853 ± 112, 634–1 010 mm; annual temperature (max. + min.
/2): 22.0 ± 0.6, 20.0–23.3°C (N=32).
Habitat: Across a post-mining, rehabiltation gradient on coastal dunes at
Richards Bay: grassland (0.00), early woodland (0.13), older woodland
(2.46), natural dune forest (2.12); on deep sand in Maputo Special
Reserve: absent from grassland (0), but present in all forest categories:
large sand forest patches (12.7), medium-sized patches (21.2), small
patches (11.1), cooler coastal dune forest (5.0); DBRU collection re-
cords exclusively from forest (9) or Eucalyptus plantation (1).
Food types: No quantitative assessment; DBRU collection records from
dung of cattle (3) and human/cattle dung mix (1).
Temporal activity: Diurnal flight activity; recorded both during the sum-
mer rainy season (Oct. to Feb.) and the dry season (May to Aug.).
Ecoregions Mozambique: Maputaland Coastal Forest Mosaic (AT0119).
2 mm
Bioregions South Africa: N Indian Ocean Coastal Belt Biome (CB); SE
Lowveld (SVl) (Savanna Biome).
Assessment rationale: EOO = 18 105 km2; although sand is widespread
along the SE coastline, AOO would be much smaller due to specialised
association with vegetation offering shade; quantitative data show that
clearance of forest and thicket is detrimental to population density;
however, S. oralensis is currently assessed as Least Concern (LC) by vir-
tue of large EOO and occurrence at various localities.
Conservation measures: Assessment of conservation status would be im-
proved by further quantitative data on ecological associations and fur-
ther information on effects of the clearance of forest and dense wood-
land; protected in Maputo Elephant Reserve (Mozambique), Tembe
Elephant Reserve and iSimangaliso Wetland Park (World Heritage Site)
(South Africa).

SISYPHINI
686 SURICATA 6 (2020)

Sisyphus perissinottoi
Montreuil, 2015c

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘RSA, EC, Amsterdamhoek (Port Elizabeth)’ [Eastern

Cape, South Africa].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991) in a species group comprising
macropterous and brachypterous taxa.
Distribution: Centred on drier areas, E bimodal rainfall region, Eastern
Cape, South Africa.
Locality data (mean ± SD, range): Altitude: 608 ± 494, 123–1 100 m;
annual rainfall: 353 ± 88, 246–426 mm; annual temperature (max. +
min. /2): 16.4 ± 1.8, 14.8–18.0°C (N=4).
Habitat: No quantitative assessment; not known.
Food types: No quantitative assessment; not known.
Temporal activity: Macropterous; diurnal flight activity; currently record-
ed only during the autumn peak of the bimodal rainfall spring/autumn
rainy season (Feb.).
Bioregions South Africa: Albany Thicket Biome (AT), Rainshadow Val-
ley Karoo (SKv) (Succulent Karoo Biome).
Assessment rationale: EOO = 9 685 km2 (underestimated); only recently
recognised as a valid species, but known by few localities with no asso-
ciated ecological data; the low number of known localities (<5) within
a small EOO would qualify for an IUCN threat category of Vulnerable
(VU), but this pattern is considered to represent a collection artefact;
2 mm
therefore, currently, assessed as Data Deficient (DD).
Conservation measures: Assessment of conservation status will only be
possible after a quantitative survey has been conducted in the bimodal
rainfall region of the Eastern Cape to determine the full EOO, AOO
and ecological associations; protected in Addo Elephant National Park.

SISYPHINI
SURICATA 6 (2020) 687

Sisyphus sordidus
Boheman, 1857

= Sisyphus natalensis Balthasar, 1968

= Sisyphus bornemisszanus Endrödi, 1983 (pars)
Global: LC (See IUCN Red List – LC as S. natalensis)

J A S O N D J F M A M J

Type localities: S. sordidus: ‘Caffraria interiore’ [inexact, may be interi-

or of SE Africa]; S. natalensis: ‘Süd-Afrika, Nord-Natal. Umgebung
von Santa Lucia Lake’; S. bornemisszanus (pars): ‘Zululand, St. Lucia,
28.22S–32.25E’ [both, environs of Lake St Lucia, KwaZulu-Natal,
South Africa].
Taxonomy: Accepted species (recent status change: subgenus Parasisyphus
Barbero, Palestrini & Zunino, 1991); as regards the synonyms, only
part of the type series of S. bornemisszanus (holotype and five paratypes)
is synonymous with S. natalensis (Daniel et al. 2016); the allotype and
five other paratypes of S. bornemisszanus have been re-described as S.
neobornemisszanus Daniel & Davis, 2016 (see species account).
Distribution: Primarily, moist lowland savanna, particularly on the Mo-
zambique Coastal Plain, but also, along the Soutpansberg and Water-
berg: South Africa, Zimbabwe, Mozambique.
Locality data (mean ± SD, range): Altitude: 225 ± 341, 0–1 148 m; an-
nual rainfall: 803 ± 191, 362–1 254 mm; annual temperature (max. +
min. /2): 22.0 ± 1.3, 18.8–25.2°C (N=59).
Habitat: In uMkhuze Game Reserve: extreme bias to deep sand (136.8)
compared to sand over clay (2.3), sandy clay loam (0.0) and clay (0.1);
across a vegetation recovery gradient at Richards Bay: extreme bias to
grassland (706) compared to young woodland (4), older woodland
(1) and natural dune forest (0); partly supported by DBRU collection 2 mm
records from sand (12), sandy loam (2) in grassland (15), open shrub/
woodland (7), forest/thicket (7).
Food types: No quantitative assessment; DBRU collection records from
various dung types including cattle (7), impala (1), horse (2), elephant
(1), rhinoceros (2), human (3), human/cattle or buffalo mix (4).
Temporal activity: Diurnal flight activity, primarily during the summer
rainy season (Oct. to May).
Ecoregions Zimbabwe, Mozambique: E Southern Miombo Woodlands
(AT0719), Maputaland Coastal Forest Mosaic (AT0119).
Bioregions South Africa: Centred on Indian Ocean Coastal Belt Biome
(CB), also, Lowveld (SVl) plus Waterberg in Central Bushveld (SVcb),
marginal in Mopane (SVmp) (Savanna Biome),.
Assessment rationale: EOO = 326 650 km2 (N extent of EOO probably
underestimated); AOO presumably smaller according to distribution
of sand and open vegetation; assessed as Least Concern (LC) owing to
large range, widespread occurrence of sand along SE coast and associ-
ation with open vegetation, which would suggest tolerance of habitat
tranformation.
Conservation measures: Assessment of conservation status would be
improved by quantitative data on dung type associations although ap-
parently a dung type generalist; protected in uMkhuze Game Reserve
and iSimangaliso Wetland Park (World Heritage Site) (South Africa),
Gorongosa National Park and Maputo Special Reserve (Mozambique).

SISYPHINI
688 SURICATA 6 (2020)

Sisyphus splendidus
Montreuil, 2015b

No synonyms
Global: DD

J A S O N D J F M A M J

Type locality: ‘Namibie,...., Otavi mts, Farm Gauss’ [NW Namibia].

Taxonomy: Accepted species (recent status change: subgenus Sisyphus
Latreille, 1807).
Distribution: Limited locality data suggest a range in mostly dry savanna
regions across mountains and plains: NW Namibia, SW Angola.
Locality data (mean ± SD, range): Altitude: 1 495 ± 370, 1 227–1 917 m;
annual rainfall: 563 ± 174, 389–737 mm; annual temperature (max. +
min. /2): 18.7 ± 1.4, 17.2–19.9°C (N=3).
Habitat: Not known.
Food types: Not known.
Temporal activity: Not known, but presumably diurnal flight activity like
other sisyphines; recorded during the late summer rainy season (Dec.,
Feb.).
Ecoregions Namibia: NE Kalahari Xeric Savanna (AT1309), N Namib-
ian Savanna Woodlands (AT1316).
Assessment rationale: EOO = 18 600 km2 (estimated); only recently
described as a new species; known by 18 individuals from only three
localities, but recorded across a fairly wide range that has probably been
underestimated; limited records probably a collecting artefact; if so,
would deserve Least Concern (LC) status, but at present assessed as
Data Deficient (DD).
Conservation measures: To assess conservation status, a quantitative
survey of N Namibia and SW Angola is required to determine EOO,
AOO and ecological associations; not currently known from any pro-
tected area.

2 mm

SISYPHINI
SURICATA 6 (2020) 689

Sisyphus umbraphilus
Daniel & Davis, 2016

No synonyms
Global: DD
Endemic: RSA

J A S O N D J F M A M J

Type locality: ‘KZN, Umfolozi’ [iMfolozi Game Reserve, KwaZulu-

Natal, South Africa].
Taxonomy: Accepted species (recent status change: subgenus Sisyphus
Latreille, 1807) referenced as Sisyphus sp. x in the unpublished taxo-
nomic revision of Paschalidis (1974a) and as Sisyphus sp. in ecological
publications for Gauteng.
Distribution: A few scattered records in forest or shaded thickets in moist
savanna regions: South Africa.
Locality data (mean ± SD, range): Altitude: 519 ± 487, 126–1 150 m;
annual rainfall: 744 ± 124, 636–904 mm; annual temperature (max. +
min. /2): 20.3 ± 1.4, 18.9–21.9°C (N=4).
Habitat: In Gauteng: recorded on deep sand (11) not sandy clay loam (0)
and in shaded thickets (11) not open woodland (0) or grassland (0); in
Ithala Game Reserve: recorded in moist, dense lowland woodland (10);
one DBRU collection record from forest.
Food types: In Ithala Game Reserve: bias shown to omnivore dung (pig:
8) compared to herbivore dung, monogastric (horse: 0), ruminant (cat-
tle: 2).
Temporal activity: Diurnal flight activity during the summer rainy season
(Oct. to Jan.); in Gauteng: activity only recorded during the early rainy
season (Nov. to Dec.).
Bioregions South Africa: Central Bushveld (SVcb), Lowveld (SVl) (Sa-
2 mm
vanna Biome); marginal in one other bioregion.
Assessment rationale: EOO = 21 880 km2 (undoubtedly underestimat-
ed); AOO would be much smaller as it is certainly a shade vegetation
specialist; possibly also a soil and dung type specialist; assessed as Data
Deficient (DD) owing to the limited number of records, which are
probably a collecting artefact reflecting its specialised habits and low
population density.
Conservation measures: To adequately assess conservation status, it is
necessary to conduct a quantitative survey of thickets in savanna re-
gions of South Africa; this survey should test soil and food associations
more thoroughly and determine level of threat through clearance of
dense woodland and forest; protected in Ithala and Hluhluwe-iMfolozi
game reserves.

SISYPHINI
690 SURICATA 6 (2020)

Poorly known species

This section comprises a number of poorly known, but from type material. They have, therefore, not yet been val-
currently valid species and their synonyms. They are most- idly matched to available non-type reference specimens.
ly, wholly or partly described from South Africa, Botswa- Distributions are based on citations in Schoolmeesters
na or Namibia. For the most part, they are known only (2017), but have not been critically assessed.

ATEUCHINI

Pedaria alternans
Waterhouse, 1890
No synonyms
Type locality: ‘S. Africa’ [?South Africa].
Distribution: South Africa, Ethiopia.
Notes: Not yet reliably matched to available southern African material.

Pedaria aspera
Péringuey, 1901
No synonyms
Type locality: ‘Cape Colony (Barkly West)’ [Northern Cape, South Africa].
Distribution: South Africa.
Notes: Not yet reliably matched to available South African material.

Pedaria conformis
Péringuey, 1901
= Pedaria somalica Balthasar, 1939a
Type localities: P. conformis: ‘Transvaal (Klerksdorp, Potchefstroom), Natal (Durban)’ [South Africa: North-West Prov-
ince and KwaZulu-Natal]; P. somalica: ‘Somalia, Obbia’.
Distribution: South Africa, Somalia.
Notes: Not yet reliably matched to available South African material; synonymy of P. conformis and P. somalica
based on identical aedeagi possessed by male types (Josso & Prévost 2003).

COPRINI

Metacatharsius pusio
(Kolbe, 1908)
No synonyms
Type locality: As Catharsius pusio: ‘Kalahari: Sekgoma-Khakea;....Khakea-Kang’ [Botswana: Sekumo to Kang].
Distribution: Namibia, Botswana.
Notes: Not yet reliably matched to available Botswana material; however, see species account for M. freyi (Fer-
reira 1964b).

Metacatharsius rugosipennis
Frey, 1975
No synonyms
Type locality: ‘Betchuanaland, Tsano’ [?Tshane, Botswana].
Distribution: Botswana.
Notes: Not yet reliably matched to available Botswana material; however, see species account for M. ferreirae
(Balthasar, 1965b).
SURICATA 6 (2020) 691

Metacatharsius transvaalensis
Balthasar, 1968
No synonyms
Type locality: ‘Süd-Afrika, Transvaal (Ost)’ [?Mpumalanga Province, South Africa].
Distribution: N South Africa.
Notes: Not yet reliably matched to available southern African material.

ONTHOPHAGINI

Caccobius inconspicuus
(Fahraeus, 1857)
No synonyms
Type locality: As Onthophagus inconspicuus: ‘in terra Natalensi’ [KwaZulu-Natal, South Africa].
Distribution: South Africa, Malawi, Mozambique, Democratic Republic of the Congo (DRC).
Notes: Not matched to available southern African material.

Jossonthophagus asperriformis
Dierkens & Moretto, 2017
= Onthophagus incertus Péringuey, 1901
= Onthophagus dubius d’Orbigny, 1902
= Onthophagus asperrimus d’Orbigny, 1902
Type localities: J. asperriformis: ‘Ovampoland (Omrramba)’ [Omuramba, Ovamboland, Namibia] nom nov. for both
O. dubius and O. incertus (pre-occupied names); O. asperrimus: ‘Afrique orientale allemande’ [Tanzania],
lectotype: same locality.
Distribution: South Africa, Botswana, Namibia, Malawi, Zambia, Zimbabwe, Mozambique, Tanzania.
Notes: Formerly assigned to Onthophagus Group 14; revised by Dierkens et al. (2017).

Jossonthophagus dispar
(Péringuey, 1901)
No synonyms
Type locality: As Onthophagus dispar: ‘Natal (Malvern)’ [Queensburgh (Durban), KwaZulu-Natal, South Africa]; lec-
totype: same locality.
Distribution: Democratic Republic of the Congo (DRC), Zambia, Zimbabwe, Namibia, now validated; Angola, Tan-
zania, Mozambique require validation as well as the South African type locality.
Notes: Formerly assigned to Onthophagus Group 14; revised by Dierkens et al. (2017).

Onthophagus aschenborni
Frey, 1975b
No synonyms
Type locality: ‘Nordtransvaal, Magoebaskloof ’ [Magoebaskloof, Limpopo Province, South Africa].
Distribution: South Africa.
Notes: Not yet reliably matched to available Group 2 South African material; cited as close to O. modestus Har-
old, 1862 (Frey 1975b).

Onthophagus asimilis
Péringuey, 1901
No synonyms
Type locality: ‘Bechuanaland’ [Botswana].
Distribution: Botswana, Namibia.
Notes: Not matched to available Group 16 southern African material.

Onthophagus aspericeps
d’Orbigny, 1908
No synonyms
Type locality: ‘Sud-Ouest africain allemand: Okahandya’ [Okahandja, Namibia].
692 SURICATA 6 (2020)

Distribution: Namibia, South Africa.

Notes: Described from a single individual; not reliably matched to available Group 22 Namibian material; rela-
tionships to O. convexus d’Orbigny, 1908, need to be determined (see species account).

Onthophagus betschuanus
Frey, 1975
No synonyms
Type locality: ‘S.-Afrika, Betschuanaland, Kukong’ [Botswana].
Distribution: Botswana.
Notes: Not matched to available Botswana material in either Groups 2 or 22.

Onthophagus biconifer
d’Orbigny, 1905
No synonyms
Type locality: ‘Transvaal’ [N South Africa].
Distribution: South Africa, Democratic Republic of the Congo (DRC).
Notes: Not matched to available Group 2 southern African material.

Onthophagus burchelli
d’Orbigny, 1908
No synonyms
Type localities: ‘Bechouana: Griquatown...Sud-Ouest africain allemand: Windhoek...Okahandja’ [Griquatown, North-
ern Cape, South Africa; Namibia: Windhoek and Okahandja].
Distribution: South Africa (NOT Botswana), Namibia.
Notes: Not matched to available Group 24 southern African material; recently recognised as a synonym of
Onthophagus leucopygus Harold, 1867 (P. Moretto, pers. comm.).

Onthophagus confertus
Péringuey, 1908
= Onthophagus pusio Péringuey, 1901
Type locality: O. confertus nom nov. for O. pusio Péringuey (pre-occupied name): ‘Natal (Maritzburg)’ [Pietermaritz-
burg, KwaZulu-Natal, South Africa].
Distribution: Zimbabwe, South Africa, Mozambique.
Notes: Not matched to available Group 19 South African material; needs to be compared with O. pusio Fahrae-
us, 1857.

Onthophagus coptorhinodes
Péringuey, 1901
No synonyms
Type locality: ‘Natal (Estcourt)’ [Estcourt, KwaZulu-Natal, South Africa].
Distribution: NE South Africa.
Notes: Not matched to available Group 3 South African material.

Onthophagus criniger
d’Orbigny, 1904
= Onthophagus crinitus d’Orbigny, 1902
See species account for O. axillaris Boheman, 1860.
Notes: Described from a single individual; not matched to available Group 16 Botswana material.

Onthophagus declivicollis
d’Orbigny, 1902
No synonyms
Type locality: ‘Natal’ [KwaZulu-Natal, South Africa].
SURICATA 6 (2020) 693

Distribution: NE South Africa, Mozambique.

Notes: Not matched to available Group 9 South African material; needs to be compared with O. asperulus d’Or-
bigny, 1905, as a possible senior name.

Onthophagus dedecor
Wallengren, 1881
= Onthophagus nigrescens d’Orbigny, 1902
Type localities: O. dedecor: ‘Transvaalia’ [N South Africa], O. nigrescens: ‘Zambèze....Transvaal’ [inexact: southern Africa,
N South Africa].
Distribution: Zimbabwe, South Africa.
Notes: Not matched to available Group 2 South African material; types need to be compared with O. aerugino­
sus Roth, 1851, and a close unnamed relative from the Waterberg, South Africa.

Onthophagus gnu
Frey, 1955
No synonyms
Type locality: ‘Süd-West-Afrika Gobabis’ [Gobabis, Namibia].
Distribution: Namibia.
Notes: Described from a single male individual, not matched to available Group 11 Namibian material; group
assignment requires validation.

Onthophagus granulum
d’Orbigny, 1904
= Onthophagus granum d’Orbigny, 1902
Type locality: O. granulum nom nov. for O. granum (pre-occupied name): ‘Mozambique...: Lourenço Marquez’ [Mapu-
to, Mozambique].
Distribution: N South Africa, Mozambique.
Notes: Matched to Group 2 South African material from Nylsvlei (Limpopo Province), but requires further
validation by comparison with type.

Onthophagus guttatus
Boheman, 1860
No synonyms
Type locality: ‘juxta lacum N’Gami’ [near Lake Ngami, N Botswana].
Distribution: Botswana, Angola.
Notes: Not placed in a group by d’Orbigny (1913); not matched to available Botswana material.

Onthophagus importunus
Péringuey, 1901
No synonyms
Type locality: ‘Ovampoland (Humbe)’ [Ovamboland, Namibia].
Distribution: Namibia, Angola.
Notes: Not reliably matched to available Namibian Group 2 material.

Onthophagus insulsus
Péringuey, 1901
No synonyms
Type locality: ‘Natal (Eshowe)’ [Eshowe, KwaZulu-Natal, South Africa].
Distribution: South Africa
Notes: Not matched to available Group 7 South African material; needs to also be compared with O. beiranus
Péringuey, 1908 (see species page), O. interstitialis Fahraeus, 1857, and two unnamed species from E
forests of South Africa of similar body size.
694 SURICATA 6 (2020)

Onthophagus lugens
Fahraeus, 1857
No synonyms
Type locality: ‘juxta fluvium Gariep’ [near Orange River, South Africa].
Distribution: South Africa, Zimbabwe.
Notes: Not matched to available Group 9 South African material; needs to be assessed as a possible synonym of
O. setosus Fahraeus, 1857, as suggested by Péringuey (1901).

Onthophagus macrothorax
d’Orbigny, 1902
No synonyms
Type locality: ‘Lac Ngami’ [Lake Ngami, N Botswana].
Distribution: Botswana, Democratic Republic of the Congo (DRC).
Notes: Not matched to available Group 16 Botswana material.

Onthophagus marshalli
d’Orbigny, 1908
No synonyms
Type localities: ‘Rhodésia: Salisbury...Bulawayo...Transvaal: Waterberg District…Sud-Ouest africain allemand: Otjosondu-
Kub’ [various localities: South Africa, Zimbabwe, Namibia].
Distribution: N South Africa, Zimbabwe, Namibia
Notes: Not matched to available Group 9 South African material; does not resemble any other species.

Onthophagus merus
Péringuey, 1901
No synonyms
Type localities: ‘Cape Colony (Kimberley), Southern Rhodesia (Lesappi River)’ [Northern Cape, South Africa; Zim­
babwe].
Distribution: South Africa, Zimbabwe, Botswana.
Notes: Matched to available Group 31 South African material from Gauteng and to Zimbabwean material from
Hwange National Park, but requires validation by comparison with type material.

Onthophagus minutulus
Harold, 1875
No synonyms
Type locality: ‘Kuruman in Südafrika’ [Kuruman, South Africa].
Distribution: South Africa, Botswana.
Notes: Not matched to available Group 3 southern African material.

Onthophagus modestus
Harold, 1862
= Onthophagus pusillus Fahraeus, 1857
Type locality: ‘juxta fluvium Gariep’ [near Orange River, South Africa].
Distribution: South Africa.
Notes: A nom nov. for pre-occupied name of O. pusillus Fabricius, 1801; not matched to available Group 2
South African material.

Onthophagus nanus
Harold, 1878
= Onthophagus decipiens Péringuey, 1901
Type localities: O. nanus: ‘Ostafrika’ [East Africa]; O. decipiens: ‘Mossamedes, Natal (Malvern)’ [Namibe, Angola;
Queensburgh, KwaZulu-Natal, South Africa].
Distribution: Kenya, Malawi, Angola, Zimbabwe, Mozambique, South Africa, Namibia.
Notes: Not yet reliably matched to available Group 11 material from South Africa, Botswana and Namibia;
synonymy of O. decipiens requires validation.
SURICATA 6 (2020) 695

Onthophagus nudus
d’Orbigny, 1908
No synonyms
Type locality: ‘Natal: Malvern’ [Queensburgh, KwaZulu-Natal, South Africa].
Distribution: South Africa.
Notes: Not yet reliably matched to available coastal Group 7 South African material.

Onthophagus otjivarongus
Balthasar, 1967
No synonyms
Type locality: ‘Süd-West-Afrika, Abachaus, Otjivarongo-District’ [Abachaus, Namibia].
Distribution: Namibia.
Notes: Described from female individuals, not matched to available coastal Group 19 Namibian material, but
type should be compared to that of Onthophagus kochi Frey, 1957 (see O. kochi species account).

Onthophagus ovigranosus
Frey, 1971
No synonyms
Type locality: ‘Tongaat, Natal’ [KwaZulu-Natal, South Africa].
Distribution: South Africa.
Notes: Described from a single female; not yet reliably matched to available Group 22 southern African material
from E coast.

Onthophagus pusio
Fahraeus, 1857
No synonyms
Type locality: ‘in terra Natalensi’ [KwaZulu-Natal, South Africa].
Distribution: South Africa.
Notes: Not placed in a group by d’Orbigny (1913); not matched to available South African material; cited as
different to O. pusio Péringuey, 1901 (Péringuey 1908).

Onthophagus rubens
d’Orbigny, 1902
= Onthophagus exiguus Péringuey, 1892
= Onthophagus kalaharicus Kolbe, 1908
Type localities: O. rubens, nom. nov. for O. exiguus (pre-occupied name): type locality not stated; O. kalaharicus: ‘Kala-
hari: Lookaneng-Severelela’ [Botswana].
Distribution: Namibia, South Africa, Botswana, Zimbabwe; also cited from Lesotho, but presumably an error.
Notes: Not yet reliably matched to available Group 31 southern African material from N Namibia, N Botswana,
N South Africa.

Onthophagus rufovirens
d’Orbigny, 1904
No synonyms
Type locality: ‘Natal: Durban’ [KwaZulu-Natal, South Africa].
Distribution: South Africa.
Notes: Not matched to available Group 2 South African material.

Onthophagus semigraniger
d’Orbigny, 1905
No synonyms
Type locality: ‘Sud-Ouest africain: Windhoek’ [Namibia].
Distribution: Namibia.
Notes: Not matched to available Group 13 Namibian material.
696 SURICATA 6 (2020)

Onthophagus senescens
Péringuey, 1908
= Onthophagus seniculus Péringuey, 1901
Type locality: O. senescens, nom. nov. for O. seniculus (pre-occupied name): ‘Ovampoland (Omrramba)’ [Ovamboland,
Namibia].
Distribution: Namibia.
Notes: Not matched to available Group 16 Namibian material.

Onthophagus setosus
Fahraeus, 1857
= Onthophagus pedestris Fahraeus, 1857
= Onthophagus scabrosus Fahraeus, 1857
= Onthophagus cretus Péringuey, 1901
Type localities: O. setosus: ‘prope fluvium Limpopo’ [near Limpopo River, southern Africa]; O. pedestris: ‘juxta fluvium
Gariep’ [near Orange River, South Africa]; O. scabrosus: ‘juxta fluvium Gariep’; O. cretus: ‘Natal (Durban,
Umkomaas River), Ovampoland (Humbe)’ [Durban, KwaZulu-Natal, South Africa; Ovamboland, Namib-
ia].
Distribution: South Africa, Namibia, Angola, Mozambique.
Notes: Not matched to available Group 9 southern African material; validation of species identities required; syn-
onymy of O. cretus and citation from Namibia considered errors; see O. cretus species account.

Onthophagus simoni
d’Orbigny, 1902
No synonyms
Type locality: ‘Hamman’s Kraal’ [Hammanskraal, Gauteng, South Africa].
Distribution: South Africa, Zimbabwe, Democratic Republic of the Congo (DRC), Tanzania.
Notes: Not matched to available Group 2 southern African material; cited as close to O. modestus Harold, 1862.

Onthophagus stenocerus
Harold, 1867
= Onthophagus gracilicornis Fahraeus, 1857
Type localities: O. stenocerus, nom nov. for O. gracilicornis (pre-occupied name): ‘prope fluvium Limpopo’ [near Limpopo
River, southern Africa].
Distribution: South Africa, Mozambique.
Notes: Not placed in a group by d’Orbigny (1913); not matched to available material from southern Africa.

Onthophagus sugillatus
Klug, 1855
= Onthophagus mactatus Klug, 1855
= Onthophagus hybridus Fahraeus, 1857
= Onthophagus intermedius Fahraeus, 1857
Type localities: O. sugillatus: ‘Inhambane’ [Mozambique coast]; O. mactatus: ‘Inhambane’; O. hybridus: ‘prope fluvium Lim-
popo’ [near Limpopo River, southern Africa]; O. intermedius: ‘prope fluvium Limpopo’.
Distribution: Mozambique, Namibia, Botswana, South Africa, Lesotho.
Notes: Not reliably matched to available material from southern Africa; Fahraeus synonyms and cited range
require validation; at least 10, currently, unnamed species recognised from Group 31 in southern African
reference material.

Onthophagus talpa
Fahraeus, 1857
No synonyms
Type locality: ‘juxta fluvium Gariep’ [near Orange River, South Africa].
Distribution: South Africa, Zimbabwe, Mozambique, Namibia.
Notes: Not placed in a group by d’Orbigny (1913); not matched to available southern African material.
SURICATA 6 (2020) 697

Onthophagus temporalis
d’Orbigny, 1902
No synonyms
Type locality: ‘Afrique australe’ [southern Africa].
Distribution: South Africa.
Notes: Described from a single female; not matched to available Group 31 southern African material.

Onthophagus ventrosus
d’Orbigny, 1905
No synonyms
Type locality: ‘Transvaal’ [N South Africa].
Distribution: South Africa.
Notes: Described from a single individual; not matched to available Group 18 South African material.

Onthophagus zumpti
Frey, 1954
No synonyms
Type locality: ‘Willowmoore Cape Provinz’ [Willowmore, Eastern Cape, South Africa].
Distribution: South Africa.
Notes: Described from three female individuals; not matched to available Group 30 South African material;
should be compared with material of O. cameloides d’Orbigny, 1900.
698 SURICATA 6 (2020)

SYNTHESIS
A. SYSTEMATICS

Higher classification of the subfamily Scarabaeinae is in dis- Topology of the basal dichotomies in the first (Monaghan
array due to the degree of polyphyly in the old tribal units et al. 2007) and most recent global molecular phyloge-
comprising the Ateuchini, Canthonini and the possibly nies (Tarasov & Dimitrov 2016) differ radically. Howev-
older (Monaghan et al. 2007) or possibly more recent tribal er, some overall patterns are similar. In the phylogeny of
unit (Tarasov & Dimitrov 2016) comprising the Coprini. Monaghan et al. (2007), colour coding of basal linkages
Although some new tribal units are proposed in the most between clades (Davis et al. 2017) shows no breaks in an-
recent global molecular phylogeny of the Scarabaeinae cestry for the genera currently assigned to the Ateuchini
(Tarasov & Dimitrov 2016), revision of the tribal classifica- and Canthonini. The same is clearly true of the phyloge-
tion system is incomplete. Therefore, the old morphologi- ny of Tarasov and Dimitrov (2016) (Figure 5: black basal
cally based, 12-tribe system (nine tribes in southern Africa) branches) in which ateuchine and canthonine genera are
is used for arranging Appendix 3 and the 542 species ac- dispersed throughout the topology in Clades 1–13 and
counts. However, notes are provided on putative closeness 15–20. They thus form at least part of the membership
or distance of relationships between genera in each of the of most clades except the most terminal. However, boot-
three extensively polyphyletic tribes (section B, below). strap support is relatively weak for defining relationships
at the level of basal nodes in both phylogenies. This in
By contrast, the phylogeny of Tarasov and Dimitrov (2016) turn weakens support for any current revision of the tribal
is used to arrange Summary Tables 1 and 2 (Appendix 1 classification system.
and 2). For the three polyphyletic tribes, arrangement is
based on the derivation of clades in numbered sequence There is, however, often high support for clades at more
along the ladderised topology (Figure 5: Clades 1–23). terminal nodes (see Tarasov & Dimitrov 2016). Each of
Each clade is further identified according to its placement the 23 clades would represent separate radiations of taxa.
in the old 12-tribe, morphological classification system. For Analyses of distribution in southern Africa identify clear
six primarily monophyletic tribes, arrangement is alphabet- differences in pattern bias between some clades. In gener-
ical, although each is further identified by clade number. al, species endemic to the southwest, in the winter rainfall
SURICATA 6 (2020) 699

Figure 5. The most recent global molecular phylogeny for the Scarabaeinae after Tarasov and Dimitrov (2016), redrawn, recaptioned
and divided into 23 clades used to define groups in the NMDS analyses of Figure 6. Most Canthonini, Ateuchini and Coprini
are incertae sedis except those designated sensu novo by Tarasov and Dimitrov (2016) (includes Ateuchini pars). Note that a few
Catharsius and Heliocopris occur in the Oriental region and a few Neotropical genera occur in the Nearctic region.
700 SURICATA 6 (2020)

and desert regions, are most prominent in clades defined rainfall endemics occur in the Onthophagini, Onitini
for three polyphyletic tribes and the Scarabaeini. The re- and Sisyphini. Some also show SW Arid representatives,
maining five tribes are mostly centred in the savanna of particularly some Gymnopleurini, Onthophagini and
the summer rainfall region in the northeast (particular- Onitini. A more detailed summary of the patterns follows
ly Oniticellini, Gymnopleurini) although a few winter below.

B. CLADISTIC GROUPS
Distances between spatial distribution patterns have been Afrotropical, one Afro-Asian) that are classified in the
summarised for all but a few of the species in 542 species Ateuchini (six genera in total in Africa). Basally derived
accounts using NMDS ordination (Statsoft Inc. 2017) with (1) Coptorhina and Delopleurus centred in Highveld
and minimum spanning trees (Excoffier et al. 2006) to Grassland and Savanna with relatives or distributions
connect data points. The analyses were based on the 23 into the tropics and (2) Frankenbergerius and Sarophorus
clades defined by the most recent cladistic analysis of re- with both grassland/savanna and winter rainfall-centred
lationships within the subfamily Scarabaeinae (Tarasov & species. All with specialist or putative specialist food as-
Dimitrov 2016, Figure 5). The analysed distribution data sociations. Described as part of ‘Basal Scarabaeinae’ by
for each clade comprised a combined matrix for: Tarasov and Dimitrov (2016).
1. Numbers of 1/16th degree squares of latitude and
Clade 2 (Canthonini) (Figure 6B): Comprises 14 ba-
longitude occupied in each of the eight biomes of
sally derived Afrotropical genera with specialist habits,
South Africa.
all assigned to the Canthonini (in 2017). The very small-
2. Numbers of 1/16th degree squares occupied in each bodied, species-rich genus, Odontoloma, of unknown hab-
of 11 main ecoregions for Namibia, Botswana, its (possibly kleptocoprid) is equally centred in the summer
Zimbabwe and Mozambique within the standard rainfall and winter plus bimodal rainfall regions with sev-
map panel south of 17°S and west of 33°E (see Fig- eral tropical relatives. The remaining species-poor genera
ure 2 in Introduction). comprise (1) six forest litter taxa along the E escarpment
3. Numbers of 1/16th degree squares occupied in each of the Highveld in the summer rainfall region and coast-
of the 86, 2° × 2° squares (each = a total of four 1° line in the bimodal to winter rainfall regions; and (2) six
squares) in the map panel graduated from 17° to genera possibly associated primarily with hyrax middens
35° S and 12° to 34° E (see Figure 2). along the arid western seaboard from winter rainfall Na-
maqualand northwards into Namibia. Described as part
Clades containing large numbers of species were subdi- of ‘Basal Scarabaeinae’ by Tarasov and Dimitrov (2016).
vided into taxonomic units (two separate NMDS ordina-
tions for different groups of genera in Scarabaeini; three Clades 4, 7, 10, 16 (Canthonini) (Figure 6C): Com-
different NMDS ordinations for the Onthophagini rep- prises four distantly related Afrotropical genera currently
resenting different genera or different species groups of assigned to the Canthonini. Possibly relict or derived with
Onthophagus). ball-rolling, forest litter or unknown habits. These com-
prise Bohepilissus and Circellium (small-bodied, flightless,
Genera from South Africa, Namibia and Botswana oc- forest litter genus or large bodied, monotypic and flight-
curred in 16 of the 23 clades defined in the global molec- less, ball-rolling genus, both in bimodal and winter rain-
ular phylogeny of Tarasov and Dimitrov (2016; Figure 5). fall regions), Chalconotus (savanna and tropical woodland
Distances between species and geographical distribution or forest ball-rolling genus), Gyronotus (flightless E coast
centres are summarised by 16 NMDS ordinations (Figure and E escarpment forest or grassland specialist genus of
6). Principal affiliations and geographical patterns are sum- unknown habits).
marised below for each of 13 cladistic groupings or tribes.
Clade 5 (Ateuchini, Coprini) (Figure 6D): Comprises
both the possibly kleptocoprid, Afrotropical, ateuchine
Polyphyletic tribes genus, Pedaria, and the tunnelling Coprini genera, Lito­
(Clades in numerical order) copris and Copris (Americas, Afro-Eurasia), but only with
54% bootstrap support. Mostly centred in the summer
Clade 1 (Ateuchini) (Figure 6A): Comprises four out rainfall region (Savanna, Highveld, NE KwaZulu-Na-
of five genera represented in southern Africa (three tal coast, N Namibia) with some winter and bimodal
SURICATA 6 (2020) 701

rainfall Copris species showing relatives on the Highveld two Afro-Eurasian and Americas). Southern African dis-
and E coast. tributions are centred on the summer rainfall region, pri-
marily in savanna and Highveld Grassland with a few in
Clades 13, 14 (Coprini) (Figure 6E): Comprises two shaded habitats. Strong bias to dung of ruminant or mo-
neighbouring lineages that include three Afro-Eurasian nogastric herbivores in some species.
tunnelling genera that are distant from two other clades
containing genera assigned to the Coprini. The three gen- Onitini (Clade 22) (Figure 6J): In the study area, the tribe
era of Coprini in Clades 13 and 14 comprise large-bod- comprises 10 tunnelling genera, eight of them species-
ied Heliocopris, medium-bodied Catharsius and smaller- poor Afrotropical endemics, mostly (seven) associated
bodied Metacatharsius with the latter more characteristic with monogastric herbivore dung. Two species-rich Af-
of the Kalahari. However, all three genera are centred in ro-Eurasian genera (Cheironitis, Onitis) include species
the summer rainfall region, Heliocopris primarily in savan- groups associated with ruminant or monogastric herbi-
na whereas Catharsius appears more diversified eco-geo- vore dung. In southern Africa, species are centred pri-
graphically. marily in the summer rainfall region with a bias to the
SW Arid region in Cheironitis and to savanna or Highveld
Clade 18 (Canthonini) (Figure 6F): Comprises the Grassland in Onitis with very few species centred in the
ball-rolling genus, Epirinus, which is assigned to the Can- winter and bimodal rainfall regions.
thonini (‘classic’ affiliation), to the Sisyphini (recent trans-
ferral by Tarasov and Dimitrov (2016)), or to the tribe Onthophagini (Clade 23) (Figures 6K, 6L, 6M): In the
Epirinini in a more recent assessment (Daniel et al. 2020). study area, the tribe comprises 15 mainly tunnelling gen-
Species are more-or-less equally shared between the winter era (two Afro-Eurasian, five Afro-Asian, seven Afrotropical)
and bimodal rainfall regions and higher rainfall regions in including the extremely species-rich, globally distributed
the summer rainfall region primarily along the E escarp- genus, Onthophagus. Distribution patterns in southern Af-
ment and E coastal forests. rica are shown by three separate NMDS ordinations owing
to the number of species. The bulk of the distributions are
Clades 19, 20, 21 (Canthonini, Coprini) (Fig 6G): centred in the summer rainfall region, particularly savanna,
Three neighbouring clades comprising six genera (five with some in Highveld Grassland, a few in the SW Arid
Afrotropical, one Afro-Asian) currently assigned to the region, and only six species centred in the winter and bi-
Canthonini or Coprini that include some genera previ- modal rainfall regions (out of a total of 123).
ously identified as doubtful canthonines (Scholtz & How-
den 1987). All genera show restricted distribution patterns Scarabaeini (Clade 4) (Figures 6N, 6O): In the study
and/or specialist habits: Aphengoecus (SW Cape-centred, area, the tribe comprises seven primarily ball-rolling gen-
possibly termitophilous (Péringuey 1901)), Dwesasilvase­ era with Afrotropical (4), Afro-Eurasian (2) or Afro-Asian
dis (monotypic in E coast forest litter), Hammondantus (1) distributions. Distribution patterns in southern Africa
(Namib Desert dune specialist), Macroderes (flightless, ac- are shown by two separate NMDS ordinations owing to
tive under cool conditions, primarily in the winter rainfall the number of species. There is a distinct SW distribution-
region), Pycnopanelus (SW Arid-centred with conspecifics al bias in most genera although most have relatives in the
in Sudan and W Oriental region), the tunnelling Xinid­ tropics. However, all genera show a few or many winter
ium (the only summer rainfall-centred clade member in rainfall representatives. Distribution is restricted to the W
high altitude grasslands and forest on E escarpment). coast in flightless Pachysoma, from the winter rainfall region
northwards into Namibia. SW distributional bias is strong
in small-bodied Scarabaeolus (winter rainfall region, SW
Primarily monophyletic tribes Arid, Kalahari, a few on the E coast). SW and NE bias is
(alphabetical order) roughly even in mostly larger-bodied Scarabaeus, Sceliages
and Escarabaeus whereas NE bias dominates in Pachylomera
Gymnopleurini (Clade 15) (Figure 6H): In the study and in Kheper, which shows various tropical relatives.
area, the tribe comprises three Afro-Eurasian or Afro-Asian
ball-rolling genera with the tunnelling genus, Heliocopris, Sisyphini (Clade 18) (Figure 6P): In the study area, the
in a neighbouring clade, but with weak support for the re- tribe comprises two ball-rolling genera (one Afro-Asian,
lationship. Species entirely centred in the summer rainfall one Afro-Eurasian and Americas) with southern African
region, primarily in savanna and SW Arid regions. distributions mostly centred in the mid-summer rain-
fall region, primarily savanna and Highveld Grassland.
Oniticellini (Clade 23) Figure 6I): In the study area, Recent changes reduce the two genera to a single genus
the tribe comprises 14 tunnelling and endocoprid genera comprising three subgenera all, of which, occur within the
(breed within droppings) (six Afrotropical, six Afro-Asian, study area (Daniel et al. 2020).
702 SURICATA 6 (2020)

Figure 6. NMDS and Minimum Spanning Tree representation of statistical distances between species distribution patterns within
the standard distribution map panel: A, Clade 1 (Ateuchini); B, Clade 2 (Canthonini).
SURICATA 6 (2020) 703

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution
patterns within the standard distribution map panel: C, Clades 4, 7, 10 and 16 (Canthonini); D, Clade 5 (Ateuchini, Coprini).
704 SURICATA 6 (2020)

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution pat-
terns within the standard distribution map panel: E, Clades 13, 14 (Coprini); F, Clade 18 (Canthonini).
SURICATA 6 (2020) 705

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution pat-
terns within the standard distribution map panel: G, Clades 19, 20, 21 (Canthonini, Coprini); H, Gymnopleurini (Clade 14).
706 SURICATA 6 (2020)

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution pat-
terns within the standard distribution map panel: I, Oniticellini (Clade 23); J, Onitini (Clade 22).
SURICATA 6 (2020) 707

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution patterns
within the standard distribution map panel: K, Onthophagini 1 (Clade 23); L, Onthophagini 2 (Clade 23, species groups 2–16).
708 SURICATA 6 (2020)

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution patterns
within the standard distribution map panel: M, Onthophagini 3 (Clade 23, species groups 17–31); N, Scarabaeini 1 (Clade 4).
SURICATA 6 (2020) 709

Figure 6 (continued). NMDS and Minimum Spanning Tree representation of statistical distances between species distribution pat-
terns within the standard distribution map panel: O, Scarabaeini 2 (Clade 4); P, Sisyphini (Clade 18).
710 SURICATA 6 (2020)

C. MAJOR SPATIAL PATTERNS

Geographical ranges varied from very small to very large. Endemic patterns: South Africa
A number of distinct regional distribution patterns were
recognised for species showing intermediate range sizes 1. SW Cape: Centred on the winter rainfall region
and these are described below. Further localisation and (Figure 7D); localisation within pattern due to soil
smaller range size within these patterns were driven by (sand/finer-grained), vegetation (forest, shrubland,
climate, soil and/or vegetation type specialisation. At the scrub, disturbed patches), climate (coastal lowlands
opposite extreme, larger range size was shown by species or mountain). Distinct sub-pattern on W Coast
with more generalist, widespread associations. These oc- caused by increasing aridity from S to N (Nama­
curred across different combinations of the intermediate- qualand) (Figure 7B) and by changing edaphic
sized regional patterns. character from W to E (sand to finer-grained soils
and cooler climate above W escarpment) (SW
CAPE, W COAST, NAMAQUALAND).
Examples of some more common
widespread patterns: Examples of the patterns are:
• SW Cape: Scarabaeus suri, Scarabaeus intricatus,
1, 2. (SW Cape, S Cape): Epirinus comosus, Odontolo­ Odontoloma pygidiale, Sarophorus diabolus.
ma dentinum, Sarophorus tuberculatus, Ontho­ • W Coast: Scarabaeolus reichei, Scarabaeus rugo­
phagus giraffa. sus, Pachysoma hippocrates, Epirinus scrobiculatus.
2, 3. (Highveld, S Cape): Sisyphus costatus, Onitis • Namaqualand: Byrrhidium ovale, Pachysoma
caffer, Onthophagus binodis, Oniticellus pictus. striatum, Scarabaeus hottentorum, Macroderes na­
2, 3. (Highveld, E Escarpment, S Cape): Epirinus makwanus, Macroderes mutilans.
relictus, Neosisyphus barbarossa, Sisyphus murica­ 2. S Cape: Centred on the bimodal spring/autumn
tus. rainfall region (Figure 7D); localisation within
3, 4. (Highveld, E Coast): Onitis pecuarius, Cyptochi­ pattern due to soil (sand/finer-grained), vegetation
rus ambiguus, Euoniticellus triangulatus.
(forest, shrubland, scrub), climate (winter to bi-
3, 5. (Highveld, Upper Karoo): Scarabaeus basuto,
modal rain from W to E) (S CAPE).
Catharsius vitulus, Phalops bubalus, Onthophagus
cyaneoniger. Examples of the pattern are:
4, 7. (E coast, E savanna): Heliocopris hamadryas, • S Cape: Circellium bacchus, Scarabaeus savignyi,
Onthophagus cribripennis. Copris jacchus, Sarophorus striatus, Sisyphus peris­
4, 7. (NE KZN coast, E savanna): Pedaria segregis, sinottoi.
Sisyphus sordidus, Phalops flavocinctus. • S Cape Forest: Bohepilissus subtilis, Epirinus sil­
vestris.
3. Highveld: Centred on the Grassland Biome (Fig-
Examples of some rarer, but
ure 3); localisation within pattern due to increasing
distinctive widespread patterns:
rainfall and higher altitude from W to E. Leads to
distinct drier NW, W or wetter SE or E escarpment
3, 4, 7. (Highveld, E coast, savanna): Onitis alexis,
Onthophagus aeruginosus, Liatongus militaris. sub-patterns; some species are endemic to forest
1, 2, 5. (SW and S Cape, Upper Karoo): Epirinus aene­ patches at different points along escarpment in the
us, Onthophagus cameloides. E and SE (HIGHVELD, HIGHVELD FOREST).
1, 6. (SW Cape W Coast / Namaqualand, Kalaha- Examples of the patterns are:
ri): Scarabaeolus flavicornis, Metacatharsius lati­ • Highveld: Coptorhina excavata, Copris antares,
frons. Copris corniger, Onthophagus obtutus.
3, 4. (NE KZN coast / Highveld E escarpment): Si­ • Highveld (NW): Epirinus gratus.
syphus fasciculatus, Afrodrepanus impressicollis. • Highveld (SE): Epirinus obtusus.
3, 7. (Highveld / N Namibia): Scarabaeus ambiguus, • Highveld (E): Epirinus asper, Epirinus validus,
Catharsius ulyssses, Neosisyphus macrorubrus, Pro­ Copris caelatus, Catharsius marcellus.
agoderus lanista. • Highveld Forest (edge of E or SE Escarpment):
5, 7. (Savanna (E & W), Upper Karoo): Scarabaeolus Epirinus punctatus, Frankenbergerius armatus tu­
bohemani, Heliocopris faunus, Onthophagus bovi­ berculatus, Xinidium dewitzi, Peckolus alpinus,
nus. Endroedyolus paradoxus.
SURICATA 6 (2020) 711

Shared pattern: South Africa, Mozambique • NE KZN Coast Forest: Gyronotus carinatus,
Scarabaeus bornemisszai, Sisyphus neobornemis­sza­
4. E Coast: Centred on the moist, E coastal belt (Figure nus, Garreta caffer.
3); localisation within pattern due to soil (sand/finer-
grained), vegetation (forest, woodland grassland),
occurrence of reserves with indigenous monogastric Shared patterns: South Africa,
herbivores; distinct sub-pattern in warmer NE (Figs Namibia, Botswana
6C, 6D) that continues northwards into the tropics
(E COAST, NE KZN COAST, FOREST). 5. SW Arid: Centred on the arid late summer rainfall
Examples of the patterns are: region (Figure 7D), primarily in South Africa and
• E Coast: Gyronotus pumilus, Copris orion caffer, Namibia; localisation within pattern due to soil
Mimonthophagus ambiguus, Onthophagus cretus. (sand/finer-grained), particularly sand on outlier
• NE KZN Coast: Kheper clericus, Anonychonitis Kalahari dunes and in saline pans in South Afri-
freyi, Copris inhalatus sanctaluciae, Tropidonitis ca. Also, a N to S climatic gradient from cooler to
paradoxus. warmer and drier to moister conditions (Figures

a b

c d

Figure 7. A, Altitude; B, Annual rainfall; C, Annual temperature (max. + min. / 2); D, Rainfall seasonality and climatic regions after
Walter and Lieth (1964) across the standard distribution map panel (see Figure 2).
712 SURICATA 6 (2020)

7B, 7C) resulting in distinct sub-patterns at higher 7. Savanna: Centred on moist to dry savanna (Figure
altitude on the Upper Karoo in South Africa and 3) in northern regions of southern Africa (Namib-
sub-patterns in the Namib and Kaokoveld Deserts ia, Botswana, South Africa); many species are bi-
in Namibia with increasing coastal coolness from E ased to the E, some to the W, whereas some occur
to W (SW ARID, KAROO, NAMIB). in both the E and W; many savanna distributions
Examples of the patterns are: continue into the tropics. Localisation within the
• SW Arid: Pycnopanelus krikkeni, Gymnopleurus pattern due to soil (sand/finer-grained), vegeta-
humanus, Gymnopleurus asperrimus, Metacathar­ tion (forest, woodland, shrubland, grassland), oc-
sius marani, Onthophagus semiflavus. currence of reserves with indigenous monogastric
• Upper Karoo: Epirinus striatus, Scarabaeus via­ herbivores and/or climate. Local patterns along the
tor, Euonthophagus vicarius, Onthophagus perin­ E–W trending Waterberg, Soutpansberg and N
gueyi. edge of Highveld in South Africa; also N Namib-
• Namib Desert: Pachysoma denticolle, Pachysoma ian Plateau and N Botswana Basin (SAVANNA, N
NAMIBIA, N BOTSWANA).
rodriguesi, Scarabaeolus rubripennis, Hammon­
dantus psammophilus. Examples of the patterns are:
• N Namib-Kaokoveld: Kheper vethi, Scarabaeus • Savanna (E & W): Copris elphenor, Onitis unci­
cognatus, Scarabaeolus namibensis, Phalops plancus. natus, Onthophagus suffusus, Oniticellus egregius.
• E Savanna: Garreta wahlbergi, Catharsius philus,
6. Kalahari: Centred on hot, deep sands of the Ka- Copris amyntor, Onitis viridulus, Onthophagus
lahari in central southern Africa with localisation lamelliger.
due to increasing aridity to the SW (Figures 7B, • Savanna game reserves: Onitis mendax, Proa­
7C) (KALAHARI). goderus rangifer, Proagoderus rectefurcatus, Eu­
Examples of the patterns are: onthophagus jeanneli.
• SW Kalahari: Scarabaeolus inoportunus, Metaca­ • Soutpansberg: Scarabaeus schulzeae.
tharsius anderseni, Copris cornifrons, Euonthopha­ • Waterberg: Onthophagus cupricollis.
gus flavimargo. • N Botswana: Escarabaeus remii, Frankenbergerius
• Kalahari: Catharsius calaharicus, Kurtops qua­ darrenmanni.
draticeps, Proagoderus sapphirinus, Onthophagus • N Namibia (W Savanna): Copris laioides, Copris
granu­lifer. subsidens, Phalops prasinus, Phalops pauliani.

D. CLIMATE, SAMPLING AND GEOGRAPHICAL BIAS

Patterns of species numbers and distribution show clear the SW desert coastline and N plateau in Namibia and
differences across the standard map panel (Figures 8, 9, along major roads in Botswana. Large areas lack collections
10). These differences are due partly to uneven intensity particularly most of the NE and E of Namibia and most of
of collecting (Figure 10A) and partly to geographical vari- the Kalahari system in Bot­swana. However, a few areas of
ation in altitude, annual rainfall, annual temperature and higher sampling and/or collecting intensity stand out as re-
rainfall seasonality (Figures 7A–7D). gards numbers of species (Figure 10D), i.e. NE Botswana,
Northern Cape and NE South Africa.
Depictions of collecting intensity are based on the reference
collections of the National Collection of Insects, Ditsong Comparisons between the nine tribes suggests three main
Museum, University of Pretoria and selected published patterns of species number distribution across South Af-
or unpublished survey data. Collecting intensity has been rica, Botswana and Namibia (Figure 9). Seven tribes (Fig-
high along the coastline and in the NE of South Africa as ures 8A, 8C–8G, 8I) show highest numbers of species
well in parts of the Northern Cape. However, the absence mostly in the NE savanna of South Africa where there is
of records across much of the Karoo and SW Highveld have high collecting intensity (Figure 10A), high topographi-
certainly resulted in lower observed numbers of species than cal and temperature diversity and relatively high summer
would be expected in some areas. In Namibia and Botswa- rainfall (Figures 7A, 7B, 7C). Of these tribes, the Ateuchi-
na, intensity of collection has been lower than in South Af- ni, Sisyphini, and to a lesser extent, the Oniticellini show
rica (Figure 10A). Collecting has been concentrated along a particularly strong bias to the moister regions in the E
SURICATA 6 (2020) 713

(Figures 7B, 8A, 8E, 8I), which may reflect their ecologi- South Africa, but they are particularly diverse in arid cli-
cal associations. The Onitini, Coprini, and, to a lesser ex- mates along the W seaboard of southern Africa although
tent, the Onthophagini, are also fairly well represented in poorly represented on the SE seaboard (Figure 8H). The
other savanna regions in the Northern Cape, N Botswana Canthonini are centred primarily in cooler regions from
and N Namibia (Figures 8C, 8F, 8G). The Gymnopleu- the SW Cape along the S coast and up the E escarpment
rini are also fairly well represented by specialist species in that borders the Highveld (Figure 8B). Fewer species oc-
the arid SW, but are absent from the SW winter rainfall cur locally on the South African highlands and the arid es-
region (Figures 7B, 7D, 8D), unlike the other eight tribes. carpment regions along the W seaboard. It should be not-
The Scarabaeini, and particularly the Canthonini, show ed that only the Scarabaeini, Coprini and Onthophagini,
quite different patterns to the other seven tribes. In savan- show appreciable numbers of species on the deep sands of
na regions, the Scarabaeini are well represented only in the S Kalahari Basin (Figures 8C, 8G, 8H).

a b c

d e f

g h i

Figure 8. Numbers of scarabaeine dung beetles species per 4° square (2×2 1° squares) within the standard distribution map panel
(see Figure 1): A, Tribe Ateuchini; B, Tribe Canthonini; C, Tribe Coprini; D, Tribe Gymnopleurini; E, Tribe Oniticellini; F, Tribe
Onitini; G, Tribe Onthophagini; H, Tribe Scarabaeini; I, Tribe Sisyphini (large text = 50–100% of square with maximum number
of species (maxima: A=12, B=20, C=36, D=10, E=22, F=20, G=73, H=29, I=16), medium text = 25–50%, small text = 0–25%).
714 SURICATA 6 (2020)

Figure 9. MDS ordination plot showing statistical distances between the patterns of species number distribution across South Africa,
Botswana and Namibia (Figures 8A–8I) in nine dung beetle tribes.

a b c

d e f

Figure 10. Collection localities of scarabaeine dung beetles represented by 1/16th of a degree squares: A, Squares in which dung
beetles were collected; B, Squares in which dung beetles endemic to South Africa, Botswana and/or Namibia were collected; C,
Squares in which threatened endemic dung beetle species were collected; D, 540 species of the Subfamily Scarabaeinae; E, Spe-
cies endemic to South Africa, Botswana and/or Namibia; F, Threatened endemic species (large text = 50–100% of square with
maximum number of species (maxima: D=213, E=59, F=20), medium text = 25–50%, small text = 0–25%).
SURICATA 6 (2020) 715

E. CLIMATE, ENDEMISM AND CONSERVATION

Based on available spot distribution data, just over half of which have, presumably, driven speciation and high en-
540 species (Figures 10A, 10D) were considered to be en- demism with geographical barriers across the hot sands of
demic to South Africa, Botswana and/or Namibia (Figures the Botswana Kalahari and the hot, mostly dry Limpopo
10B, 10E). Putative endemic species were recorded across Valley in the NE.
the entire area of the three countries. Numbers of endem-
ics in the SW of South Africa were frequently similar to This hypothesis is supported by the summary (Table 2)
numbers in the NE. However, for example, the 59 or 43 of geographical and climatic associations for endemic and
species recorded in the 4° squares near Cape Town (Fig- non-endemic species (Appendix 1). Table 2 shows that av-
ure 6SD, 6SE) represented 86.8% and 79.6% endemism erage annual temperatures range from 15.71 to 16.77°C
compared to the 59 and 48 recorded in the 4° squares near in the five most southwesterly patterns of distribution
Eswatini, which represented only 29.6% or 22.5% ende- shown by endemic species (E1–E5). This reflects their
mism. Thus, endemism clearly increases greatly to the SW spatial restriction by cool southern African temperatures
but is comparatively low in the hot central basin of the (Figure 7B). The other five patterns shown by endemic
Botswana Kalahari and towards the N borders of the three species (E6–E10) have a more northeasterly bias with
countries (Figures 7C, 10D, 10E). warmer average annual temperatures (18.43 to 19.52°C).
Nevertheless, these remain lower than those in putative
This trend in endemism is, presumably, related to the cli- geographical barrier regions (>20°C) (Figure 7B). For
matic and ecological conditions recorded within the map non-endemic species, average annual temperatures for
panel. The uplands (Figure 7A) and cold west coastal sea three patterns (17.15 to 19.71°C) are also lower than 20°C
currents create cooler conditions than in the central Ka- (NE1–NE3). However, these patterns comprise relatively
lahari Basin and the tropical regions to the north (Figure few species that show distributions extending just over the
7C). The cold west coastal sea currents are also responsible Namibian border into S Angola (SW Arid; N Namibia =
for drier conditions and changing rainfall seasonality to arid, savanna and psammophilous species combined), or
the SW (Figures 7B, 7D) where winter rainfall replaces on highlands in Zimbabwe and/or E Africa (Highveld).
the summer rainfall experienced to the NE. These factors In the remaining four lowland patterns shown by non-
create unique regional and local ecological conditions, endemic species (NE4–NE7), average annual temperatures

Table 2. Average data for different patterns of distribution shown by non-endemic and endemic species of dung beetles in South Africa,
Botswana and Namibia (see also summary of distribution patterns for each species in Appendix 3)

Average climatic and topographical data

Endemic patterns centred on: Number of Annual Annual rainfall Altitude (m) Average
species, subsp. temperature (oC) (mm) EOO (km2)
E1. SW Cape 65 16.77 240 317 29 839
E2. S Cape 22 16.14 432 492 15 776
E3. Upper Karoo 10 16.03 318 1 206 254 724
E4. Highveld 88 15.68 708 1 345 110 292
E5. SW Arid – S Namib 20 16.44 114 992 11 858
E6. N Namibia 11 19.52 347 1 202 67 067
E7. E & NE Coast 17 19.45 867 308 23 585
E8. Kalahari 20 19.51 319 1 055 393 283
E9. SW Arid 14 18.43 219 1 065 232 507
E10. E Savanna 22 19.33 640 970 77 635

Non-endemic patterns centred on:

NE1. Highveld 11 17.15 751 1 028 1 232 819
NE2. N Namibia 21 19.71 458 1 022 520 357
NE3. NE Coast 34 21.95 805 178 324 560
NE4. Kalahari 23 20.18 429 994 1 289 828
NE5. SW Arid 8 18.50 223 1 030 648 278
NE6. E Savanna 102 20.47 669 781 1 232 115
NE7. Savanna (E & W) 55 20.40 585 861 2 542 927
716 SURICATA 6 (2020)

Table 3. Threatened or potentially threatened species with IUCN threat categories*

SOUTH AFRICA SPECIES IUCN SOUTH AFRICA SPECIES IUCN

ATEUCHINI CANTHONINI (DELTOCHILINI) (continued)
Frankenbergerius barratti (Waterhouse, 1876) DD/VU Peckolus poenskopius Deschodt & Scholtz, 2008 DD/EN
Frankenbergerius gomesi (Ferreira, 1954a) DD/VU Silvaphilus oubosiensis Roets & Oberlander, 2010 VU
Frankenbergerius nanus (Péringuey, 1888) DD/EN COPRINI
Frankenbergerius opacus Frolov & Scholtz, 2005 DD/CR Copris jacchus (Fabricius 1775) NT
Sarophorus bidentatus Frolov & Scholtz, 2003a DD/VU Copris sexdentatus Thunberg, 1818 EN
Sarophorus carinatus Frolov & Scholtz, 2003a DD/EN Copris sphaeropterus Harold, 1877b EN
Sarophorus diabolus Roets, 2017 DD/CR Copris victorini Boheman, 1857 DD/EN
Sarophorus frolovi Roets, 2017 DD/VU Macroderes amplior Frolov & Scholtz, 2004 DD/VU
Sarophorus latus Frolov & Scholtz, 2003a DD/VU Macroderes arrowi Janssens, 1939a DD/VU
Sarophorus punctatus Frolov & Scholtz, 2003a EN Macroderes cornutus Frolov & Scholtz, 2004 DD/VU
CANTHONINI (DELTOCHILINI) Macroderes endroedyi Frolov & Scholtz, 2004 NT
Aliuscanthoniola similaris Deschodt & Scholtz, 2008 EN Macroderes fornicatus Sharp, 1880 DD/EN
Aphengoecus clypeatus Péringuey, 1901 DD/CR Macroderes foveatus Frolov & Scholtz, 2004 DD/VU
Aphengoecus multiserratus Scholtz & Howden, 1987 VU Macroderes greeni (Kirby, 1818) DD/VU
Bohepilissus nitidus (Balthasar, 1965a) VU Macroderes minutus Frolov & Scholtz, 2004 DD/EN
Byrrhidium ovale Harold, 1869a DD/VU Macroderes namakwanus Frolov & Scholtz, 2004 DD/VU
Circellium bacchus (Fabricius, 1781) VU Macroderes politulus Preudhomme de Borre, 1880 DD/VU
Dwesasilvasedis medinae Deschodt & Scholtz, 2008 VU Macroderes undulatus Preudhomme de Borre, 1880 NT
Endroedyolus paradoxus Scholtz & Howden, 1987 VU Xinidium dewitzi (Harold, 1878) DD/VU
Epirinus aquilus Medina & Scholtz, 2005 VU ONITINI
Epirinus bentoi Ferreira, 1964a VU Anonychonitis freyi Janssens, 1950 NT
Epirinus comosus Péringuey, 1901 DD/NT Gilletellus porculus (Boheman, 1857) DD/CR
Epirinus convexus Scholtz & Howden, 1987 DD/VU Kolbeellus ateuchoides (van Lansberge 1875b) DD/EN
Epirinus davisi Scholtz & Howden, 1987 DD/VU Tropidonitis paradoxus (Boheman, 1857) VU
Epirinus granulatus Scholtz & Howden, 1987 VU SCARABAEINI
Epirinus hluhluwensis Medina & Scholtz, 2005 DD/VU Kheper clericus (Boheman, 1857) DD/VU
Epirinus minimus Medina & Scholtz, 2005 VU Kheper zurstrasseni Davis, 1986 DD/EN
Epirinus montanus Scholtz & Howden, 1987 DD/EN Pachysoma aesculapius (Olivier, 1789) VU
Epirinus ngomae Medina & Scholtz, 2005 DD/VU Pachysoma endroedyi (Harrison, Scholtz & Chown,
EN
Epirinus pseudorugosus Medina & Scholtz, 2005 DD/VU 2003)
Epirinus punctatus Scholtz & Howden, 1987 DD/VU Pachysoma glentoni (Harrison, Scholtz & Chown,
VU
Epirinus sebastiani Medina & Scholtz, 2005 DD/VU 2003)
Gyronotus carinatus Felsche, 1911 VU Scarabaeus bornemisszai zur Strassen, 1980 NT
Gyronotus glabrosus Scholtz & Howden, 1987 VU Scarabaeus heqvisti zur Strassen, 1962 DD/NT
Gyronotus perissinottoi Moretto, 2013 DD/VU Scarabaeus karae Davis & Deschodt 2017 DD/EN
Nebulasilvius insularis Deschodt & Scholtz, 2008 VU Scarabaeus piliventris zur Strassen, 1962 DD/VU
Nebulasilvius johani Deschodt & Scholtz, 2008 VU Scarabaeus schulzeae Ferreira, 1969 DD/VU
Parvuhowdenius harrisoni Deschodt & Scholtz, 2008 DD/VU SISYPHINI
Peckolus alpinus Howden & Scholtz, 1988 DD/EN Sisyphus neobornemisszanus Daniel & Davis, 2016 NT
Peckolus parvus Scholtz & Howden, 1987 DD/EN Sisyphus perissinottoi Montreuil, 2015c DD/VU

BOTSWANA SPECIES NAMIBIA SPECIES

ONTHOPHAGINI CANTHONINI
Mimonthophagus limbibasis (d’Orbigny, 1905) DD/NT Dicranocara deschodti Frolov & Scholtz, 2003b VU
ONITINI
Platyonitis bicuariensis Ferreira, 1976b VU
SCARABAEINI
Pachysoma schinzi Fairmaire, 1888 DD/VU
*All endemic to South Africa, Botswana and/or Namibia (including Eswatini and Lesotho) except Mimonthophagus limbibasis (also
Zimbabwe), Platyonitis bicuariensis (also Angola), Tropidonitis paradoxus, Scarabaeus bornemisszai and Sisyphus neobornemisszanus
(also S Mozambique).
SURICATA 6 (2020) 717

exceed 20°C (20.18 to 22.00°C). It is also noteworthy are mostly those centred on or localised within those spatial
that average EOO (Extent of Occurrence) is greater in patterns (Table 4). Distributions of these species largely co-
non-endemics than in endemics (Table 2). incide with the Fynbos and Succulent Karoo Biomes (pat-
terns 1–2), Indian Ocean Coastal Belt Biome (pattern 4) or
Of 275 (out of 281) recorded endemic species (Figures small forest patches along the E escarpment encompassed by
10B, 10E, Table 1), a total of 72 are currently listed as fac- the Grassland Biome (pattern 3) (Figures I3, 9C, 9F). The
ing threats or potential threats (Figures 10C, 10F, Table few threatened species in patterns 6–7 are probably related
3). More than half of these species have been described to loss of preferred dung types due to range contraction in
since 1985 (Table 3). Clearly, species showing ecological particular indigenous herbivores. Habitat transformation
specialisation, restricted range and/or endemism to high- in the Grassland Biome remains worrisome as regards the
ly transformed regions face the greatest threats. Therefore, various South African Highveld endemics. Transformation
threatened species are primarily recorded from South Africa, in the NE savanna regions has an impact, but most savanna
which is the most transformed of the three focal countries. species are also distributed beyond the borders of South Af-
Spatial patterns 1–4 (from pg. 710) contain the most trans- rica. Transformation in the SW Arid and Kalahari Savanna
formed habitats in South Africa together with some in pat- regions remains limited. However, the effects of range con-
tern 7 (see Mucina & Rutherford 2006: page 731, Figure traction of indigenous mammals overlie all these effects of
16.5). Thus, species that are allotted IUCN threat categories vegetation modification and are mostly poorly known.

Table 4. Number of threatened or potentially threatened endemic species per spatial pattern in South Africa, Botswana and Namibia

IUCN category
Spatial pattern
CR EN VU NT DD/CR DD/EN DD/VU DD/NT
1. SW Cape 1 8 2 3 3 10
2. S Cape 3 3 1 2 1
3. Highveld 4 6 8 1
4. E coast 1 2 1 5
5. Kalahari
6. SW Arid 1 1 1
7. Savanna 1 2
For an alternative, less qualitative assessment of biogeographical patterns and conservation of dung beetles in southern Africa, see Davis
& Scholtz (2020).
APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th
degree squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; puta-
tive ecological bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Ateuchini (Clade 1)
718

Coptorhina auspicata 0 8 4 85 4 0 0 0 0 26 LC Ss GW F
Coptorhina excavata 0 13 75 13 0 0 0 0 0 8 DD Ss G F
Coptorhina klugii 0 2 33 45 14 4 2 0 0 51 LC ?Gen GW F
Coptorhina nitidipennis 0 0 9 87 4 0 0 0 0 23 LC ? ? ?F
Delopleurus pullus 0 10 0 80 10 0 0 0 0 10 LC ? ? ?F
Frankenbergerius armatus 0 0 46 31 12 12 0 0 0 26 LC ? F>G ?
Frankenbergerius barratti 0 0 100 0 0 0 0 0 0 7 DD (R) ?Fg ?FG ?
Frankenbergerius forcipatus 0 0 33 17 0 50 0 0 0 6 DD (R) ? ?F ?
Frankenbergerius gomesi 0 0 25 67 8 0 0 0 0 12 DD (U) ? FT ?F
Frankenbergerius nanus 0 0 0 0 0 0 100 0 0 4 DD ?Ss ?S ?
(R, RR)
Frankenbergerius nitidus 0 0 0 0 0 0 0 100 0 1 DD ?Sa ?S ?
(R, RR)
Frankenbergerius opacus 0 0 0 0 0 0 100 0 0 1 DD ?Fg ?S ?
(R, RR)
Sarophorus bidentatus 0 0 0 0 0 0 0 100 0 1 DD ? ? ?
(R, RR)
Sarophorus carinatus 0 0 100 0 0 0 0 0 0 2 DD ? ?T ?
(R, RR)
Sarophorus costatus 0 3 20 73 5 0 0 0 0 102 LC Gen T>W>G O>M>R
Sarophorus diabolus 0 0 0 0 0 0 100 0 0 1 DD ?Fg ?T ?O
(R, RR)
Sarophorus frolovi 0 0 100 0 0 0 0 0 0 1 DD ? ?F ?O
(R, RR)
Sarophorus latus 0 0 25 75 0 0 0 0 0 4 DD (R) ? ? ?OC
Sarophorus punctatus 0 0 0 0 0 0 100 0 0 1 DD ? ?F ?
(R, RR)
Sarophorus striatus 0 0 0 13 0 63 25 0 0 8 LC (RR) ? ? ?
Sarophorus tuberculatus 0 0 0 0 0 29 71 0 0 14 LC Ss TS ?R
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Canthonini (Clade 2)
Odontoloma apiculum 0 0 0 0 0 100 0 0 0 1 DD ? ?T ?
(R, RR)
SURICATA 6 (2020)

Odontoloma dentinum 0 0 0 0 0 13 60 27 0 15 LC Ss G>S ?

Odontoloma disalatum 0 0 0 0 0 0 100 0 0 5 DD ? ? ?
Odontoloma doubei 0 0 83 0 0 0 17 0 0 6 DD ? ?Gen ?Gen
Odontoloma endroedyi 11 0 0 0 0 11 56 22 0 9 LC ? ? ?O
Odontoloma louwi 0 13 50 38 0 0 0 0 0 8 LC FG>Ss ?Gen ?
Odontoloma obscurum 0 0 50 0 0 25 25 0 0 4 LC ? ? ?
Odontoloma pauxillum 0 0 0 80 20 0 0 0 0 5 LC ?Sa ? ?
Odontoloma peckorum 0 0 75 25 0 0 0 0 0 12 LC ?FG ?G ?
Odontoloma planatum 0 0 83 17 0 0 0 0 0 2 DD ?Sa ? ?
Odontoloma pusillum 0 0 0 0 0 0 100 0 0 4 LC Sa>Ss SG ?
Odontoloma pygidiale 3 0 6 0 0 0 68 24 0 34 LC ? ? ?
Odontoloma quadridens 0 0 0 100 0 0 0 0 0 2 DD ?Sa ? ?
Odontoloma sculpturatum 0 0 43 29 0 14 14 0 0 7 DD ? ? ?
Odontoloma spinicaudum 0 0 0 0 0 0 0 100 0 1 DD ?sa ?S ?
(R, RR)
Aliuscanthoniola similaris 0 0 0 0 100 0 0 0 0 1 EN (R, RR) ?FG F ?
Endroedyolus paradoxus 0 0 100 0 0 0 0 0 0 2 EN (R, RR) ? F ?
Nebulasilvius insularis 0 0 0 100 0 0 0 0 0 1 VU (U, ? F ?
RR)
Nebulasilvius johani 0 0 100 0 0 0 0 0 0 1 VU (R, RR) ? F ?
Outenikwanus tomentosus 0 0 0 0 0 0 100 0 0 1 DD ? F ?
(R, RR)
Parvuhowdenius harrisoni 0 0 100 0 0 0 0 0 0 1 DD ? F ?
(R, RR)

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
719

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Canthonini (Clade 2) (continued)
720

Peckolus alpinus 0 0 100 0 0 0 0 0 0 1 DD ? F ?

(R, RR)
Peckolus parvus 0 0 100 0 0 0 0 0 0 1 DD ? F ?
(R, RR)
Peckolus poenskopius 0 0 100 0 0 0 0 0 0 2 DD ? F ?
(R, RR)
Silvaphilus oubosiensis 0 0 0 0 0 0 100 0 0 1 VU (R, RR) ? F ?Gen
Byrrhidium convexum 0 0 0 0 0 0 0 33 67 3 DD ?Ss ? ?Hyrax
(R, RR)
Byrrhidium ovale 0 0 0 0 0 0 0 100 0 5 DD ?FG S Dpellets
(R, RR)
Dicranocara tatasensis 0 0 0 0 0 0 0 100 0 1 LC (R, RR) ?Ss ?S ?Hyrax
Canthonini (Clade 4)
Circellium bacchus 0 0 0 0 0 39 61 0 0 18 VU ?Ss SW RM
Ateuchini (Clade 5)
Pedaria barrei 0 0 0 100 0 0 0 0 0 6 DD ?Ss ?W ?
Pedaria cylindrica 0 0 0 93 7 0 0 0 0 15 LC ?Ss ?GW ?
Pedaria picea 0 17 13 53 15 2 0 0 0 47 LC Sa GW>T O>R>M
Pedaria segregis 0 0 0 85 15 0 0 0 0 13 LC Sa G?W ?
Coprini (Clade 5)
Copris amyntor 0 2 3 94 1 0 0 0 0 120 LC FG W>TG M>OR
Copris anceus 0 0 0 0 0 0 86 14 0 21 LC Sa S ?
Copris antares 4 2 65 8 2 14 6 0 0 51 LC ?Ss ?G ?
Copris bootes 0 0 0 100 0 0 0 0 0 4 LC (LLR) ?Ss W M
Copris caelatus 0 0 85 15 0 0 0 0 0 34 LC FG G ?
Copris cambeforti 0 0 0 100 0 0 0 0 0 2 DD ? ? ?
(R, RR)
Copris capensis 0 0 0 0 0 0 58 42 0 31 LC Ss GS ?
Copris cassius 0 82 4 13 0 0 0 0 0 45 LC Sa G>SW M>RO
Copris cornifrons 4 96 0 0 0 0 0 0 0 25 LC (U) Sa GS ?
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Copris corniger 1 0 85 8 1 4 1 0 0 78 LC ?FG ?G ?
Copris crassus 0 0 88 0 0 0 13 0 0 8 LC ?FG ? ?
(U, LLR)
SURICATA 6 (2020)

Copris denticulatus 0 5 8 86 1 1 0 0 0 104 LC ?FG ?Gen ?

Copris elphenor 1 22 11 64 2 0 0 0 0 184 LC Ss GW R>M>O
Copris evanidus 0 14 5 81 0 0 0 0 0 118 LC ?Ss>FG GW M>R
Copris fidius 0 0 19 35 15 4 27 0 0 78 LC ? F RO
Copris inhalatus 0 13 6 31 50 0 0 0 0 16 LC (U) Sa G ?
Copris jacchoides 2 0 88 10 0 0 0 0 0 42 LC FG G ?R
Copris jacchus 0 0 0 0 0 40 60 0 0 5 NT ?FG ? ?
(U, LLR)
Copris macer 0 0 57 43 0 0 0 0 0 21 LC ?FG ?G ?R
Copris mesacanthus 0 0 20 80 0 0 0 0 0 92 LC FG W>G>T ?RM
Copris obesus 0 3 54 42 1 0 0 0 0 78 LC ?Ss GW ?
Copris orion caffer 0 0 0 17 44 33 6 0 0 18 LC ? ? ?
Copris puncticollis 0 0 0 44 56 0 0 0 0 16 LC Sa G ?
Copris ritsemae 0 0 44 56 0 0 0 0 0 9 LC ? G ?
Copris sexdentatus 0 0 0 0 0 33 67 0 0 3 EN (R, RR) ?FG ?S ?
Copris sphaeropterus 0 0 0 0 0 0 100 0 0 1 EN (R, RR) ? ? ?
Copris urus 0 0 0 36 64 0 0 0 0 14 LC (RR) Sa GW ?
Copris victorini 0 0 0 0 0 0 100 0 0 4 DD ?Ss ?S ?
(U, RR)
Copris vilhenai 0 0 0 100 0 0 0 0 0 23 LC Ss TW>G M>R
Copris vrydaghi 0 0 75 25 0 0 0 0 0 4 DD ? ? ?
Litocopris muticus 0 0 0 100 0 0 0 0 0 2 LC ?FG ?FT ?M
Litocopris simplex 0 0 41 9 9 21 21 0 0 34 LC ?FG ?G ?
Canthonini (Clade 7)
Gyronotus carinatus 0 0 0 20 80 0 0 0 0 5 VU (RR) ? F ?

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
721

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Canthonini (Clade 7) (continued)
Gyronotus glabrosus 0 0 80 20 0 0 0 0 0 5 VU (RR) ? G ?
722

Gyronotus perissinottoi 0 0 0 0 100 0 0 0 0 1 DD (RR) ? G ?

Gyrontus pumilus 0 0 0 15 69 15 0 0 0 13 LC ?FG F ?
Gyronotus schueli 0 0 50 50 0 0 0 0 0 2 DD ? ?G ?
(R, RR)
Canthonini (Clade 10)
Bohepilissus nitidus 0 0 0 0 0 0 100 0 0 1 VU (RR) ? F ?
Bohepilissus subtilis 0 0 0 7 0 0 93 0 0 14 LC ? F ?
Coprini (Clade 13)
Catharsius aegeus 0 3 6 91 0 0 0 0 0 35 LC ? ? C
Catharsius calaharicus 17 64 0 19 0 0 0 0 0 53 LC Sa GSW OM>R
Catharsius harpagus 0 0 0 40 60 0 0 0 0 5 DD (RR) Sa G ?O
Catharsius heros 0 50 0 50 0 0 0 0 0 12 LC (LLR) Ss W M>O
Catharsius laticeps 0 0 0 100 0 0 0 0 0 3 DD ? ? ?
Catharsius longiceps 0 0 75 25 0 0 0 0 0 8 DD ? G ?
Catharsius marcellus 0 0 94 6 0 0 0 0 0 32 LC ?FG G ?
Catharsius melancholicus 0 93 0 7 0 0 0 0 0 14 LC Sa ? C>O
Catharsius pandion 0 0 0 19 81 0 0 0 0 21 LC Sa FT ?
Catharsius philus 0 0 7 93 0 0 0 0 0 60 LC FG ? RO>M
Catharsius platycerus 0 0 0 100 0 0 0 0 0 1 DD ? ?W M
(U, LLR)
Catharsius sesostris 0 0 30 64 6 0 0 0 0 50 LC ? T>W>G OC>MR
Catharsius tricornutus 0 7 29 43 5 2 14 0 0 114 LC Ss W>GT O>MR
Catharsius ulysses 0 40 15 45 0 0 0 0 0 40 LC Ss ?WS>G ?
Catharsius vitulus 50 0 50 0 0 0 0 0 0 26 LC Ss GS ?
Metacatharsius anderseni 0 100 0 0 0 0 0 0 0 17 LC Sa ?SG M>OR
M. dentinum & zuluanus 36 32 2 22 8 0 0 0 0 50 LC ?Sa ?S C>O>MR
Metacatharsius exiguiformis 25 75 0 0 0 0 0 0 0 53 LC Sa SG Dgen
Metacatharsius exiguus 0 35 0 41 24 0 0 0 0 17 LC Sa SG ?RM
Metacatharsius ferreirae 0 0 0 100 0 0 0 0 0 1 DD ?Sa ?S ?C
Metacatharsius freyi 0 0 0 100 0 0 0 0 0 3 DD ?Sa ? ?
Metacatharsius latifrons 2 54 4 0 0 0 17 23 0 48 LC Sa SG ?OM>R
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Metacatharsius marani 72 26 1 0 0 0 0 1 0 90 LC Ss ?S>G ?
M. opacus & pseudoopacus 2 4 4 78 12 0 0 0 0 51 LC Ss WS CDGen
M. pumilionis & pumilioni- 0 62 3 35 0 0 0 0 0 37 LC ? ? ?
SURICATA 6 (2020)

formis
Metacatharsius troglodytes 0 29 4 68 0 0 0 0 0 77 LC Sa G M>O>R
Coprini (Clade 14)
Heliocopris andersoni 0 11 16 64 7 2 0 0 0 44 LC ? ? ?M>O
Heliocopris atropos 6 28 0 67 0 0 0 0 0 18 LC Ss WG ?M>RO
Heliocopris faunus 55 0 7 38 0 0 0 0 0 29 LC ?Gen ?WS ?
Heliocopris hamadryas 1 9 26 48 7 4 5 0 0 92 LC ?Ss ? ?
Heliocopris japetus 0 24 0 75 2 0 0 0 0 55 LC Sa ?W>G OM>R
Heliocopris neptunus 0 0 4 96 0 0 0 0 0 54 LC ?FG GSW ?
Heliocopris pirmal 0 6 32 62 0 0 0 0 0 50 LC ?FG ? ?
Canthonini (Clade 16)
Chalconotus convexus 0 1 6 82 10 1 0 0 0 136 LC Gen T>W>G OC>M>R
Canthonini (Clade 18)
Epirinus aeneus 41 6 0 0 0 1 39 10 3 80 LC Ss G>S ?
Epirinus aquilus 0 0 0 0 0 100 0 0 0 2 VU (R, RR) Sa F ?
Epirinus asper 0 0 73 27 0 0 0 0 0 15 LC ?FG G ?
Epirinus bentoi 0 0 0 0 0 0 71 29 0 7 VU Sa S>G ?
(U, RR)
Epirinus comosus 0 0 0 0 0 0 100 0 0 10 DD Sa S ?
Epirinus convexus 0 0 25 25 50 0 0 0 0 4 DD ? F ?
(R, RR)
Epirinus davisi 0 0 0 100 0 0 0 0 0 2 DD ? F ?
(R, RR)
Epirinus drakomontanus 0 0 100 0 0 0 0 0 0 2 DD ?FG G ?
(R, RR)

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
723

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Canthonini (Clade 18) (continued)
724

Epirinus flagellatus 4 0 7 0 0 4 55 29 0 69 LC Ss GS ?
Epirinus granulatus 0 0 0 0 0 0 50 50 0 8 VU (RR) Sa S ?
Epirinus gratus 0 53 47 0 0 0 0 0 0 17 LC ? G ?
Epirinus hilaris 0 0 0 0 0 15 85 0 0 13 LC ? SF ?
Epirinus hluhluwensis 0 0 0 100 0 0 0 0 0 1 DD ? F ?
(R, RR)
Epirinus minimus 0 0 0 0 0 50 50 0 0 2 VU (R, RR) ? F ?
Epirinus montanus 0 0 0 0 0 0 100 0 0 3 DD ? S ?
(R, RR)
Epirinus mucrodentatus 0 0 50 50 0 0 0 0 0 2 DD ? ? ?
(R, RR)
Epirinus ngomae 0 0 0 100 0 0 0 0 0 1 DD (RR) ? F ?
Epirinus obtusus 18 0 45 5 0 27 5 0 0 22 LC ?FG GS ?
Epirinus pseudorugosus 0 0 0 0 0 0 100 0 0 2 DD Sa G>S ?
(R, RR)
Epirinus punctatus 0 0 80 20 0 0 0 0 0 5 DD (R) ? F ?
Epirinus pygidialis 0 0 100 0 0 0 0 0 0 3 DD (R) ? ?FS ?
Epirinus relictus 0 0 44 11 0 0 44 0 0 9 LC (U) ? ? ?
Epirinus rugosus 0 0 0 0 0 0 100 0 0 1 DD ? ? ?
(R, RR)
Epirinus scrobiculatus 0 0 0 0 0 0 41 59 0 17 LC Sa S>G ?
Epirinus sebastiani 0 0 100 0 0 0 0 0 0 1 DD ? F ?
(R, RR)
Epirinus silvestris 0 0 0 0 0 0 100 0 0 8 LC (U, RR) ? F ?
Epirinus striatus 77 3 16 0 0 0 3 0 0 31 LC Ss ? ?
Epirinus sulcipennis 0 0 0 0 0 0 100 0 0 1 DD ?Ss ?S ?
(R, RR)
Epirinus validus 0 0 86 10 0 5 0 0 0 21 LC ?Ss G ?
Canthonini (Clade 19)
Aphengoecus clypeatus 0 0 0 0 0 0 100 0 0 1 DD ? ? ?
(R, RR)
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Aphengoecus multserratus 0 0 0 0 0 0 88 13 0 8 VU Sa S ?
(U, RR)
Canthonini (Clade 20)
SURICATA 6 (2020)

Dwesasilvasedis medinae 0 0 0 0 100 0 0 0 0 1 VU (R, RR) ?Ss F ?

Pycnopanelus krikkeni 68 32 0 0 0 0 0 0 0 34 LC Ss ?S>G ?
Coprini (Clade 21)
Macroderes amplior 0 0 0 0 0 0 0 100 0 3 DD ?Ss ?S ?
(R, RR)
Macroderes arrowi 0 0 0 0 0 0 0 100 0 1 DD ?Sa ?S ?
(R, RR)
Macroderes bias 13 0 38 0 0 25 25 0 0 8 DD (U) ?Ss ?S ?
Macroderes cornutus 0 0 0 0 0 0 0 100 0 4 DD ?Sa ?S ?
(R, RR)
Macroderes endroedyi 0 0 0 0 0 0 71 29 0 7 NT (R, RR) ? ? ?
Macroderes fornicatus 0 0 0 0 0 0 100 0 0 2 DD Sa S ?
(R, RR)
Macroderes foveatus 0 0 0 0 0 0 50 50 0 4 DD ? ? ?
(R, RR)
Macroderes greeni 0 0 0 0 0 0 100 0 0 3 DD Sa S ?
(R, RR)
Macroderes minutus 0 0 0 0 0 0 22 78 0 9 DD R, RR) ? ? ?
Macroderes mutilans 0 0 0 0 0 0 0 100 0 4 DD ? ? ?
(R, RR)
Macroderes namakwanus 0 0 0 0 0 0 0 100 0 4 DD Sa ? ?
(R, RR)
Macroderes nitidus 0 0 0 0 0 0 0 100 0 1 DD ? ? ?
(R, RR)
Macroderes politulus 0 0 0 0 0 0 100 0 0 6 DD ? ? ?
(R, RR)
CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
725

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Coprini (Clade 21) (continued)
726

Macroderes undulatus 0 0 0 0 0 0 100 0 0 4 NT (R, RR) ?FG ? ?

Xinidium dentilabris 0 0 75 24 1 0 0 0 0 68 LC FG G ?
Xinidium dewitzi 0 0 75 25 0 0 0 0 0 4 DD ? F ?
(U, RR)
Xinidium howdeni 0 0 100 0 0 0 0 0 0 2 DD ? F ?
(R, RR)
Gymnopleurini (Clade 15)
Allogymnopleurus consocius 0 0 44 56 0 0 0 0 0 9 DD (LLR) FG ? ?R
Allogymnopleurus splendidus 21 19 9 47 3 0 0 0 0 180 LC Ss G>W O>R>M
Garreta australugens 0 0 0 100 0 0 0 0 0 4 DD (U) ?FG ? ?
Garreta caffer 0 0 0 18 82 0 0 0 0 11 LC Sa F ?
Garreta laetus 0 0 50 50 0 0 0 0 0 2 LC ? F ?
Garreta unicolor 0 0 41 50 9 0 0 0 0 74 LC ?FG G>S>W O>R>M
Garreta wahlbergi 0 0 5 94 1 0 0 0 0 102 LC FG W>T>G O>R>M
Gymnopleurus aenescens 20 34 0 46 0 0 0 0 0 59 LC Ss GS>W O>R>M
Gymnopleurus andreaei 76 24 0 0 0 0 0 0 0 50 LC ?Ss GS ?DGen
Gymnopleurus asperrimus 65 35 0 0 0 0 0 0 0 71 LC ?Ss GS ?DGen
Gymnopleurus humanus 71 21 5 0 0 2 1 1 0 111 LC ?Ss GS ?DGen
Gymnopleurus humeralis 0 0 0 100 0 0 0 0 0 67 LC ?Ss SW O>R
Gymnopleurus leei 23 0 63 0 0 2 13 0 0 48 LC ?Gen G ?
Gymnopleurus pumilus 0 0 0 100 0 0 0 0 0 66 LC FG ?W O>R>M
Gymnopleurus reichei 0 0 0 100 0 0 0 0 0 4 LC ? ? ?Gen
(R, LLR)
Gymnopleurus thelwalli 0 0 0 100 0 0 0 0 0 4 DD (R) ? ? ?
Gymnopleurus virens 0 0 33 64 3 0 0 0 0 66 LC FG W>G>T ?
Oniticellini (Clade 23)
Afrodrepanus impressicollis 0 0 10 67 24 0 0 0 0 21 LC ?FG>Ss FW O>R>M
Cyptochirus ambiguus 0 0 56 30 3 2 8 0 0 86 LC FG G ?R
Drepanocerus kirbyi 0 1 36 48 4 8 3 0 0 75 LC Gen WT>G R>M>O
Drepanocerus patrizii 14 14 22 44 0 6 0 0 0 36 LC Gen W>G>T RM
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Eodrepanus bechynei 0 0 0 100 0 0 0 0 0 4 LC ?Gen ? ?RM
Eodrepanus fastiditus 0 2 30 61 7 0 0 0 0 44 LC FG>Sa GW M>R
Eodrepanus parallelus 0 0 0 100 0 0 0 0 0 2 LC ? ? ?RM
SURICATA 6 (2020)

Epidrepanus pulvinarius/ 0 0 40 55 5 0 0 0 0 20 DD ?FG W DGen

caelatus
Euoniticellus africanus 15 6 55 5 1 5 11 1 0 149 LC ?FG G ?RM
Euoniticellus intermedius 19 13 19 33 3 4 9 1 0 281 LC Gen WG>T R>M>O
Euonticellus kawanus 0 0 0 100 0 0 0 0 0 1 DD ? ?W M
(U, LLR)
Euoniticellus triangulatus 1 0 40 25 5 5 24 1 0 128 LC ?Gen G ?RM
Euoniticellus zumpti 0 0 0 93 7 0 0 0 0 14 DD ? ?W M>O
Ixodina abyssinica 0 0 0 100 0 0 0 0 0 4 DD ? ? ?RM
Latodrepanus laticollis 0 3 0 97 0 0 0 0 0 33 LC Sa>FG W>G>T RM>O
Liatongus militaris 1 3 31 50 7 3 6 0 0 184 LC FG>Sa G>W>T M>R
Oniticellus egregius 0 0 4 92 4 0 0 0 0 25 LC (U) ? ? M
Oniticellus formosus 2 16 7 68 7 0 0 0 0 57 LC FG>Sa G>W>T M>R
Oniticellus pictus 0 0 42 13 0 19 26 0 0 31 LC ?FG ?G>W ?RM
Oniticellus planatus 0 0 29 47 8 6 10 0 0 113 LC Gen ?G>W>T ?MR
Paraixodina freyi 0 0 0 86 14 0 0 0 0 7 DD (LLR) ?FG ?W ?M
Paraixodina saegeri 0 0 0 100 0 0 0 0 0 6 DD (LLR) ? ?WF M
Tibiodrepanus sulcicollis 0 0 52 17 7 7 17 0 0 29 LC ?FG ?G ?MR
Tiniocellus eurypygus 0 0 6 94 0 0 0 0 0 50 LC Sa>FG W>GT M>O>R
Tiniocellus spinipes 0 0 0 100 0 0 0 0 0 10 LC ?FG ?W ?M
Tragiscus dimidiatus 0 0 0 100 0 0 0 0 0 13 LC ?Gen ?W M
(U, LLR)
Onitini (Clade 22)
Anonychonitis freyi 0 0 0 100 0 0 0 0 0 2 NT (R, RR) ? ? M
Cheironitis audens 86 14 0 0 0 0 0 0 0 49 LC Gen SW MR

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
727

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onitini (Clade 22) (continued)
728

Cheirontis hoplosternus 36 31 20 10 0 0 3 0 0 80 LC Ss SW MR
Cheironitis imitator 0 0 0 100 0 0 0 0 0 11 DD Ss W M
Cheironitis indicus 79 21 0 0 0 0 0 0 0 28 LC Ss SW MR
Cheironitis scabrosus 57 14 5 0 0 0 14 10 0 107 LC Gen G R
Heteronitis castelnaui 0 0 0 97 3 0 0 0 0 36 LC FG W M
Kolbeellus ateuchoides 50 0 50 0 0 0 0 0 0 2 DD ? ? ?M
(R, RR)
Megalonitis bohemani 0 0 0 100 0 0 0 0 0 1 DD ?Gen ?W M
(R, LLR)
Neonitis nigritiae 0 0 0 100 0 0 0 0 0 1 DD (R) ? ? M
Onitis aeruginosus 0 0 0 100 0 0 0 0 0 2 LC ?Ss W MO
Onitis alexis 5 11 27 49 2 3 3 0 0 248 LC Gen G>W>T RM
Onitis aygulus 28 18 19 0 0 5 25 5 0 111 LC Gen GS R
Onitis caffer 5 7 38 25 2 5 17 2 0 210 LC Gen G R
Onitis confusus 17 17 4 0 0 9 43 9 0 23 LC Sa SG RM
Onitis cribratus 0 0 100 0 0 0 0 0 0 3 DD FG G ?R
(R, RR)
Onitis curvipes 15 8 38 0 0 0 31 8 0 13 DD (R) ?FG G ?M
Onitis deceptor 0 40 0 50 10 0 0 0 0 30 LC Sa W>T>G ?MR
Onitis fabricii 0 0 8 92 0 0 0 0 0 12 LC FG GW RM
Onitis fulgidus 0 0 7 93 0 0 0 0 0 70 LC FG TW O>M>R
Onitis inversidens 0 0 0 100 0 0 0 0 0 13 LC (LLR) FG WT M
Onitis licitus 0 0 100 0 0 0 0 0 0 2 DD (R) ?FG ?G ?
Onitis longitibialis 100 0 0 0 0 0 0 0 0 1 DD (R) ? ?S ?M
Onitis mendax 0 0 0 100 0 0 0 0 0 22 LC (LLR) FG W M
Onitis minutus 0 0 0 0 0 9 82 9 0 11 DD ?Ss GS ?R
Onitis obenbergeri 0 0 0 100 0 0 0 0 0 14 LC Ss W M>O>R
Onitis paraconfusus 0 0 0 100 0 0 0 0 0 2 DD Sa W M
(R, RR)
Onitis parainflaticollis 0 27 9 45 18 0 0 0 0 11 DD (R) ? ? ?
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onitis pecuarius 0 0 47 21 6 6 19 0 0 78 LC FG G R
Onitis perpunctatus 0 0 51 43 0 2 4 0 0 47 LC FG F>G RO
Onitis picticollis 0 0 20 78 3 0 0 0 0 40 LC Gen GW M
SURICATA 6 (2020)

Onitis pseudosetosus 0 0 0 100 0 0 0 0 0 12 LC Ss W ?MO

Onitis robustus 0 0 0 100 0 0 0 0 0 2 LC FG W M
(U, LLR)
Onitis tortuosus 0 1 64 29 5 1 1 0 0 133 LC FG G R
Onitis uncinatus 1 13 13 71 1 0 0 0 0 134 LC FG W>G>T RM
Onitis viridulus 0 1 10 84 5 0 0 0 0 92 LC FG Gen MR
Onitis westermanni 0 0 0 100 0 0 0 0 0 6 LC Gen GW ?R
Tropidonitis paradoxus 0 0 0 67 33 0 0 0 0 3 VU (RR) Sa ?WF M
Onthophagini (Clade 23)
Caccobiomorphus megaponerae 0 0 0 100 0 0 0 0 0 3 LC (R) ? ? ?Ant prey
Caccobius castaneus 0 0 0 100 0 0 0 0 0 5 DD (U) ?Gen ? ?RM
Caccobius cavatus 0 8 0 42 50 0 0 0 0 12 LC Sa ? O>MR
Caccobius ferrugineus 0 30 2 67 0 0 0 0 0 46 LC Sa G>WT O>M>R
Caccobius histerinus 0 0 0 80 20 0 0 0 0 25 LC Sa ? ?
Caccobius nigritulus 2 9 0 83 6 0 0 0 0 47 LC Sa W>T>G O>M>R
Caccobius obtusus 0 0 55 22 5 0 17 0 0 58 LC ?FG ?G ?
Cleptocaccobius convexifrons 0 0 0 100 0 0 0 0 0 4 LC Sa WG>T O>R>M
Cleptocaccobius postlutatus 0 0 35 58 8 0 0 0 0 26 LC ? ? ?
Cleptocaccobius viridicollis 9 20 13 52 6 0 0 0 0 64 LC FG>Sa W>G>T O>R>M
Digitonthophagus gazella 4 9 16 56 4 5 7 0 0 221 LC FG>Sa G>W>T ?
Digitonthophagus namaquensis 48 30 0 0 0 4 4 13 0 23 LC ? ? ?
Euonthophagus carbonarius 0 0 3 95 2 0 0 0 0 63 LC ? ? ?
Euonthophagus flavimargo 19 81 0 0 0 0 0 0 0 47 LC ?Ss ?W O>M>R
Euonthophagus jeanneli 0 0 0 100 0 0 0 0 0 12 LC Sa GS OM>R
Euonthophagus vicarius 83 0 6 0 0 3 6 3 0 36 LC ?Gen ?S ?
CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
729

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onthophagini (Clade 23) (continued)
730

Hamonthophagus acutus 29 71 0 0 0 0 0 0 0 21 LC ?Sa ?S ?

Hamonthophagus depressus 1 0 9 73 18 0 0 0 0 80 LC ?Ss ?W ?
Haroldius convexus 0 0 0 100 0 0 0 0 0 1 DD (R) ? ? ?Ant prey
Heteroclitopus remipes 0 20 0 60 20 0 0 0 0 5 DD (R) ? ? ?Termites
Hyalonthophagus alcyon 0 0 0 100 0 0 0 0 0 1 DD ?Sa ?W ?M>R
Hyalonthophagus alcyonides 0 0 12 83 5 0 0 0 0 42 LC ?Gen W>G>T O>MR
Kurtops caffrarius - - - - - - - - - 0 DD (R) ? ? ?
Kurtops quadraticeps 9 79 0 12 0 0 0 0 0 34 LC Sa SG O>M>R
Kurtops signatus 30 32 0 33 5 0 0 0 0 106 LC Sa W>G O>M>R
Milichus apicalis 0 0 0 100 0 0 0 0 0 17 LC (LLR) FG ?W M
Mimonthophagus ambiguus 0 0 0 20 80 0 0 0 0 10 LC Sa G>F ?
Mimonthophagus anomalus 0 0 0 100 0 0 0 0 0 3 LC (LLR) ? ? O>M>R
Onthophagus absyrtus 0 0 91 9 0 0 0 0 0 11 DD ?FG ?G ?
Onthophagus aequepubens 0 0 0 100 0 0 0 0 0 12 LC ? T ?
Onthophagus aeruginosus 1 0 27 67 5 0 0 0 0 113 LC Gen WT>G O>MR
Onthophagus albipennis 91 2 2 0 0 2 2 0 0 44 LC Gen ?GS ?
Onthophagus albipodex 0 0 6 94 0 0 0 0 0 16 LC ?Sa>FG G>SW ?
Onthophagus apiciosus 0 0 20 80 0 0 0 0 0 20 LC ? ? ?C
Onthophagus asperulus 0 0 67 24 5 5 0 0 0 88 LC ?FG ?G ?
Onthophagus beiranus 0 0 0 76 24 0 0 0 0 21 LC FG WT ?RM
Onthophagus bicavifrons 1 6 11 74 8 0 0 0 0 87 LC ? ? C
Onthophagus binodis 1 0 62 15 2 3 17 0 0 115 LC ?FG ?G ?R
Onthophagus bovinus 24 24 6 47 0 0 0 0 0 17 LC ? ? ?
Onthophagus cameloides 21 0 0 0 0 3 51 25 0 71 LC ?Ss G ?RM
Onthophagus cinctipennis 0 5 59 27 9 0 0 0 0 22 LC ?Sa, Ss G>W>T ?
Onthophagus cineraceus 0 0 0 0 0 0 100 0 0 2 DD ? ? ?
(R, RR)
Onthophagus convexus 47 38 0 15 0 0 0 0 0 68 LC ?Sa, Ss G>W O>M>R
Onthophagus corniculiger 0 0 0 100 0 0 0 0 0 12 DD ? ? M
(U, LLR)
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onthophagus cretus 0 0 14 57 29 0 0 0 0 14 LC ? F ?
Onthophagus cribripennis 0 1 43 44 8 2 1 0 0 84 LC ?FG ?GF O>R
Onthophagus croesulus 0 0 27 73 0 0 0 0 0 11 DD ? ? C
SURICATA 6 (2020)

Onthophagus cupricollis 0 0 20 80 0 0 0 0 0 10 DD ? ? C
Onthophagus cyaneoniger 7 7 68 14 0 0 4 0 0 28 LC Ss G ?
O. deterrens & variolosus 0 2 47 30 5 0 16 0 0 57 DD ?FG ?G C
Onthophagus discretus 0 0 10 90 0 0 0 0 0 10 LC FG>Sa T ?
Onthophagus ebenicolor 0 0 3 88 10 0 0 0 0 40 LC ?FG ?W ?C
Onthophagus ebenus 0 1 26 64 9 0 0 0 0 70 LC FG>Sa GW OC
Onthophagus fimetarius 0 2 29 67 2 0 0 0 0 96 LC ?Ss>FG Gen O>RM
Onthophagus flavolimbatus 0 5 2 83 10 0 0 0 0 42 LC Ss>FG GW OR>M
Onthophagus fritschi 43 2 47 0 0 5 0 3 0 58 LC ?Gen GS ?
Onthophagus fugitivus 0 0 19 81 0 0 0 0 0 16 DD ?FG ?SW ?
Onthophagus giraffa 0 0 0 0 0 3 97 0 0 34 LC ?Ss G>S ?
O. giuseppecarpenetoi 0 0 0 56 44 0 0 0 0 9 LC (RR) Sa G>F ?M
Onthophagus granulifer 19 73 4 4 0 0 0 0 0 48 LC Ss ?GW O>M>R
Onthophagus graphicus 0 33 0 67 0 0 0 0 0 6 DD ? ? C
Onthophagus herus 0 0 0 100 0 0 0 0 0 2 LC (LLR) ? ?TF ?OC
Onthophagus hyaena 0 0 73 0 0 0 27 0 0 30 LC ?FG ?G ?
Onthophagus immundus 0 0 0 0 0 0 90 10 0 10 LC ?Ss ?G ?
Onthophagus interstitialis 1 0 59 38 1 0 0 0 0 68 LC FG>Ss G>W O>RM
Onthophagus juvencus 0 0 0 38 63 0 0 0 0 8 LC Sa G ?
Onthophagus lacustris 0 0 0 40 60 0 0 0 0 10 LC ?Sa FT ?O>R
Onthophagus lamelliger 0 0 0 100 0 0 0 0 0 49 LC ?Gen ?W O>M>R
Onthophagus lamnifer 0 0 100 0 0 0 0 0 0 27 LC ?FG ?G ?R
Onthophagus leroyi 0 0 0 100 0 0 0 0 0 20 LC ?FG ?WFT ?
Onthophagus leucopygus 9 70 17 4 0 0 0 0 0 23 LC ?Ss ?GSW ?DGen

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
731

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onthophagini (Clade 23) (continued)
732

Onthophagus lugubris 0 0 80 10 0 5 5 0 0 40 LC ?FG ?G ?

Onthophagus minutus 0 0 0 0 0 3 90 7 0 30 LC Sa GS ?
Onthophagus monodon 0 0 100 0 0 0 0 0 0 10 LC ? ? ?
Onthophagus naso 0 0 31 38 25 6 0 0 0 16 LC ?FG ?G ?OC
Onthophagus obtusicornis 1 11 25 54 6 0 2 1 0 100 LC ?Gen G>W>T CO>M>R
Onthophagus obtutus 0 0 100 0 0 0 0 0 0 28 LC FG G ?
Onthophagus pallidipennis 0 22 4 72 2 0 0 0 0 50 LC Ss>FG W>G>T RO>M
Onthophagus parumnotatus 0 0 45 49 6 0 0 0 0 53 LC FG G O>R>M
Onthophagus pauxillus 0 2 13 83 2 0 0 0 0 46 LC ?Gen W>G>T RO>M
Onthophagus pellax 0 13 7 80 0 0 0 0 0 15 LC Ss ?G>W ?
Onthophagus peringueyi 84 3 5 0 0 5 2 0 0 58 LC ?Gen GS ?
Onthophagus pilosus 0 0 65 24 6 0 6 0 0 34 LC Ss ?G>W ?
Onthophagus plebejus 0 0 0 88 13 0 0 0 0 8 LC (LLR) ?Ss ?W O>M (M)
Onthophagus probus 58 42 0 0 0 0 0 0 0 71 LC Ss GS O>MR
Onthophagus producticollis 0 0 13 87 0 0 0 0 0 15 DD FG WT ?
Onthophagus pugionatus 4 1 23 60 11 0 0 0 0 73 LC Ss>FG T>W>G ?O>RM
Onthophagus pullus 0 0 7 89 4 0 0 0 0 27 LC Sa T>W>G ?O
Onthophagus quadrimaculatus 0 13 6 63 19 0 0 0 0 16 DD ?FG>Ss T ?C
Onthophagus quadrinodosus 0 3 9 76 12 0 0 0 0 34 LC Sa TW>G O>RM
Onthophagus rasipennis 0 0 4 94 2 0 0 0 0 47 LC ?FG or Ss W>TG ?DGen
Onthophagus scapularis 0 0 0 0 0 100 0 0 0 2 DD ? ? ?C
Onthophagus semiflavus 36 64 0 0 0 0 0 0 0 39 LC ?Ss GS O>MR
Onthophagus suffusus 0 0 0 100 0 0 0 0 0 11 LC ?Ss ?W O>M>R
Onthophagus trinodosus 0 0 0 100 0 0 0 0 0 12 DD (U) ?FG ?S ?O>R
Onthophagus ursinus 0 0 0 22 78 0 0 0 0 9 LC ?Sa ?WG ?
Onthophagus venustulus 18 63 3 16 0 0 0 0 0 67 LC Ss ?WS>G OM>R
Onthophagus verticalis 4 31 0 65 0 0 0 0 0 26 LC Ss TW>G MO>R
Onthophagus vigens 0 0 67 23 3 7 0 0 0 30 LC FG G ?
Onthophagus vinctus 2 6 11 69 12 1 0 0 0 147 LC Ss T>W>G O>MR
Onthophagus virescens 0 0 0 100 0 0 0 0 0 4 LC Sa ?W O>M>R
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Onthophagus vylderi 0 0 0 100 0 0 0 0 0 3 LC Sa ?W C
Phalops ardea 0 0 0 100 0 0 0 0 0 47 LC ?Ss ?W O>RM
Phalops boschas 0 0 0 100 0 0 0 0 0 57 LC ?Ss ?W O>RM
SURICATA 6 (2020)

Phalops bubalus 19 1 61 5 0 13 0 0 0 77 LC ?Gen ?G ?R

Phalops dregei 37 15 11 34 0 3 0 0 0 100 LC ?Gen GW O>RM
Phalops flavocinctus 0 1 15 81 3 0 0 0 0 78 LC FG G>W O>MR
Phalops pryroides 80 20 0 0 0 0 0 0 0 20 LC Ss ?SW ?
Phalops rufosignatus 28 53 6 0 0 11 3 0 0 36 LC ?Gen ?GW O>M>R
Phalops smaragdinus 0 0 4 89 7 0 0 0 0 46 LC ?Ss W>G O>MR
Phalops wittei 33 45 11 12 0 0 0 0 0 103 LC Sa GS>W O>M>R
Proagoderus aciculatus 0 0 0 40 52 8 0 0 0 25 LC Sa F ?O>RM
Proagoderus aureiceps 0 0 0 62 38 0 0 0 0 29 LC Sa G ?
Proagoderus bicallosus 0 0 0 81 19 0 0 0 0 21 LC (LLR) Ss W MO
Proagoderus chalcostolus 0 0 40 45 15 0 0 0 0 73 LC FG G ?
Proagoderus dives 0 0 0 78 22 0 0 0 0 18 LC Sa FW ?O>MR
Proagoderus furcifer 0 0 0 100 0 0 0 0 0 1 DD ?Ss ?SW M
(R, LLR)
Proagoderus lanista 10 0 68 12 8 2 0 0 0 50 LC FG G ?
Proagoderus loricatus 0 0 3 97 0 0 0 0 0 29 LC (LLR) ?Ss ?SW O>MR
(M)
Proagoderus quadrituber 0 0 17 67 17 0 0 0 0 6 LC FG WF ?
Proagoderus rangifer 0 0 0 100 0 0 0 0 0 17 LC (LLR) ?Ss SW O>MR
(M)
Proagoderus rectefurcatus 0 0 0 100 0 0 0 0 0 6 LC (LLR) ?Ss SW M
Proagoderus sapphirinus 0 52 4 44 0 0 0 0 0 48 LC Sa GW>T O>CMR
Proagoderus tersidorsis 0 0 2 94 4 0 0 0 0 50 LC FG ?W ?
Scarabaeini (Clade 4)
Escarabaeus satyrus 55 22 3 6 0 1 3 8 1 119 LC Ss GS ?RM

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
733

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Scarabaeini (Clade 4) (continued)
734

Kheper bonellii 0 0 0 0 0 0 54 46 0 24 LC Ss SG ?RM

Kheper clericus 0 0 0 100 0 0 0 0 0 4 DD (RR) FG W MR
Kheper cupreus 0 0 0 100 0 0 0 0 0 25 LC Ss SW OR
Kheper kalaharicus 0 100 0 0 0 0 0 0 0 2 LC (RR) Sa GS OR
Kheper lamarcki 0 32 3 54 11 0 0 0 0 65 LC Sa W>GT O>MR>C
Kheper nigroaeneus 0 3 9 88 0 0 0 0 0 103 LC Gen WG O>R>M
Kheper prodigiosus 0 25 0 75 0 0 0 0 0 20 LC Sa G>W M>O>R
Kheper subaeneus 0 0 14 86 0 0 0 0 0 44 LC FG WG O>MR
Kheper zurstrasseni 0 0 33 67 0 0 0 0 0 3 DD (R, ? ? ?
RR)
Pachylomera femoralis 0 16 6 65 13 0 0 0 0 68 LC Sa GW O>CMR
Pachylomera opacus 21 74 0 5 0 0 0 0 0 43 LC Sa GS O>MR
Pachysoma aesculapius 0 0 0 0 0 0 100 0 0 15 VU (RR) Sa S Rpellets
Pachysoma bennigseni 0 0 0 0 0 0 0 75 25 4 LC (RR) Sa ?S Detritus
Pachysoma endroedyi 0 0 0 0 0 0 0 100 0 1 EN (R, RR) Sa S Detritus
Pachysoma gariepinum 0 0 0 0 0 0 0 92 8 12 LC Sa ?S Detritus
Pachysoma glentoni 0 0 0 0 0 0 100 0 0 3 VU (R, RR) Sa S Detritus
Pachysoma hippocrates 0 0 0 0 0 0 34 66 0 29 LC (RR) Sa S Detritus
Pachysoma striatum 0 0 0 0 0 0 0 100 0 23 LC (RR) Sa S Rpellets
Scarabaeolus anderseni 7 85 0 7 0 0 0 0 0 27 LC Sa SG C
Scarabaeolus andreaei 0 0 0 100 0 0 0 0 0 1 LC Sa ?S ?O
Scarabaeolus bohemani 50 11 16 21 0 0 1 0 0 123 LC Gen GS CO>MR
Scarabaeolus carniphilus 0 100 0 0 0 0 0 0 0 1 DD Sa G C>R
Scarabaeolus clanceyi 0 0 0 100 0 0 0 0 0 6 LC Sa G O
Scarabaeolus damarensis 12 86 0 2 0 0 0 0 0 42 LC Sa GS OR>M
Scarabaeolus flavicornis 23 47 5 5 0 0 3 15 3 75 LC Sa G OC>MR
Scarabaeolus fritschi 79 0 0 0 0 0 0 18 3 34 LC Sa GS ?
Scarabaeolus gracai 0 0 0 100 0 0 0 0 0 2 DD (RR) Sa ? ?
Scarabaeolus inoportunus 27 70 3 0 0 0 0 0 0 33 LC Sa GS DGenC
Scarabaeolus inquisitus 22 44 11 22 0 0 0 0 0 18 LC Sa GS C>dung
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Scarabaeolus intricatus 0 0 0 0 0 0 68 33 0 40 LC Ss S>G ?
Scarabaeolus karrooensis 86 14 0 0 0 0 0 0 0 21 LC ? ? ?
Scarabaeolus kochi 35 65 0 0 0 0 0 0 0 26 LC Sa GS CO>MR
SURICATA 6 (2020)

Scarabaeolus lucidulus 0 0 0 100 0 0 0 0 0 2 DD Sa ?W Banana-

CO
Scarabaeolus megaparvulus 100 0 0 0 0 0 0 0 0 18 LC ?Gen ?SG ?
Scarabaeolus niemandi 0 0 0 100 0 0 0 0 0 4 DD (RR) ?Sa ?W ?
Scarabaeolus pabulator 81 5 0 5 0 0 0 5 5 21 LC Sa S ?
Scarabaeolus parvulus 50 46 0 0 0 0 0 2 2 46 LC Sa SG ?DGen
Scarabaeolus planipennis 0 0 0 80 20 0 0 0 0 5 LC (RR) Sa GW>F ?O
Scarabaeolus reichei 0 0 0 0 0 0 46 54 0 24 LC Ss GS ?C+dung
Scarabaeolus rubripennis 0 0 0 0 0 0 0 75 25 4 LC Sa ? ?
Scarabaeolus soutpansbergensis 0 0 0 100 0 0 0 0 0 7 DD (RR) ?Sa ?SW ?
Scarabaeus alienus 0 0 0 0 0 0 0 100 0 2 DD (R) Ss S ?M
Scarabaeus ambiguus 7 26 35 32 0 0 0 0 0 57 LC ?Ss G ?
Scarabaeus basuto 43 0 57 0 0 0 0 0 0 42 LC ?FG G ?R
Scarabaeus bornemisszai 0 0 0 0 100 0 0 0 0 10 NT (RR) Sa F ?O
Scarabaeus caffer 0 0 45 50 5 0 0 0 0 22 LC FG G OR
Scarabaeus spretus & convexus 0 0 36 9 5 5 41 5 0 22 DD ? ? ?
Scarabaeus costatus 29 65 3 0 0 0 0 3 0 31 LC Sa GS O>R
Scarabaeus deludens 0 3 9 88 0 0 0 0 0 32 LC FG SW MR
Scarabaeus ebenus 0 0 0 82 18 0 0 0 0 11 LC (U) Sa W O
Scarabaeus funebris 13 13 40 33 0 0 0 0 0 15 DD (U) FG GSW ?
Scarabaeus galenus 0 0 3 97 0 0 0 0 0 30 LC Ss W ?Rpellets
Scarabaeus geminogalenus 0 0 0 77 23 0 0 0 0 13 LC Sa G ?
Scarabaeus goryi 0 15 6 65 15 0 0 0 0 62 LC Sa Gen O>RM
Scarabaeus heqvisti 0 0 33 67 0 0 0 0 0 3 DD (RR) Sa G ?
Scarabaeus hottentorum 0 0 0 0 0 0 0 100 0 5 DD (RR) Ss S OR
CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
735

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 1: Summary Table 1. Distributions of 484 scarabaeine dung beetle species across biomes of South Africa (Mucina & Rutherford 2006) represented by proportions of occupied 1/16th degree
squares; data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; putative ecolog-
ical bias assessed from available published or unpublished data (see Field Survey section under Introduction); see Synthesis section for key to division of tribes into clades of related species (continued)

Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Scarabaeini (Clade 4) (continued)
736

Scarabaeus interstitialis 0 13 13 73 0 0 0 0 0 15 LC (U) ? GW ?

Scarabaeus karae 0 0 100 0 0 0 0 0 0 3 DD ? G ?
(R, RR)
Scarabaeus piliventris 0 0 0 0 0 0 40 60 0 5 DD ?Ss GS ?
(U, RR)
Scarabaeus proboscideus 25 37 0 0 0 0 14 22 2 81 LC Sa ?S ?DGen
Scarabaeus proximus 0 0 0 0 0 0 0 90 10 10 LC Sa S ?
Scarabaeus rixosus - - - - - - - - - 0 DD ? ? ?
(R, RR)
Scarabaeus rugosus 0 0 0 0 0 0 55 45 0 33 LC Sa S ?
Scarabaeus rusticus 0 2 26 72 0 0 0 0 0 85 LC ?Gen W ?
Scarabaeus savignyi 0 0 0 0 0 33 67 0 0 6 DD (U) ?FG ?S ?
Scarabaeus schulzae 0 0 0 100 0 0 0 0 0 8 DD (RR) ? ?WF ?
Scarabaeus suri 0 0 0 0 0 0 83 17 0 23 LC Ss SG ?RM
Scarabaeus viator 81 6 4 0 0 4 3 1 0 67 LC Gen GS ?
Scarabaeus vicinus 52 33 2 12 0 0 0 0 0 42 LC Sa GS ?Rpellets
Scarabaeus westwoodi 0 0 100 0 0 0 0 0 0 6 LC FG G ?R
Scarabaeus zambesianus 0 33 0 67 0 0 0 0 0 33 LC Sa SW DGen
Sceliages adamastor 20 0 30 0 0 10 30 10 0 10 DD ? ?S Mi
Sceliages brittoni 0 0 0 0 0 0 42 58 0 12 LC Sa S Mi
Sceliages difficilis 0 0 29 71 0 0 0 0 0 24 LC ?Sa ?GS ?Mi
Sceliages gagates 0 0 0 50 50 0 0 0 0 4 LC Sa ?G ?Mi
Sceliages granulatus 25 75 0 0 0 0 0 0 0 4 LC Sa GS Mi
Sceliages hippias 0 0 19 81 0 0 0 0 0 31 LC ? ?SW Mi
Sisyphini (Clade 18)
Neosisyphus barbarossa 0 0 83 0 0 6 11 0 0 18 LC ?FG G ?R
Neosisyphus calcaratus 0 0 6 92 2 0 0 0 0 50 LC Ss> FG SW R>O>M
Neosisyphus confrater 0 0 38 33 29 0 0 0 0 24 LC FG, Ss G ?
Neosisyphus fortuitus 0 0 10 78 12 0 0 0 0 41 LC ?Gen T>W>G ?
Neosisyphus infuscatus 0 0 12 70 12 6 0 0 0 33 LC FG, Ss ?W>GF R>OM
SURICATA 6 (2020)
Tribes and species % of 1/16th degree squares occupied in summer rainfall biomes % in winter rainfall biomes Total N Conser- Putative ecological bias
(Clades, see Figure 5) Nama Kalahari Grassland East Indian Ocean Albany Fynbos Succulent Desert 1/16th vation Soil Veg. Food
Karoo Savanna Savanna Coastal Belt Thicket Karoo degree status (low
squares frequency)
Neosisyphus kuehni 0 0 100 0 0 0 0 0 0 7 DD (U) ?Gen G ?R
Neosisyphus macrorubrus 32 35 31 2 0 0 0 0 0 65 LC ?Gen ?SG>W ?R
Neosisyphus mirabilis 0 0 0 36 45 9 9 0 0 11 LC Ss>FG FT ?
SURICATA 6 (2020)

Neosisyphus quadricollis 17 0 0 0 0 0 83 0 0 6 DD ? ? ?
Neosisyphus rubrus 0 5 38 50 3 3 1 0 0 172 LC FG>Ss G>W>T R>MO
Neosisyphus spinipes 0 0 7 63 20 3 7 0 0 71 LC Gen ?G>W>T ?R>MO
Sisyphus caffer 0 2 75 17 5 0 2 0 0 59 LC ?FG G OR>M
Sisyphus costatus 0 0 82 15 3 0 0 0 0 34 LC FG G O>R>M
Sisyphus fasciculatus 0 0 17 75 8 0 0 0 0 24 LC FG FT>W OR>M
Sisyphus goryi 0 2 9 86 1 0 1 0 0 85 LC Gen W>GT OR>M
Sisyphus impressipennis 0 0 3 97 0 0 0 0 0 31 LC ?Ss TF ?O>RM
Sisyphus manni 0 0 41 54 5 0 0 0 0 37 LC ?Ss GW ?
Sisyphus muricatus 0 0 61 4 0 0 35 0 0 23 LC ?FG G ?
Sisyphus nanniscus 0 0 2 77 21 0 0 0 0 43 LC Gen,Ss>FG T O>R>M
Sisyphus neobornemisszanus 0 0 0 0 100 0 0 0 0 5 NT (LLR) Sa F ?
Sisyphus oralensis 0 0 0 42 58 0 0 0 0 12 LC Sa FT ?
Sisyphus perissinottoi 0 0 0 0 0 67 33 0 0 3 DD ? ? ?
Sisyphus sordidus 0 0 0 62 38 0 0 0 0 29 LC Sa G ?
Sisyphus umbraphilus 0 0 0 100 0 0 0 0 0 4 DD ?Sa T O>R
Total numbers of species 108 148 241 320 153 86 130 71 12 484

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range
SOIL: Sa = Sand; Ss = Sandy Soils (Sand, Sandy loam); FG = Finer Grained (Sandy loam, Sandy clay loam, Clay); Gen = Generalist; ? = unknown or uncertain; > = is greater than;.
VEG. (VEGETATION): G = Pasture/Grassland; S = Scrub, Shrubland; W = Open Woodland, Woodland; F = Forest; T = Shaded Thicket; Gen = Generalist; > = is greater than; ? = unknown or
uncertain.
FOOD: O = Omnivore dung, R = Ruminant herbivore dung; M = Monogastric herbivore dung; C = Carrion; Mi = Millipedes; F = Fungus; DGen = Dung Generalist; Dpellets = Dung pellets; Hyrax
737

= Hyrax middens; > = is greater than; ? = unknown or uncertain.

APPENDIX 2: Summary Table 2. Distributions of 217 scarabaeine dung beetle species across ecoregions of Namibia and Botswana (Olson et al. 2001) represented by proportions of occupied 1/16th degree squares;
data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; see Synthesis section for key to
division of tribes into clades of related species

Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
738

Ateuchini (Clade 1)
Coptorhina auspicata 0 0 0 0 0 67 0 33 0 0 0 6 LC
Coptorhina nitidipennis 0 0 0 0 0 33 67 0 0 0 0 3 LC
Delopleurus darrenmanni 0 0 0 0 0 0 0 100 0 0 0 2 DD
Delopleurus gilleti 0 0 0 0 100 0 0 0 0 0 0 1 LC
Delopleurus pullus 0 0 0 0 50 0 0 0 50 0 0 4 LC
Sarophorus angolensis 0 0 0 0 100 0 0 0 0 0 0 1 DD (RR)
Sarophorus costatus 0 0 0 0 0 50 0 0 0 50 0 2 LC
Canthonini (Clade 2)
Dicranocara deschodti 0 100 0 0 0 0 0 0 0 0 0 2 VU (R, RR)
Dicranocara inexpectata 0 100 0 0 0 0 0 0 0 0 0 1 DD (R, RR)
Dicranocara vandersmisseni 0 100 0 0 0 0 0 0 0 0 0 2 DD (R,RR)
Drogo stalsi 0 100 0 0 0 0 0 0 0 0 0 1 DD (R, RR)
Namakwanus irishi 0 0 0 0 0 100 0 0 0 0 0 1 DD (R, RR)
Namakwanus scholtzi 0 0 100 0 0 0 0 0 0 0 0 1 DD (R, RR)
Namaphilus ameibensis 0 0 0 0 100 0 0 0 0 0 0 1 DD (R, RR)
Namaphilus davisi 0 0 0 0 0 100 0 0 0 0 0 1 DD (R, RR)
Namaphilus endroedyi 0 0 0 0 100 0 0 0 0 0 0 2 DD (R, RR)
Odontoloma louwi 0 0 0 0 56 22 11 0 11 0 0 9 LC
Versicorpus erongoensis 0 0 0 0 100 0 0 0 0 0 0 1 DD (R, RR)
Versicorpus streyi 0 0 0 0 100 0 0 0 0 0 0 1 DD (R, RR)
Ateuchini (Clade 5)
Pedaria brancoi 0 0 0 0 0 50 0 0 0 0 50 2 DD
Pedaria cylindrica 0 0 0 0 0 0 50 0 0 0 50 2 LC
Pedaria picea 0 0 0 0 0 50 50 0 0 0 0 4 LC
Coprini (Clade 5)
Copris amyntor 0 0 0 0 0 22 56 11 0 11 0 9 LC
Copris bootes 0 0 0 0 0 0 0 100 0 0 0 3 LC (LLR)
Copris cassius 0 0 0 0 0 85 15 0 0 0 0 20 LC
SURICATA 6 (2020)
Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
Copris cornifrons 0 0 0 0 0 71 29 0 0 0 0 7 LC (U)
Copris denticulatus 0 0 0 0 0 0 75 0 0 25 0 4 LC
SURICATA 6 (2020)

Copris elphenor 0 0 0 0 5 44 20 15 10 5 0 59 LC
Copris evanidus 0 0 0 0 0 7 57 0 0 36 0 14 LC
Copris gracilis 0 0 0 0 29 45 2 0 24 0 0 42 LC
Copris inhalatus 0 0 0 0 0 71 14 0 0 0 14 7 LC (U)
Copris laioides 0 0 0 0 0 40 13 0 47 0 0 15 LC
Copris mesacanthus 0 0 0 0 0 0 0 75 0 0 25 4 LC
Copris puncticollis 0 0 0 0 0 0 0 71 29 0 0 7 LC
Copris subsidens 0 0 0 0 40 33 0 0 26 0 0 42 LC
Copris vilhenai 0 0 0 0 0 0 0 80 0 20 0 5 LC
Coprini (Clade 13)
Catharsius aegeus 0 0 0 0 0 0 29 43 0 14 14 7 LC
Catharsius calaharicus 0 0 0 0 6 62 24 6 3 0 0 34 LC
Catharsius heros 0 0 0 0 11 0 33 22 22 0 11 9 LC (LLR)
Catharsius melancholicus 0 0 0 0 17 50 17 8 0 0 8 12 LC
Catharsius philus 0 0 0 0 0 0 83 0 0 17 0 6 LC
Catharsius tricornutus 0 0 0 0 0 0 0 60 40 0 0 5 LC
Catharsius ulysses 0 0 0 0 13 58 17 0 13 0 0 24 LC
Metacatharsius anderseni 0 0 0 0 0 100 0 0 0 0 0 8 LC
Metacatharsius dentinum 0 0 4 0 32 43 11 7 0 0 4 28 LC
Metacatharsius exiguiformis 0 0 9 4 27 53 4 0 0 2 0 45 LC
Metacatharsius exiguus 0 0 0 0 0 82 9 0 9 0 0 11 LC
Metacatharsius ferreirae 0 0 0 0 0 25 50 25 0 0 0 4 DD
Metacatharsius freyi 0 0 0 0 0 25 75 0 0 0 0 4 DD
Metacatharsius latifrons 0 0 0 0 0 100 0 0 0 0 0 11 LC
Metacatharsius marani 0 0 4 0 28 60 0 0 8 0 0 25 LC
Metacatharsius opacus 0 0 0 0 10 31 25 17 12 2 4 52 LC
Metacatharsius pumilioni- 0 0 0 0 25 50 25 0 0 0 0 8 LC
formis
Metacatharsius troglodytes 0 0 0 0 0 29 32 18 21 0 0 34 LC

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
739

FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range

APPENDIX 2: Summary Table 2. Distributions of 217 scarabaeine dung beetle species across ecoregions of Namibia and Botswana (Olson et al. 2001) represented by proportions of occupied 1/16th degree squares;
data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; see Synthesis section for key to
division of tribes into clades of related species (continued)

Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
740

Coprini (Clade 14)

Heliocopris andersoni 0 3 0 0 23 39 6 3 23 0 3 31 LC
Heliocopris atropos 0 0 0 0 6 63 6 13 6 0 6 16 LC
Heliocopris faunus 0 0 7 0 34 31 3 0 21 3 0 29 LC
Heliocopris hamadryas 0 0 0 0 0 50 0 50 0 0 0 2 LC
Heliocopris japetus 0 0 0 0 0 53 17 20 10 0 0 30 LC
Heliocopris neptunus 0 0 0 0 25 0 25 25 0 25 0 4 LC
Canthonini (Clade 16)
Chalconotus convexus 0 0 2 0 37 14 5 16 14 7 5 43 LC
Canthonini (Clade 20)
Hammondantus psammophilus 17 0 67 0 17 0 0 0 0 0 0 6 LC (R)
Pycnopanelus krikkeni 0 4 12 4 62 19 0 0 0 0 0 26 LC
Gymnopleurini (Clade 15)
Allogymnopleurus splendidus 0 0 0 0 0 52 17 13 0 13 4 23 LC
Garreta nitens 0 0 0 0 28 17 17 0 39 0 0 18 LC
Garreta wahlbergi 0 0 0 0 0 0 0 0 0 67 33 3 LC
Gymnopleurus aenescens 0 0 4 4 34 28 14 10 2 2 2 50 LC
Gymnopleurus andreaei 0 8 21 4 58 8 0 0 0 0 0 24 LC
Gymnopleurus asperrimus 0 14 0 0 0 86 0 0 0 0 0 7 LC
Gymnopleurus humanus 0 4 13 5 54 20 0 0 5 0 0 80 LC
Gymnopleurus humeralis 0 0 0 0 0 0 0 0 0 100 0 1 LC
Gymnopleurus ignitus 0 0 0 0 0 0 13 50 0 25 13 8 LC
Gymnopleurus imitator 0 0 0 0 0 100 0 0 0 0 0 1 DD (R, RR)
Gymnopleurus pumilus 0 0 0 0 23 23 19 15 19 0 0 26 LC
Gymnopleurus virens 0 0 0 0 0 50 50 0 0 0 0 2 LC
Oniticellini (clade 23)
Drepanocerus kirbyi 0 0 0 0 0 0 0 100 0 0 0 2 LC
Euoniticellus intermedius 3 3 3 0 21 42 8 5 11 3 3 38 LC
Euonticellus kawanus 0 0 0 0 33 0 0 33 33 0 0 6 DD (U, LLR)
SURICATA 6 (2020)
Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
Euoniticellus triangulatus 0 0 0 0 0 0 0 0 0 0 100 1 LC
Latodrepanus laticollis 0 0 0 0 0 0 50 50 0 0 0 4 LC
SURICATA 6 (2020)

Liatongus militaris 0 0 0 0 0 0 0 100 0 0 0 1 LC

Oniticellus egregius 0 0 0 0 20 0 20 0 60 0 0 5 LC (U)
Oniticellus formosus 0 0 0 0 0 0 50 50 0 0 0 2 LC
Oniticellus planatus 0 0 0 0 0 0 0 50 0 0 50 2 LC
Paraixodina saegeri 0 0 0 0 0 0 0 100 0 0 0 2 DD (LLR)
Tiniocellus spinipes 0 0 0 0 0 0 0 0 0 0 100 3 LC
Tragiscus dimidiatus 0 0 0 0 0 0 0 50 50 0 0 2 LC
Onitini (Clade 22)
Cheironitis audens 0 0 0 0 14 0 0 0 86 0 0 7 LC
Cheirontis hoplosternus 0 0 0 0 0 20 20 0 60 0 0 5 LC
Cheironitis imitator 0 0 0 0 0 0 0 100 0 0 0 3 DD
Cheironitis indicus 0 20 40 0 40 0 0 0 0 0 0 5 LC
Cheironitis scabrosus 10 30 10 0 40 10 0 0 0 0 0 10 LC
Gilletellus porculus 0 0 0 0 0 0 0 0 100 0 0 1 DD (R, LLR)
Heteronitis castelnaui 0 0 0 0 15 0 8 15 46 0 15 13 LC
Megalonitis bohemani 0 0 0 0 0 0 25 25 25 0 25 4 DD (R, LLR)
Onitis aeruginosus 0 0 0 0 0 0 50 0 0 0 50 2 LC
Onitis alexis 0 0 0 0 23 28 20 14 11 2 2 64 LC
Onitis aygulus 0 50 0 0 0 50 0 0 0 0 0 4 LC
Onitis bilobatus 0 0 0 0 100 0 0 0 0 0 0 1 DD (R, RR)
Onitis caffer 0 0 0 0 0 100 0 0 0 0 0 2 LC
Onitis confusus 0 0 100 0 0 0 0 0 0 0 0 1 LC
Onitis deceptor 0 0 0 0 0 0 0 100 0 0 0 2 LC
Onitis fulgidus 0 0 0 0 0 0 0 57 0 29 14 7 LC
Onitis inversidens 0 0 0 0 0 0 0 100 0 0 0 2 LC (LLR)
Onitis mendax 0 0 0 0 0 0 0 0 80 0 20 5 LC (LLR)
Onitis mniszechi 0 0 20 0 20 40 0 0 20 0 0 5 DD
Onitis obenbergeri 0 0 0 0 0 0 0 100 0 0 0 1 LC
Onitis obscurus 0 0 0 0 24 12 18 0 47 0 0 17 LC

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
741

FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range

APPENDIX 2: Summary Table 2. Distributions of 217 scarabaeine dung beetle species across ecoregions of Namibia and Botswana (Olson et al. 2001) represented by proportions of occupied 1/16th degree squares;
data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; see Synthesis section for key to
division of tribes into clades of related species (continued)

Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
742

Onitini (Clade 22) (continued)

Onitis orthopus 0 0 0 0 0 0 50 25 0 0 25 4 LC
Onitis parainflaticollis 0 0 0 0 0 100 0 0 0 0 0 1 DD (R)
Onitis picticollis 0 0 0 0 100 0 0 0 0 0 0 1 LC
Onitis robustus 0 0 0 0 0 0 25 50 25 0 0 4 LC (U, LLR)
Onitis setosus 0 0 0 0 57 0 0 0 43 0 0 7 LC
Onitis uncinatus 0 0 0 0 20 24 16 10 26 4 0 50 LC
Onitis viridulus 0 0 0 0 0 20 20 60 0 0 0 5 LC
Onitis westermanni 0 0 0 0 0 0 33 0 0 67 0 3 LC
Platyonitis bicuariensis 0 0 0 0 0 0 0 0 100 0 0 1 VU (R, LLR)
Onthophagini (Clade 23)
Caccobius castaneus 0 0 0 0 40 0 20 0 20 0 20 5 DD (U)
Caccobius cavatus 0 0 0 0 0 20 20 40 0 0 20 5 LC
Caccobius ferrugineus 0 0 0 3 16 31 31 9 3 0 6 32 LC
Caccobius histerinus 0 0 0 0 0 0 0 100 0 0 0 2 LC
Caccobius nigritulus 0 0 0 0 0 45 18 27 0 0 9 11 LC
Cleptocaccobius convexifrons 0 0 0 0 0 33 33 0 0 0 33 3 LC
Cleptocaccobius viridicollis 0 0 3 0 67 13 10 0 3 0 3 30 LC
Digitonthophagus gazella 0 0 0 0 0 17 48 30 0 0 4 23 LC
Digitonthophagus namaquensis 0 6 10 0 65 13 0 0 6 0 0 31 LC
Digitonthophagus viridicollis 0 0 0 0 39 25 7 0 21 0 7 28 LC
Euonthophagus carbonarius 0 0 0 0 0 0 44 56 0 0 0 9 LC
Euonthophagus flavimargo 0 0 0 0 0 100 0 0 0 0 0 7 LC
Hamonthophagus acutus 0 11 22 0 28 33 6 0 0 0 0 18 LC
Hamonthophagus depressus 0 0 0 0 75 8 17 0 0 0 0 12 LC
Hamonthophagus fallax 0 0 0 0 0 0 0 40 0 0 60 5 DD
Hyalonthophagus alcyon 0 0 0 0 0 0 0 25 25 0 50 4 DD
Hyalonthophagus alcyonides 0 0 0 0 0 0 100 0 0 0 0 2 LC
Kurtops quadraticeps 0 0 0 0 7 63 19 7 0 0 4 27 LC
Kurtops signatus 0 0 0 0 11 53 21 11 3 0 3 38 LC
Milichus apicalis 0 0 0 0 50 0 0 50 0 0 0 4 LC (LLR)
SURICATA 6 (2020)
Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
Mimonthophagus anomalus 0 0 0 0 0 0 0 100 0 0 0 3 LC (LLR)
Mimonthophagus limbibasis 0 0 0 0 0 0 100 0 0 0 0 1 DD (R, RR)
SURICATA 6 (2020)

Onthophagus aequepubens 0 0 0 0 80 20 0 0 0 0 0 5 LC
Onthophagus aeruginosus 0 0 7 0 20 33 13 27 0 0 0 15 LC
Onthophagus albipodex 0 0 0 0 0 0 100 0 0 0 0 1 LC
Onthophagus apiciosus 0 0 0 0 0 0 0 100 0 0 0 1 LC
Onthophagus axillaris 0 0 14 0 43 43 0 0 0 0 0 14 DD
Onthophagus bayeri 0 0 0 0 50 50 0 0 0 0 0 8 DD
Onthophagus bicavifrons 0 0 0 0 47 29 18 6 0 0 0 17 LC
Onthophagus bovinus 0 0 0 0 30 30 10 0 30 0 0 10 LC
Onthophagus convexus 0 0 6 0 13 44 13 19 0 0 6 16 LC
Onthophagus ebenicolor 0 0 0 0 13 0 13 25 13 38 0 8 LC
Onthophagus ebenus 0 0 0 0 0 0 0 100 0 0 0 2 LC
Onthophagus fimetarius 0 0 0 0 58 11 0 11 16 0 5 19 LC
Onthophagus flavolimbatus 0 0 0 0 0 0 67 33 0 0 0 3 LC
Onthophagus gonopygus 0 0 27 13 60 0 0 0 0 0 0 15 LC
Onthophagus granulifer 0 0 0 0 4 61 22 4 4 0 4 23 LC
Onthophagus graphicus 0 0 0 0 100 0 0 0 0 0 0 1 DD
Onthophagus juvencus 0 0 0 0 0 0 0 0 0 0 100 1 LC
Onthophagus kochi 0 0 0 0 60 0 40 0 0 0 0 5 DD
Onthophagus lamelliger 0 0 0 0 0 0 22 33 0 33 11 9 LC
Onthophagus leucopygus 0 0 0 0 0 75 25 0 0 0 0 4 LC
Onthophagus obtusicornis 0 0 0 0 20 80 0 0 0 0 0 15 LC
Onthophagus pallidipennis 0 0 0 0 0 71 18 6 0 0 6 17 LC
Onthophagus plebejus 0 0 0 0 0 0 0 57 43 0 0 7 LC (LLR)
Onthophagus probus 0 4 31 0 50 15 0 0 0 0 0 26 LC
Onthophagus pullus 0 0 0 0 80 20 0 0 0 0 0 5 LC
Onthophagus quadrimaculatus 0 0 0 0 38 0 13 0 25 25 0 8 DD
Onthophagus quadrinodosus 0 0 0 0 0 60 0 20 0 0 20 5 LC
Onthophagus rasipennis 0 0 0 0 20 20 40 0 20 0 0 5 LC
Onthophagus semiflavus 0 3 26 3 42 23 3 0 0 0 0 31 LC

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
743

FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range

APPENDIX 2: Summary Table 2. Distributions of 217 scarabaeine dung beetle species across ecoregions of Namibia and Botswana (Olson et al. 2001) represented by proportions of occupied 1/16th degree squares;
data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; see Synthesis section for key to
division of tribes into clades of related species (continued)

Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
744

Onthophagini (Clade 23) (continued)

Onthophagus suffusus 0 0 0 0 0 0 29 29 29 0 14 7 LC
Onthophagus tricorniger 0 0 0 0 40 20 0 0 40 0 0 5 DD
Onthophagus venustulus 0 0 2 0 28 34 19 6 6 0 4 47 LC
Onthophagus verticalis 0 0 0 0 0 56 17 11 6 0 11 18 LC
Onthophagus vinctus 0 0 0 0 31 16 20 18 4 2 8 49 LC
Onthophagus virescens 0 0 0 0 0 0 0 75 0 0 25 4 LC
Onthophagus vylderi 0 0 0 0 0 33 0 33 0 0 33 3 LC
Phalops ardea 0 0 0 0 0 0 0 0 50 50 0 2 LC
Phalops boschas 0 0 0 0 0 0 40 50 0 0 10 10 LC
Phalops dregei 0 0 0 0 0 0 33 17 33 17 0 6 LC
Phalops flavocinctus 0 0 0 0 0 0 40 40 0 20 0 5 LC
Phalops pauliani 0 0 0 0 9 64 0 0 27 0 0 11 LC
Phalops plancus 0 0 14 7 79 0 0 0 0 0 0 14 LC
Phalops prasinus 0 0 4 0 63 26 0 0 7 0 0 27 LC
Phalops pryroides 0 7 0 0 40 40 0 0 13 0 0 15 LC
Phalops rufosignatus 0 4 4 0 15 56 0 0 22 0 0 27 LC
Phalops smaragdinus 0 0 0 0 0 0 100 0 0 0 0 1 LC
Phalops wittei 0 2 4 0 15 50 13 2 12 0 2 52 LC
Phalops zuninoi 0 0 0 0 0 0 0 0 100 0 0 1 LC
Proagoderus bicallosus 0 0 0 0 0 0 0 50 13 0 38 8 LC (LLR)
Proagoderus furcifer 0 0 0 0 0 0 100 0 0 0 0 2 DD (R, LLR)
Proagoderus lanista 0 0 0 0 0 33 0 0 67 0 0 3 LC
Proagoderus loricatus 0 0 0 0 0 0 0 100 0 0 0 2 LC (LLR)
Proagoderus plato 0 0 0 0 0 0 0 0 0 0 100 1 DD (R, LLR)
Proagoderus rangifer 0 0 0 0 0 0 0 0 0 100 0 1 LC (LLR)
Proagoderus sapphirinus 0 0 0 0 17 63 8 8 4 0 0 24 LC
Scarabaeini (Clade 4)
Escarabaeus remii 0 0 0 0 0 0 17 33 0 0 50 6 LC
Escarabaeus satyrus 3 10 12 5 39 23 5 1 0 1 0 77 LC
SURICATA 6 (2020)
Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
Kheper cupreus 0 0 0 0 27 36 14 5 14 5 0 22 LC
Kheper kalaharicus 0 0 0 0 0 100 0 0 0 0 0 1 LC (RR)
SURICATA 6 (2020)

Kheper lamarcki 0 0 0 0 0 46 12 38 0 4 0 26 LC
Kheper nigroaeneus 0 0 0 0 0 20 40 20 0 20 0 5 LC
Kheper prodigiosus 0 0 0 0 3 38 21 29 6 3 0 34 LC
Kheper subaeneus 0 0 0 0 0 0 67 0 0 33 0 3 LC
Kheper vethi 0 0 0 50 50 0 0 0 0 0 0 2 DD
Pachylomera femoralis 0 0 0 0 0 52 19 21 7 0 0 42 LC
Pachylomera opacus 0 0 0 0 0 80 20 0 0 0 0 5 LC
Pachysoma bennigseni 92 0 8 0 0 0 0 0 0 0 0 13 LC (RR)
Pachysoma denticolle 5 0 83 0 13 0 0 0 0 0 0 40 LC
Pachysoma fitzsimonsi 0 0 78 0 22 0 0 0 0 0 0 9 LC (RR)
Pachysoma gariepinum 85 15 0 0 0 0 0 0 0 0 0 20 LC
Pachysoma rodriguesi 0 0 96 0 4 0 0 0 0 0 0 23 LC
Pachysoma rotundigena 0 0 71 0 29 0 0 0 0 0 0 14 LC (RR)
Pachysoma schinzi 0 40 60 0 0 0 0 0 0 0 0 5 DD (RR)
Pachysoma valeflorae 100 0 0 0 0 0 0 0 0 0 0 2 DD (R, RR)
Scarabaeolus afronitidus 0 0 0 0 0 100 0 0 0 0 0 3 DD
Scarabaeolus anderseni 0 0 0 0 0 79 21 0 0 0 0 14 LC
Scarabaeolus bohemani 0 4 4 0 46 29 11 0 4 4 0 28 LC
Scarabaeolus canaliculatus 50 0 50 0 0 0 0 0 0 0 0 2 LC (R, RR)
Scarabaeolus carniphilus 0 0 0 0 0 100 0 0 0 0 0 2 DD
Scarabaeolus damarensis 0 0 0 0 0 60 10 25 5 0 0 20 LC
Scarabaeolus flavicornis 0 0 8 0 21 53 11 5 3 0 0 38 LC
Scarabaeolus fritschi 0 0 100 0 0 0 0 0 0 0 0 2 LC
Scarabaeolus inoportunus 0 0 0 0 0 78 22 0 0 0 0 9 LC
Scarabaeolus inquisitus 0 0 0 0 0 0 100 0 0 0 0 3 LC
Scarabaeolus karrooensis 0 0 25 0 50 25 0 0 0 0 0 4 LC
Scarabaeolus kochi 0 0 0 0 0 78 22 0 0 0 0 9 LC
Scarabaeolus lucidulus 0 0 0 0 0 0 75 0 0 0 25 4 DD
Scarabaeolus megaparvulus 0 13 25 0 50 13 0 0 0 0 0 8 LC
Scarabaeolus namibensis 0 0 78 0 22 0 0 0 0 0 0 9 LC
CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
745

FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range

APPENDIX 2: Summary Table 2. Distributions of 217 scarabaeine dung beetle species across ecoregions of Namibia and Botswana (Olson et al. 2001) represented by proportions of occupied 1/16th degree squares;
data comprising 20% or greater of total records are highlighted in yellow; conservation status assessed using IUCN Red List categories (IUCN 2001) and frequency of records; see Synthesis section for key to
division of tribes into clades of related species (continued)

Tribes and species % of Arid Namibia & Botswana ecoregions (AT 13++) % of Mesic Namibia & Botswana savanna ecoregions (AT07++) Total N Conservation
(clades, see Figure 5) Succulent Nama Namib Kaokoveld Namibian Kalahari Kalahari Zambezian Angolan Southern Zambezian 1/16th status
Karoo Karoo Desert Desert Savanna Xeric Acacia & Mopane Mopane African Baikiaea degree (low frequency)
Woodlands Savanna Baikiaea Woodlands Woodlands Bushveld Woodlands squares
Woodlands
746

Scarabaeini (Clade 4) (continued)

Scarabaeolus obsoletepunctatus 0 0 0 0 100 0 0 0 0 0 0 1 DD (R, RR)
Scarabaeolus pabulator 0 0 60 0 40 0 0 0 0 0 0 5 LC
Scarabaeolus parvulus 0 10 20 5 50 15 0 0 0 0 0 20 LC
Scarabaeolus rubripennis 17 0 71 0 13 0 0 0 0 0 0 24 LC
Scarabaeus alienus 0 0 100 0 0 0 0 0 0 0 0 1 DD (R)
Scarabaeus ambiguus 0 0 5 0 5 63 11 0 16 0 0 19 LC
Scarabaeus cognatus 0 0 40 20 40 0 0 0 0 0 0 5 DD
Scarabaeus costatus 0 27 27 9 18 18 0 0 0 0 0 11 LC
Scarabaeus deludens 0 0 0 0 25 0 25 0 50 0 0 4 LC
Scarabaeus funebris 0 0 0 0 0 0 0 0 100 0 0 3 DD (U)
Scarabaeus galenus 0 0 0 0 0 0 0 50 50 0 0 2 LC
Scarabaeus goryi 0 0 0 0 9 31 31 25 3 0 0 32 LC
Scarabaeus plausibilis - - - - - - - - - - - 0 DD
Scarabaeus proboscideus 0 0 5 3 18 65 8 0 0 3 0 40 LC
Scarabaeus proximus 83 0 17 0 0 0 0 0 0 0 0 6 LC
Scarabaeus vicinus 0 0 0 0 0 78 11 0 11 0 0 9 LC
Scarabaeus zambesianus 0 0 0 0 0 22 44 30 0 0 4 27 LC
Sceliages granulatus 0 0 0 0 0 100 0 0 0 0 0 2 LC
Sisyphini (Clade 18)
Neosisyphus calcaratus 0 0 0 0 0 29 14 43 0 14 0 7 LC
Neosisyphus macrorubrus 0 0 0 0 0 0 0 0 100 0 0 3 LC
Neosisyphus rubrus 0 0 0 0 0 50 0 0 0 50 0 2 LC
Sisyphus alveatus 0 0 0 0 0 0 0 0 0 0 100 1 LC
Sisyphus goryi 0 0 0 0 22 0 22 33 0 22 0 18 LC
Sisyphus impressipennis 0 0 0 0 100 0 0 0 0 0 0 1 LC
Sisyphus splendidus 0 0 0 0 50 50 0 0 0 0 0 2 DD
Total number of species 12 27 53 15 111 129 120 99 83 42 60 216

CONSERVATION STATUS: DD = Data Deficient; LC = Least Concern; NT = Near Threatened; VU = Vulnerable; EN = Endangered; CR = Critically Endangered
SURICATA 6 (2020)

FREQUENCY: U = Uncommon; R = Rare; RR = Range Restricted; LLR = Localised in Large Range

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe ATEUCHINI (Figs 6A, 6D)
Coptorhina auspicata (NE6) 19.55 610 1 139 E Savanna / Kalahari 56 898 960
Coptorhina excavata (E4) 16.43 586 1 451 Endemic Highveld (Grassland) 8 115 840
SURICATA 6 (2020)

Coptorhina klugii (NE6) 19.53 751 646 E Savanna / Highveld (N Grassland) 73 681 450
Coptorhina nitidipennis (NE6) 20.25 722 837 E Savanna 39 4 106 325
Delopleurus darrenmanni (NE2) 22.30 609 907 W Savanna - N Botswana 3 22 265
Delopleurus gilleti (NE2) 21.02 678 791 W Savanna - N Namibia 4 1 773 210
Delopleurus pullus (NE7) 19.66 701 1 063 Savanna (E & W) 23 186 855
Frankenbergerius armatus armatus (E7) 17.99 778 376 Endemic E Coast (Forest) 14 43 660
Frankenbergerius armatus tuberculatus (E4) 17.25 839 1 216 Endemic Highveld (E Forest) 17 21 915
Frankenbergerius barratti (E4) 16.33 768 1 320 Endemic Highveld (Grassland) 8 16 485
Frankenbergerius forcipatus (E4) 15.22 662 1 220 Endemic Highveld (SE?Forest) 6 21 625
Frankenbergerius gomesi (E4) 18.54 805 943 Endemic Highveld (NE Forest) / E Savanna 13 18 695
Frankenbergerius nanus (E1) 15.05 436 339 Endemic SW Cape (Fynbos) 4 2 995
Frankenbergerius nitidus (E1) 18.15 140 1 Endemic SW Cape - Namaqualand (Succulent Karoo) 1 ?
Frankenbergerius opacus (E1) 16.90 435 228 Endemic SW Cape (Fynbos) 1 ?
Pedaria barrei (NE6) 21.83 560 496 E Savanna 7 1 460 055
Pedaria brancoi (NE7) 20.86 729 1 087 Savanna (E & W) 8 809 330
Pedaria cylindrica (NE6) 20.26 612 810 E Savanna 22 127 365
Pedaria picea (E10) 19.29 664 860 Endemic E Savanna 73 388 675
Pedaria segregis (NE6) 21.99 704 213 E Savanna 33 730 030
Sarophorus angolensis (NE2) 18.15 583 1 372 W Savanna - Arid NW Namibia 2 3 760
Sarophorus bidentatus (E1) 15.50 162 1 070 Endemic SW Cape - Namaqualand (Succulent Karoo) 1 ?
Sarophorus carinatus (E4) 16.78 875 1 281 Endemic Highveld (E Woodland) 2 755
Sarophorus costatus (NE6) 19.26 674 926 E Savanna / Highveld (N Grassland) 139 283 960
Sarophorus diabolus (E1) 16.90 350 334 Endemic SW Cape (Fynbos) 2 ?
Sarophorus frolovi (E4) 15.30 815 1 187 Endemic Highveld (SE Forest) 1 ?
Sarophorus latus (E10) 19.03 633 1 091 Endemic E Savanna 4 12 270
Sarophorus punctatus (E2) 17.95 458 0 Endemic S Cape (Forest) 1 ?
Sarophorus striatus (E2) 17.18 501 371 Endemic S Cape (Albany Thicket, Fynbos) 13 6740
Sarophorus tuberculatus (E2) 15.92 386 445 Endemic SW Cape / S Cape (Fynbos, Albany Thicket) 17 46 695
*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
747

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe CANTHONINI (Figs 6B, 6C, 6F, 6G)
Aliuscanthoniola similaris (E7) 19.70 961 58 Endemic E Coast (Forest) 1 ?
Aphengoecus clypeatus (E1) 16.90 435 228 Endemic SW Cape (Fynbos) 1 ?
Aphengoecus multiserratus (E1) 17.08 245 194 Endemic SW Cape - West Coast (Fynbos) 16 3 700
748

Bohepilissus nitidus (E1) 14.07 487 485 Endemic SW Cape (Forest) 6 25

Bohepilissus subtilis (E2) 16.21 492 422 Endemic S Cape (Forest) 29 38 100
Byrrhidium convexum (E1) 16.61 57 452 Endemic SW Cape - Richtersveld (Succulent Karoo) 7 163
Byrrhidium ovale (E1) 16.77 143 689 Endemic SW Cape - Namaqualand (Succulent Karoo) 6 2 370
Chalconotus convexus (NE7) 20.30 646 790 Savanna (E & W) 335 6 750 790
Circellium bacchus (E2) 17.24 431 151 Endemic S Cape (Fynbos, Albany Thicket) 24 11 950
Dicranocara deschodti (E5) 16.54 81 769 Endemic SW Arid - S Namibia (Nama Karoo) 5 190
Dicranocara inexpectata (E5) 16.90 99 867 Endemic SW Arid - S Namibia (Nama Karoo) 1 ?
Dicranocara tatasensis (E1) 17.70 53 372 Endemic SW Cape - Richtersveld (Succulent Karoo) 1 3.3
Dicranocara vandersmisseni (E5) 19.55 91 449 Endemic SW Arid - S Namibia (Nama Karoo) 2 300
Drogo stalsi (E5) 12.85 123 1 608 Endemic SW Arid - S Namibia (Nama Karoo) 1 ?
Dwesasilvisedis medinae (E7) 18.90 899 112 Endemic E Coast (Forest) 1 28.6
Endroedyolus paradoxus (E4) 15.55 1 111 1 170 Endemic Highveld (SE Forest) 2 80
Epirinus aeneus (E1) 17.55 231 652 Endemic SW Cape (Fynbos) / Nama Karoo 125 335 500
Epirinus aquilus (E2) 16.75 539 402 Endemic S Cape (Forest) 2 130
Epirinus asper (E4) 13.74 854 1 763 Endemic Highveld (NE Grassland) 18 40 500
Epirinus bentoi (E1) 17.33 226 91 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 12 2 700
Epirinus comosus (E1) 16.01 375 381 Endemic SW Cape / S Cape (Fynbos) 14 27 500
Epirinus convexus (E7) 18.78 869 225 Endemic E Coast (Forest) 6 2 300
Epirinus davisi (E7) 20.55 988 258 Endemic NE Coast - KZN (Forest) 2 81
Epirinus drakomontanus (E4) 7.78 877 2 731 Endemic Highveld (E Grassland) 3 80
Epirinus flagellatus (E1) 16.08 302 540 Endemic SW Cape / S Cape (Fynbos, Succulent Karoo) 110 258 000
Epirinus granulatus (E1) 17.25 216 130 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 11 3 400
Epirinus gratus (E4) 17.22 542 1 379 Endemic Highveld (NW Grassland) 23 74 300
Epirinus hilaris (E2) 15.28 451 609 Endemic S Cape (Fynbos) 17 29 550
Epirinus hluhluwensis (E7) 20.45 900 273 Endemic NE Coast - KZN (Forest) 1 1 580
Epirinus minimus (E2) 17.35 462 61 Endemic S Cape (Forest) 2 1 130
Epirinus montanus (E2) 12.45 331 1 442 Endemic S Cape (Fynbos) 2 1 750
Epirinus mucrodentatus (E4) 15.28 885 1 534 Endemic Highveld (NE) 3 170
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Epirinus ngomae (E4) 17.55 858 999 Endemic Highveld (E Forest) 1 765
Epirinus obtusus (E4) 15.60 494 1 039 Endemic Highveld (SE Grassland) / Albany Thicket 26 106 900
Epirinus pseudorugosus (E1) 15.80 302 63 Endemic SW Cape - West Coast (Fynbos) 7 230
Epirinus punctatus (E4) 13.20 954 1 820 Endemic Highveld (E Forest) 5 17 500
SURICATA 6 (2020)

Epirinus pygidialus (E4) 11.50 815 2 100 Endemic Highveld (E?Forest) 4 7 810
Epirinus relictus (E4) 11.84 701 1 761 Endemic Highveld (E Grassland) / S Cape (Fynbos) 11 50 250
Epirinus rugosus (E1) 13.20 273 1 453 Endemic SW Cape (Fynbos) 3 15
Epirinus scrobiculatus (E1) 17.33 197 177 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 25 15 275
Epirinus sebastiani (E4) 14.28 705 1 242 Endemic Highveld (SE Forest) 2 130
Epirinus silvestris (E2) 16.87 481 237 Endemic S Cape (Forest) 14 1 175
Epirinus striatus (E3) 15.22 306 1 352 Endemic Upper Karoo 53 98 000
Epirinus sulcipennis (E2) 10.90 379 1 546 Endemic S Cape (Fynbos) 1 ?
Epirinus validus (E4) 12.95 793 1 772 Endemic Highveld (E Grassland) 26 52 550
Gyronotus carinatus (E7) 21.05 964 115 Endemic NE Coast - KZN (Forest) 5 2 900
Gyronotus glabrosus (E4) 16.20 869 1 363 Endemic Highveld (NE Grassland) 6 1 270
Gyronotus perissinottoi (E7) 19.35 966 161 Endemic E Coast (Grassland) 1 ?
Gyronotus pumilus (E7) 19.51 914 119 Endemic E Coast (Forest) 18 6 350
Gyronotus schuelei (E4) 19.10 922 870 Endemic Highveld (NE Grassland) 2 83
Hammondantus psammophilus (E5) 16.11 76 845 Endemic SW Arid - Namib Desert 6 13 600
Namakwanus irishi (E5) 18.30 353 1 815 Endemic SW Arid - Arid Namibian Savanna 1 ?
Namakwanus scholtzi (E5) 16.65 18 400 Endemic SW Arid - Namib Desert 1 ?
Namaphilus ameibensis (E5) 17.50 210 1 197 Endemic SW Arid - Arid Namibian Savanna 1 ?
Namaphilus davisi (E5) 21.00 176 1 164 Endemic SW Arid - Arid Namibian Savanna 1 25
Namaphilus endroedyi (E5) 15.72 155 1 474 Endemic SW Arid - Arid Namibian Savanna 3 280
Nebulasilvius insularis (E4) 16.85 847 974 Endemic Highveld (SE Forest) 1 ?
Nebulasilvius johani (E4) 15.35 815 1 179 Endemic Highveld (SE Forest) 2 1.7
Odontoloma apiculum (E2) 18.28 411 227 Endemic S Cape (Forest) 2 905
Odontoloma dentinum (E1) 16.95 316 179 Endemic SW Cape / S Cape (Fynbos, Succulent Karoo) 21 52 460
Odontoloma disalatum (E1) 16.89 266 289 Endemic SW Cape (Fynbos) 11 43 165
Odontoloma doubei (E4) 14.80 833 1 206 Endemic Highveld (E Grassland) 6 13 375
Odontoloma endroedyi (E2) 15.04 313 913 Endemic S Cape (Fynbos, Succulent Karoo) 11 45 810
Odontoloma louwi (E4) 18.57 339 1 305 Endemic Highveld (Grassland) / W Savanna - N Namibia 11 131 940

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
749

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe CANTHONINI (Figs 6B, 6C, 6F, 6G) (continued)
Odontoloma obscurum (E4) 11.39 688 1 838 Endemic Highveld (SE Grassland) 5 8 020
Odontoloma pauxillum (NE6) 20.13 629 924 E Savanna 6 1 021 080
Odontoloma peckorum (E4) 14.40 816 1 544 Endemic Highveld (E Grassland) 12 19 885
750

Odontoloma planatum (E4) 16.31 900 1 096 Endemic Highveld (E Grassland) 7 14 615
Odontoloma pusillum (E1) 16.60 273 94 Endemic SW Cape - West Coast (Fynbos) 5 1 685
Odontoloma pygidiale (E1) 16.81 243 334 Endemic SW Cape (Fynbos, Succulent Karoo) 47 71 165
Odontoloma quadridens (NE6) 20.30 486 1 040 E Savanna 3 15 205
Odontoloma sculpturatum (E4) 16.29 597 861 Endemic Highveld (E Grassland) / S Cape 7 64 415
Odontoloma spinicaudum (E1) 18.15 142 38 Endemic SW Cape - Namaqualand (Succulent Karoo) 1 ?
Outenikwanus tomentosus (E2) 13.75 404 999 Endemic S Cape (Forest) 1 ?
Parvuhowdenius harrisoni (E4) 14.65 855 1 364 Endemic Highveld (SE Forest) 1 16.9
Peckolus alpinus (E4) 15.98 891 1 410 Endemic Highveld (NE Forest) 2 15
Peckolus parvus (E4) 9.15 810 2 497 Endemic Highveld (E Forest) 1 1.4
Peckolus poenskopius (E4) 17.98 1 005 1 085 Endemic Highveld (NE Forest) 3 85
Pycnopanelus krikkeni (E9) 18.84 191 987 Endemic SW Arid (Nama Karoo, Arid Namibian Savanna) 110 264 500
Silvaphilus oubosiensis (E1) 16.30 367 389 Endemic SW Cape (Forest) 1 6
Versicorpus erongoensis (E5) 17.50 210 1 197 Endemic SW Arid - Arid Namibian Savanna 1 ?
Versicorpus streyi (E5) 15.85 179 1 621 Endemic SW Arid - Arid Namibian Savanna 1 ?
Tribe COPRINI (Figs 6D, 6E, 6G)
Catharsius aegeus (NE6) 20.72 631 792 E Savanna 48 812 345
Catharsius calaharicus (E8) 19.86 329 1 062 Endemic Kalahari 152 639 850
Catharsius harpagus (NE3) 22.23 801 28 NE Coast - KZN 12 8 250
Catharsius heros (NE7) 21.26 538 895 Savanna (E & W) / Kalahari 32 3 687 200
Catharsius laticeps (NE6) 22.83 513 308 E Savanna 4 15 750
Catharsius longiceps (E4) 16.82 657 1 470 Endemic Highveld (Grassland) 10 6 130
Catharsius marcellus (E4) 15.16 802 1 533 Endemic Highveld (Grassland) 37 126 300
Catharsius melancholicus (NE4) 19.98 338 1 050 Kalahari 45 970 820
Catharsius pandion (NE3) 21.91 853 43 NE Coast - KZN 52 18 650
Catharsius philus (NE6) 20.51 627 783 E Savanna 94 774 500
Catharsius platycerus (NE6) 23.16 704 568 E Savanna 8 3 012 810
Catharsius sesostris (NE6) 19.31 757 993 E Savanna / Highveld (Grassland) 96 5 293 770
Catharsius tricornutus (NE7) 18.92 701 873 Savanna (E & W), Highveld (Grassland), S Cape 205 1 064 120
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Catharsius ulysses (E4) 18.87 454 1 307 Endemic Highveld (Grassland) / W Savanna - N Namibia 70 367 200
Catharsius vitulus (E4) 15.30 404 1 385 Endemic Highveld (Grassland) / Upper Karoo 40 92 235
Copris amyntor (NE6) 20.73 631 739 E Savanna 255 1 016 860
Copris anceus (E1) 16.70 270 77 Endemic SW Cape - W Coast (Fynbos) 35 7 625
SURICATA 6 (2020)

Copris antares (E4) 16.47 628 1 136 Endemic Highveld (Grassland) 61 269 985
Copris bootes (NE6) 22.39 740 497 E Savanna 19 959 800
Copris caelatus (E4) 14.85 792 1 480 Endemic Highveld (E Grassland)) 40 91 950
Copris cambeforti (E10) 20.00 578 952 Endemic E Savanna 2 540
Copris capensis (E1) 17.14 205 175 Endemic SW Cape (Fynbos, Succulent Karoo) 43 45 395
Copris cassius (E8) 19.22 372 1 161 ?Endemic Kalahari (SW) 92 547 290
Copris cornifrons (E8) 19.37 312 1 111 Endemic Kalahari (SW) 38 286 300
Copris corniger (E4) 15.50 718 1 356 Endemic Highveld (Grassland) 90 318 935
Copris crassus (E4) 14.98 721 1 437 Endemic Highveld (E Grassland)) 8 41 240
Copris denticulatus (NE6) 19.64 690 927 E Savanna 182 940 275
Copris elphenor (NE7) 20.02 581 972 Savanna (E & W) 420 3 968 365
Copris evanidus (NE6) 20.56 564 802 E Savanna 217 2 357 905
Copris fidius (NE1) 17.63 755 650 Highveld (E Forest) / E Coast / S Cape (Forest) 110 116 200
Copris gracilis (E6) 19.29 340 1 394 Endemic W Savanna - N Namibia 54 168 350
Copris inhalatus perturbator (NE4) 19.91 492 1 147 Kalahari 14 614 150
Copris inhalatus sanctaeluciae (NE3) 22.02 831 34 NE Coast - KZN 27 8 520
Copris jacchoides (E4) 15.27 724 1 430 Endemic Highveld (Grassland) 52 258 970
Copris jacchus (E2) 17.51 462 244 Endemic S Cape (Albany Thicket, Fynbos) 6 9 990
Copris laioides (E6) 20.59 446 1 429 Endemic W Savanna - N Namibia 20 49 485
Copris macer (NE6) 17.89 846 1 153 E Savanna / Highveld (Grassland) 39 65 380
Copris mesacanthus mesacanthus (NE6) 20.90 791 894 E Savanna 20 378 400
Copris mesacanthus transvaalensis (NE6) 19.43 670 944 E Savanna 133 223 300
Copris obesus (NE1) 17.68 752 1 237 Highveld (NE Grassland) 171 1 252 590
Copris orion caffer (E7) 19.38 806 189 Endemic E Coast / Albany Thicket 23 31 800
Copris puncticollis (NE7) 22.26 778 245 Savanna (E & W) / NE Coast - KZN 43 211 185
Copris ritsemae (E4) 17.27 724 1 356 Endemic Highveld (N Grassland) 10 18 840
Copris sexdentatus (E2) 17.62 427 88 Endemic S Cape (Fynbos) 3 3 945
Copris sphaeropterus (E2) 16.40 422 152 Endemic S Cape (Fynbos) 1 ?

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
751

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe COPRINI (Figs 6D, 6E, 6G)
Copris subsidens (E6) 19.05 340 1 366 Endemic W Savanna - N Namibia 58 105 070
Copris urus (NE3) 21.91 847 42 NE Coast KZN 25 8 690
Copris victorini (E2) 16.06 425 437 Endemic S Cape (Fynbos) 4 2 100
752

Copris vilhenai (NE6) 20.81 630 867 E Savanna 43 677 310

Copris vrydaghi (E4) 16.90 758 1 070 Endemic Highveld (E Grassland) 4 5 085
Heliocopris andersoni (NE7) 19.61 567 1 057 Savanna (E & W) 103 3 715 845
Heliocopris atropos (NE4) 20.01 493 1 113 Kalahari / East Savanna 44 1 602 870
Heliocopris faunus (NE7) 18.36 376 1 177 Savanna (E & W) / Upper Karoo 69 391 450
Heliocopris hamadryas (NE6) 18.60 678 1 068 E Savanna / Highveld (Grassland) 155 3 416 220
Heliocopris japetus (NE4) 20.46 569 1 020 Kalahari / E Savanna 137 3 260 895
Heliocopris neptunus (NE6) 19.92 653 824 E Savanna 83 328 720
Heliocopris pirmal (NE6) 18.70 702 1 085 E Savanna / Highveld (NE Grassland) 64 236 025
Litocopris muticus (NE6) 22.17 912 257 E Savanna 11 793 435
Litocopris simplex (E7) 16.74 684 695 Endemic E Coast / Highveld (E Grassland) 45 85 760
Macroderes amplior (E1) 17.58 134 388 Endemic SW Cape - Namaqualand (Succulent Karoo) 3 1 170
Macroderes arrowi (E1) 18.73 161 218 Endemic SW Cape - Namaqualand (Succulent Karoo) 2 52.5
Macroderes bias (E2) 15.54 432 839 Endemic S Cape (Albany Thicket, Fynbos) 9 40 815
Macroderes cornutus (E1) 17.41 115 73 Endemic SW Cape - Namaqualand (Succulent Karoo) 4 2 605
Macroderes endroedyi (E1) 17.69 204 162 Endemic SW Cape - W Coast (Fynbos, Succulent Karoo) 9 2 775
Macroderes fornicatus (E1) 15.23 454 235 Endemic SW Cape (Fynbos) 2 535
Macroderes foveatus (E1) 16.55 255 20 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 2 105
Macroderes greeni (E1) 16.07 324 47 Endemic SW Cape (Fynbos) 7 1 535
Macroderes minutus (E1) 18.42 155 257 Endemic SW Cape - Namaqualand (Succ. Karoo, Fynbos) 6 5 045
Macroderes mutilans (E1) 16.96 129 871 Endemic SW Cape - Namaqualand (Succulent Karoo) 5 1 501
Macroderes namakwanus (E1) 17.94 126 148 Endemic SW Cape - Namaqualand (Succulent Karoo) 5 53
Macroderes nitidus (E1) 17.03 169 791 Endemic SW Cape - Namaqualand (Succulent Karoo) 2 80
Macroderes politulus (E1) 16.47 356 320 Endemic SW Cape (Fynbos) 8 7 940
Macroderes undulatus (E1) 16.65 388 277 Endemic SW Cape (Fynbos) 4 2 300
Metacatharsius anderseni (E8) 19.57 266 1 077 Endemic Kalahari (SW) 52 255 955
Metacatharsius dentinum (NE4) 19.51 294 1 034 Kalahari / Nama Karoo 114 1 080 805
Metacatharsius exiguiformis (E8) 19.30 248 1 041 Endemic Kalahari (SW) / Nama Karoo / Arid W Savanna 168 471 130
Metacatharsius exiguus (NE4) 20.67 557 827 Kalahari / East Savanna / NE Coast - KZN 40 680 625
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Metacatharsius ferreirae (NE4) 21.95 459 951 Kalahari / East Savanna 8 923 845
Metacatharsius freyi (E8) 20.78 412 994 Endemic Kalahari / East Savanna 10 257 510
Metacatharsius latifrons (E8) 18.45 279 975 Endemic Kalahari (SW) / SW Cape - West Coast (SKaroo) 91 322 620
Metacatharsius marani (E9) 18.45 223 1 083 Endemic SW Arid (Nama Karoo, Arid W Savanna) 239 312 390
SURICATA 6 (2020)

Metacatharsius opacus (NE7) 21.33 560 771 Savanna (E & W) 189 3 129 685
M. pumilioniformis / pumilionis (E8) 19.69 398 1 000 Endemic Kalahari / E Savanna 43 688 900
Metacatharsius troglodytes (NE7) 20.62 538 934 Savanna (E & W) 173 1 228 465
Metacatharsius zuluanus (NE3) 22.49 708 132 NE Coast - KZN / E Savanna 19 30 915
Xinidium dentilabris (E4) 15.47 838 1 394 Endemic Highveld (E Grassland) 103 107 815
Xinidium dewitzi (E4) 15.05 813 1 287 Endemic Highveld (E Forest) 4 5 990
Xinidium howdeni (E4) 10.90 768 1 707 Endemic Highveld (E Forest) 3 20
Tribe GYMNOPLEURINI (Fig. 6H)
Allogymnopleurus consocius (E7) 20.19 710 540 Endemic NE Coast - KZN / Highveld (N Grassland) 11 35 250
Allogymnopleurus splendidus (NE7) 19.49 477 956 Savanna (E & W) / Upper Karoo 413 1 523 140
Garreta australugens (NE6) 22.64 659 405 E Savanna 7 88 890
Garreta caffer (NE3) 22.04 849 12 NE Coast - KZN 26 8 394
Garreta laetus olivaceus (NE6) 21.23 913 754 E Savanna (Forest) 18 2 848 875
Garreta nitens (NE2) 20.36 545 1 282 W Savanna - N Namibia 26 4 761 275
Garreta unicolor (NE6) 17.32 797 1 111 E Savanna / Highveld (N Grassland) 98 209 670
Garreta wahlbergi (NE6) 20.33 643 857 E Savanna 209 737 635
Gymnopleurus aenescens (NE7) 19.75 354 978 Savanna (E & W) / Kalahari / Upper Karoo 157 1 407 890
Gymnopleurus andreaei (NE5) 18.60 188 1 003 SW Arid (Nama Karoo, Arid W Savanna) 143 263 705
Gymnopleurus asperrimus (E9) 18.92 206 1 020 Endemic SW Arid (Nama Karoo) / Kalahari (SW) 187 223 500
Gymnopleurus humanus (NE5) 17.93 231 1 106 SW Arid (Nama Karoo, Arid W Savanna) 380 587 750
Gymnopleurus humeralis (NE6) 20.94 601 735 E Savanna 115 534 260
Gymnopleurus ignitus (NE4) 22.80 652 706 Kalahari (NE) 24 659 605
Gymnopleurus leei (E4) 15.59 514 1 335 Endemic Highveld (Grassland) / S Cape 74 256 000
Gymnopleurus pumilus (NE7) 20.77 550 850 Savanna (E & W) 141 451 400
Gymnopleurus reichei (NE6) 22.89 521 256 E Savanna 4 253 700
Gymnopleurus thelwalli (NE6) 21.52 661 617 E Savanna 7 555 105
Gymnopleurus virens (NE6) 18.53 739 1 089 E Savanna 114 267 770

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
753

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe ONITICELLINI (Fig. 6I)
Afrodrepanus impressicollis (E7) 19.50 875 595 Endemic NE Coast - KZN / E Savanna (NE Forest) 32 27 860
Cyptochirus ambiguus (NE1) 16.94 775 1 046 Highveld (Grassland) / E Savanna 112 323 350
Drepanocerus kirbyi (NE6) 18.45 733 883 E Savanna / Highveld (Grassland) 112 3 526 325
754

Drepanocerus patrizii (NE6) 18.01 536 1 129 E Savanna / Highveld (Grassland) 41 503 040
Eodrepanus bechynei (NE3) 20.24 1 021 421 NE Coast - KZN / East Savanna 6 2 080 435
Eodrepanus fastiditus (NE6) 18.47 763 941 E Savanna / Highveld (Grassland) 56 1 517 320
Eodrepanus parallelus (NE3) 20.98 755 754 NE Coast - KZN / E Savanna 8 797 475
Epidrepanus caelatus (NE1) 17.23 786 1 199 Highveld (Grassland) 24 160 840
Euoniticellus africanus (E4) 15.75 518 1 218 Endemic Highveld (Grassland) 215 677 390
Euoniticellus intermedius (NE7) 18.70 507 967 Savanna (E & W), SW Arid, SW Cape, S Cape 580 9 060 360
Euoniticellus kawanus (NE7) 20.64 444 968 Savanna (E & W) 8 2 691 645
Euoniticellus triangulatus (NE1) 16.91 673 887 Highveld (Grassland) / E Savanna / S Cape 178 2 234 985
Euoniticellus zumpti (NE3) 21.37 778 310 NE Coast - KZN / E Savanna 15 273 965
Ixodina abyssinica (NE3) 21.34 888 229 NE Coast - KZN 6 347 255
Latodrepanus laticollis (NE6) 20.40 701 832 E Savanna 80 4 469 700
Liatongus militaris (NE6) 18.42 720 949 E Savanna / Highveld (Grassland) 315 3 632 435
Oniticellus egregius (NE7) 20.67 597 748 Savanna (E & W) 38 2 407 180
Oniticellus formosus (NE6) 20.39 608 752 E Savanna 77 4 827 535
Oniticellus pictus (NE1) 16.49 661 834 Highveld (E Grassland) / S Cape 35 131 510
Oniticellus planatus (NE6) 18.84 731 791 E Savanna / Highveld (NE Grassland) / S Cape 186 4 837 160
Paraixodina freyi (NE3) 21.18 795 247 NE Coast - KZN / E Savanna 9 450 215
Paraixodina saegeri (NE6) 21.73 668 480 E Savanna 12 1 073 235
Tibiodrepanus sulcicollis (NE1) 16.55 725 893 Highveld (SE Grassland) / E Coast 38 5 683 200
Tiniocellus eurypygus eurypygus (E10) 18.7 638 1 212 Endemic E Savanna 32 39 045
T. eurypygus transdrakensbergensis (E10) 21.13 700 467 Endemic E Savanna 35 84 580
Tiniocellus spinipes (NE6) 22.60 636 470 E Savanna 27 2 951 525
Tragiscus dimidiatus (NE7) 22.39 602 471 Savanna (E & W) 24 2 192 230
Tribe ONITINI (Fig. 6J)
Anonychonitis freyi (E7) 20.80 878 199 Endemic NE Coast - KZN / E Savanna 5 960
Cheironitis audens (E9) 17.93 241 1 100 Endemic SW Arid (Nama Karoo, Arid W Savanna) 105 208 930
Cheironitis hoplosternus (E4) 17.67 353 1 187 Endemic Highveld (Grassland) / Upper Karoo / N Namibia 138 651 685
Cheironitis imitator (NE6) 22.29 486 541 E Savanna 16 669 950
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Cheironitis indicus (E9) 18.51 196 1 028 Endemic SW Arid (Nama Karoo, Arid W Savanna) 61 113 785
Cheironitis scabrosus (E9) 17.59 244 1 017 Endemic SW Arid (Nama Karooa) / SW Cape 242 478 375
Gilletellus porculus (E6) 22.2 391 1 156 Endemic W Savanna - N Namibia 1 ?
Heteronitis castelnaui (NE7) 22.03 605 565 Savanna (E & W) 74 3 997 830
SURICATA 6 (2020)

Kolbeellus ateuchoides (E3) 11.85 429 1 890 Endemic Upper Karoo 2 1 395
Megalonitis bohemani (NE7) 22.39 528 928 Savanna (E & W) 7 317 740
Neonitis nigritiae (NE6) 18.68 748 1 355 E Savanna 4 123 785
Onitis aeruginosus (NE6) 22.96 586 701 E Savanna 10 134 325
Onitis alexis (NE7) 19.27 600 981 Savanna (E & W) / Highveld (Grassland) 574 9 990 325
Onitis aygulus (E3) 16.88 356 971 Endemic Upper Karoo / S Cape 157 756 700
Onitis bilobatus (E6) 17.00 367 1 264 Endemic W Savanna - N Namibia 1 ?
Onitis caffer (E4) 17.01 585 990 Endemic Highveld (Grassland) / E Savanna / S Cape 275 576 190
Onitis confusus (E3) 16.55 298 603 Endemic SW Cape / Upper Karoo 34 529 675
Onitis cribratus (E4)*** 14.63 762 1 430 Endemic Highveld (E Grassland) 3 265
Onitis curvipes (E4) 16.69 458 914 Endemic Highveld (Grassland) / S Cape 14 150 365
Onitis deceptor (NE6) 19.86 625 961 E Savanna / Kalahari (SW) 62 1 264 130
Onitis fabricii (NE6) 22.05 713 999 E Savanna 22 2 595 525
Onitis fulgidus (NE6) 20.44 633 832 E Savanna 112 366 000
Onitis inversidens (NE6) 22.35 608 545 E Savanna 24 1 704 640
Onitis licitus (E4) 13.7 528 1 619 Endemic Highveld (S Grassland) 2 105
Onitis longitibialis (E3) 17.55 353 1 222 Endemic Upper Karoo 1 ?
Onitis mendax (NE7) 21.48 628 657 Savanna (E & W) 39 2 759 790
Onitis minutus (E1) 16.89 386 393 Endemic SW Cape / S Cape (Fynbos) 11 70 125
Onitis mniszechi (E6) 18.66 265 1 236 Endemic W Savanna - N Namibia 5 67 800
Onitis obenbergeri (NE6) 22.27 431 533 E Savanna 26 59 050
Onitis obscurus (NE2) 20.40 478 1 299 W Savanna - N Namibia 21 157 030
Onitis orthopus (NE2) 21.38 618 1 056 W Savanna - N Botswana 7 281 070
Onitis paraconfusus (E10) 22.35 578 377 Endemic E Savanna 2 1 950
Onitis parainflaticollis (E10) 18.25 631 1 115 Endemic E Savanna / Kalahari 12 116 085
Onitis pecuarius (E4) 16.68 730 876 Endemic Highveld (E Grassland) / E Savanna / S Cape 116 198 180
Onitis perpunctatus (E4) 16.47 741 1 268 Endemic Highveld (Grassland) / E Savanna 67 96 210
Onitis picticollis (NE7) 19.71 657 871 Savanna (E & W) 65 327 800

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
755

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe ONITINI (Fig. 6J) (continued)
Onitis pseudosetosus (NE6) 19.38 639 1 063 E Savanna 18 191 355
Onitis robustus (NE7) 22.19 697 691 Savanna (E & W) 15 3 525 050
Onitis setosus (NE2) 19.19 417 1 131 W Savanna - N Namibia 8 119 160
756

Onitis tortuosus (NE1) 16.98 751 1 248 Highveld (Grassland) / E Savanna 204 348 320
Onitis uncinatus (NE7) 19.79 599 1 015 Savanna (E & W) 313 4 665 140
Onitis viridulus (NE6) 20.20 715 803 E Savanna 206 4 084 025
Onitis westermanni (NE6) 20.48 655 954 E Savanna 48 944 160
Platyonitis bicuariensis (NE2) 22.23 609 1 092 W Savanna - N Namibia 3 88 760
Tropidonitis paradoxus (NE3) 22.45 720 36 NE Coast - KZN 6 2 455
Tribe ONTHOPHAGINI (Figs 6K, 6L, 6M)
Caccobiomorphus megaponerae (NE6) 22.06 600 703 E Savanna 7 299 670
Caccobius castaneus (NE7) 21.18 494 863 Savanna (E & W) 12 2 314 560
Caccobius cavatus (NE6) 21.36 693 539 E Savanna / NE Coast - KZN 26 861 245
Caccobius ferrugineus (NE7) 20.37 510 951 Savanna (E & W) / Kalahari 137 4 823 540
Caccobius histerinus (NE6) 21.25 696 473 E Savanna 38 1 133 180
Caccobius nigritulus (NE7) 21.11 598 695 Savanna (E & W) / Kalahari 102 2 013 940
Caccobius obtusus (NE1) 16.62 796 1 047 Highveld (E Grassland) / S Cape 75 781 120
Cleptocaccobus convexifrons (NE6) 20.95 622 806 E Savanna 12 3 973 335
Cleptocaccobus postlutatus (NE7) 18.90 724 885 Savanna (E & W) 51 1 627 025
Cleptocaccobius viridicollis (NE7) 19.06 448 971 Savanna (E & W) / Kalahari 134 1 295 810
Digitonthophagus gazella (NE6) 19.52 626 804 E Savanna 404 3 711 400
Digitonthophagus namaquensis (NE5) 18.32 215 984 SW Arid (Nama Karoo, Arid W Savanna) 69 795 870
Digitonthophagus viridicollis (NE2) 19.56 390 1 222 W Savanna - N Namibia 29 419 910
Euonthophagus carbonarius (NE6) 21.25 619 627 E Savanna 115 946 605
Euonthophagus flavimargo (E8) 19.30 264 1 080 Endemic Kalahari (SW) 114 211 965
Euonthophagus jeanneli (NE6) 22.37 582 371 E Savanna 19 409 200
Euonthophagus vicarius (E3) 15.80 282 1 236 Endemic Upper Karoo 64 97 405
Hamonthophagus acutus (E9) 19.08 219 1 005 Endemic SW Arid (Nama Karoo) / Kalahari (SW) 50 390 575
Hamonthophagus depressus (NE7) 20.11 676 701 Savanna (E & W) 128 669 355
Hamonthophagus fallax (NE2) 22.50 666 884 W Savanna - N Botswana 7 327 565
Haroldius convexus (E10) 17.20 699 1 452 Endemic E Savanna 1 ?
Heteroclitopus remipes (NE6) 18.99 756 809 E Savanna / E Coast 6 42 635
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Hyalonthophagus alcyon (NE7) 21.91 635 938 Savanna (E & W) 12 474 810
Hyalonthophagus alcyonides (NE6) 20.89 694 576 E Savanna 62 232 385
Kurtops quadraticeps (NE4) 19.81 321 1 079 Kalahari 115 799 195
Kurtops signatus (NE4) 19.81 362 965 Kalahari / E Savanna / Upper Karoo 293 1 133 835
SURICATA 6 (2020)

Milichus apicalis (NE7) 21.55 657 509 Savanna (E & W) 32 7 382 875
Mimonthophagus ambiguus (NE3) 22.06 857 34 E Coast 31 298 230
Mimonthophagus anomalus (NE6) 22.68 647 730 E Savanna 22 658 380
Mimonthophagus limbibasis (NE6) 21.55 540 1 044 E Savanna 3 2 790
Onthophagus absyrtus (E4) 15.53 759 1 556 Endemic Highveld (Grassland) 11 65 350
Onthophagus aequepubens (NE7) 20.50 570 780 Savanna (E & W) 25 245 690
Onthophagus aeruginosus (NE7) 19.43 664 936 Savanna (E & W) / Highveld (Grassland) 232 4 968 210
Onthophagus albipennis (E3) 16.99 245 1 158 Endemic Upper Karoo 113 126 235
Onthophagus albipodex (NE6) 19.63 686 988 E Savanna 37 324 950
Onthophagus apiciosus (NE6) 19.15 709 1 038 E Savanna 33 2 057 540
Onthophagus asperulus (E4) 16.01 779 1 241 Endemic Highveld (Grassland) 139 170 285
Onthophagus axillaris (NE2) 18.77 415 1 249 W Savanna - N Namibia 23 393 675
Onthophagus bayeri (E6) 17.58 240 1 520 Endemic W Savanna - N Namibia 8 10 885
Onthophagus beiranus (NE3) 22.32 887 154 NE Coast - KZN / E Savanna 47 98 720
Onthophagus bicavifrons (NE7) 19.57 633 928 Savanna (E & W) 149 4 290 470
Onthophagus binodis (E4) 15.85 669 1 092 Endemic Highveld (Grassland) / S Cape 160 274 485
Onthophagus bovinus (NE7) 19.77 416 1 093 Savanna (E & W) / Upper Karoo / Kalahari (SW) 32 642 681
Onthophagus cameloides (E1) 16.78 237 506 Endemic SW & S Cape (Fynbos, S Karoo) / Upper Karoo 97 241 800
Onthophagus cinctipennis (NE1) 17.62 761 1 240 Highveld (Grassland) / E Savanna 35 217 320
Onthophagus cineraceus (E1) 14.43 473 571 Endemic SW Cape (Fynbos) 2 180
Onthophagus convexus (NE4) 19.25 279 1 015 Kalahari / Upper Karoo 183 988 530
Onthophagus corniculiger (NE6) 21.68 629 450 E Savanna 14 145 035
Onthophagus cretus (E7) 18.03 876 586 Endemic E Coast (Forest) 21 44 150
Onthophagus cribripennis (NE6) 17.46 800 1 122 E Savanna / Highveld (E Grassland) 146 206 820
Onthophagus croesulus (E10) 18.43 680 1 113 Endemic E Savanna / Highveld (E Grassland) 13 128 590
Onthophagus cupricollis (E10) 17.59 762 1 228 Endemic E Savanna 11 26 645
Onthophagus cyaneoniger (E4) 16.07 590 1 411 Endemic Highveld (Grassland) 42 181 630
Onthophagus deterrens (E4) 16.62 793 1 275 Endemic Highveld (Grassland) / E Savanna 55 293 940

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
757

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe ONTHOPHAGINI (Figs 6K, 6L, 6M) (continued)
Onthophagus discretus (NE6) 18.96 648 1 166 E Savanna 13 42 255
Onthophagus ebenicolor (NE7) 20.77 619 716 Savanna (E & W) 69 1 691 195
Onthophagus ebenus (NE6) 19.25 704 953 E Savanna / Highveld (N Grassland) 123 847 490
758

Onthophagus fimetarius (NE7) 18.79 683 1 069 Savanna (E & W) / Highveld (N Grassland) 199 5 546 895
Onthophagus flavolimbatus (NE7) 21.32 627 620 Savanna (E & W) 72 860 785
Onthophagus fritschi (E3) 15.38 413 1 333 Endemic Upper Karoo 80 174 610
Onthophagus fugitivus (E10) 19.18 607 1 042 Endemic E Savanna 18 118 900
Onthophagus giraffa (E1) 15.78 408 480 Endemic SW Cape / S Cape (Fynbos) 52 59 495
Onthophagus giuseppecarpanetoi (NE3) 22.26 754 36 NE Coast - KZN 27 8 945
Onthophagus gonopygus (NE2) 16.99 185 881 W Savanna - Arid W Namibia 17 102 535
Onthophagus granulifer (NE4) 19.65 299 1 060 Kalahari 127 1 080 205
Onthophagus graphicus (NE7) 19.96 486 935 Savanna (E & W) 9 1 088 955
Onthophagus herus (NE6) 23.32 722 439 E Savanna 8 1 168 730
Onthophagus hyaena (E4) 15.18 614 1 250 Endemic Highveld (S Grassland) / S Cape 38 38 760
Onthophagus immundus (E1) 16.02 340 521 Endemic SW Cape / S Cape (Fynbos) 13 26 615
Onthophagus interstitialis (E4) 17.85 653 1 042 Endemic Highveld (N Grassland) / E Savanna 95 251 810
Onthophagus juvencus (NE3) 22.54 865 67 NE Coast - KZN / E Savanna 25 2 971 515
Onthophagus kochi (E6) 20.41 442 949 Endemic W Savanna - N Namibia / N Botswana 7 42 635
Onthophagus lacustris (NE3) 22.33 846 245 NE Coast - KZN / E Savanna 38 874 260
Onthophagus lamelliger (NE6) 21.68 593 591 E Savanna 107 1 440 975
Onthophagus lamnifer (E4) 15.90 691 1 429 Endemic Highveld (NE Grassland) 38 96 245
Onthophagus leroyi (NE6) 21.77 612 382 E Savanna 27 228 595
Onthophagus leucopygus (NE4) 18.78 381 1 224 Kalahari 35 587 600
Onthophagus lugubris (E4) 14.90 719 1 429 Endemic Highveld (Grassland) 47 180 110
Onthophagus minutus (E1) 16.61 303 118 Endemic SW Cape / S Cape (Fynbos) 39 44 485
Onthophagus monodon (E4) 16.29 753 1 416 Endemic Highveld (Grassland) 16 100 835
Onthophagus naso (E7) 18.48 833 578 Endemic Highveld (Grassland) / E Savanna / E Coast 18 70 340
Onthophagus obtusicornis (NE7) 19.03 634 1 014 Savanna (E & W) / Highveld (Grassland) 179 418 675
Onthophagus obtutus (E4) 14.99 677 1 579 Endemic Highveld (Grassland) 33 89 365
Onthophagus pallidipennis (NE6) 20.11 502 947 E Savanna / Kalahari 97 1 056 735
Onthophagus parumnotatus (NE6) 17.48 812 1 213 E Savanna / Highveld (Grassland) 103 2 391 165
Onthophagus pauxillus (NE6) 19.34 657 949 E Savanna 67 316 415
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Onthophagus pellax (E10) 19.06 591 1 151 Endemic E Savanna 24 40 975
Onthophagius peringueyi (E3) 17.00 247 1 157 Endemic Upper Karoo 136 219 865
Onthophagus pilosus (E4) 16.16 757 1 353 Endemic Highveld (Grassland) / E Savanna 51 156 440
Onthophagus plebejus (NE7) 22.63 619 654 Savanna (E & W) 32 1 949 910
SURICATA 6 (2020)

Onthophagus probus (NE5) 18.76 205 1 010 SW Arid (Nama Karoo, Arid W Savanna) 218 830 525
Onthophagus producticollis (NE6) 17.53 719 1 283 E Savanna 18 85 445
Onthophagus pugionatus (NE6) 19.31 714 916 E Savanna / Highveld (Grassland) 129 4 391 285
Onthophagus pullus (NE7) 20.09 636 852 Savanna (E & W) 46 2 692 290
Onthophagus quadrimaculatus (NE7) 20.33 618 827 Savanna (E & W) 34 1 875 455
Onthophagus quadrinodosus (NE6) 19.54 676 955 E Savanna 52 2 016 665
Onthophagus rasipennis (NE6) 20.02 655 811 E Savanna 73 230 805
Onthophagus scapularis (E2) 17.55 493 233 Endemic S Cape (Albany Thicket) 2 1 725
Onthophagus semiflavus (NE5) 19.04 210 966 SW Arid (Kalahari (SW), Nama Karoo) 109 226 630
Onthophagus suffusus (NE7) 21.84 586 719 Savanna (E & W) 31 962 255
Onthophagus tricorniger (E9) 19.18 333 1 430 Endemic SW Arid (Arid W Savanna) 6 65 590
Onthophagus trinodosus (E10) 19.24 635 1 020 Endemic E Savanna 14 32 470
Onthophagus ursinus (NE3) 22.22 861 39 NE Coast - KZN 17 38 715
Onthophagus variolosus (E4) 15.68 519 820 Endemic Highveld (S Grassland) / S Cape 16 58 525
Onthophagus venustulus (NE4) 19.57 329 1 084 Kalahari 184 1 414 010
Onthophagus verticalis (NE6) 20.20 520 1 045 E Savanna / Kalahari 65 1 288 690
Onthophagus vigens (E4) 15.44 834 1 367 Endemic Highveld (E Grassland) 44 61 695
Onthophagus vinctus (NE7) 20.16 663 797 Savanna (E & W) 305 8 771 775
Onthophagus virescens (NE6) 22.61 651 572 E Savanna 14 137 725
Onthophagus vylderi (E8) 20.81 588 1 050 Endemic Kalahari 8 241 110
Phalops ardea (NE7) 21.53 618 662 Savanna (E & W) 97 1 179 470
Phalops boschas (NE6) 21.10 630 724 E Savanna 130 2 069 530
Phalops bubalus (E4) 15.59 527 1 256 Endemic Highveld (Grassland) / Upper Karoo 108 355 495
Phalops dregei (NE7) 17.93 431 1 102 Savanna (E & W) / Upper Karoo 178 500 430
Phalops flavocinctus (NE6) 20.55 677 716 E Savanna 156 1 564 855
Phalops pauliani (E6) 19.90 400 1 437 Endemic W Savanna - N Namibia 11 66 480
Phalops plancus (NE2) 17.00 232 885 W Savanna - Arid W Namibia 14 39 635
Phalops prasinus (NE2) 18.53 324 1 203 W Savanna - N Namibia 32 115 340

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
759

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe ONTHOPHAGINI (Figs 6K, 6L, 6M) (continued)
Phalops pyroides (E9) 18.71 242 1 133 Endemic SW Arid (Nama Karoo, Arid W Savanna) 61 122 540
Phalops rufosignatus (NE5) 18.63 311 1 161 SW Arid (Nama Karoo, Arid W Savanna) 76 591 895
Phalops smaragdinus (NE6) 20.36 731 825 E Savanna 107 1 479 555
760

Phalops wittei (NE4) 19.18 300 1 102 Kalahari / Upper Karoo / Highveld (Grassland) 254 1 562 330
Phalops zuninoi (NE2) 21.83 674 1 158 W Savanna - N Namibia 2 197 250
Proagoderus aciculatus (NE3) 21.76 862 43 E Coast 74 33 450
Proagoderus aureiceps (NE3) 21.97 763 157 E Savanna / NE Coast - KZN 63 192 750
Proagoderus bicallosus (NE6) 22.27 656 517 E Savanna 52 3 227 420
Proagoderus chalcostolus (NE1) 17.99 829 1 026 Highveld (Grassland) / E Savanna 136 232 570
Proagoderus dives (NE3) 22.05 805 411 E Savanna / NE Coast - KZN 57 1 444 830
Proagoderus furcifer (NE7) 22.20 555 951 Savanna (E & W) 7 267 405
Proagoderus lanista (E4) 17.67 672 1 193 Endemic Highveld (Grassland) / W Savanna - N Namibia 63 300 285
Proagoderus loricatus (NE6) 21.42 604 595 E Savanna 51 1 895 725
Proagoderus plato (NE2) 21.63 613 1 023 W Savanna - N Namibia / N Botswana 2 235 670
Proagoderus quadrituber (NE3) 21.58 858 315 E Savanna 11 134 250
Proagoderus rangifer (NE6) 21.86 598 586 E Savanna 35 1 303 445
Proagoderus rectefurcatus (NE6) 22.51 580 630 E Savanna 20 847 660
Proagoderus sapphirinus (NE4) 19.56 411 1 111 Kalahari / E Savanna 131 1 117 105
Proagoderus tersidorsis (E10) 20.69 697 532 Endemic E Savanna 74 85 690
Tribe SCARABAEINI (Figs 6N, 6O)
Escarabaeus remii (E6) 22.38 503 937 Endemic W Savanna - N Botswana 9 74 550
Escarabaeus satyrus (NE5) 18.17 230 1 017 SW Arid (Nama Karoo, W Savanna, Kalahari) 342 1 491 750
Kheper bonellii (E1) 17.07 223 162 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 36 25 000
Kheper clericus (E7) 21.18 833 161 Endemic NE Coast - KZN - E Savanna 5 750
Kheper cupreus (NE7) 21.04 487 866 Savanna (E & W) 59 1 865 025
Kheper kalaharicus (E8) 20.42 209 953 Endemic Kalahari (SW) 6 6 000
Kheper lamarcki (NE4) 20.67 548 849 Kalahari / E Savanna / NE Coast - KZN 153 2 173 065
Kheper nigroaeneus (NE6) 20.07 672 840 E Savanna 207 662 125
Kheper prodigiosus (NE4) 21.32 484 891 Kalahari / E Savanna 77 2 045 625
Kheper subaeneus (NE6) 20.77 641 750 E Savanna 88 1 257 850
Kheper vethi (NE2) 16.93 193 519 W Savanna - Arid NW Namibia 2 35 400
Kheper zurstrasseni (E4) 18.22 626 1 206 Endemic Highveld (N Grassland) / E Savanna 3 1 450
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Pachylomera femoralis (NE4) 20.94 583 796 E Savanna / Kalahari / NE Coast - KZN 197 2 663 660
Pachylomera opacus (NE4) 19.30 275 1 076 Kalahari / Upper Karoo 78 800 000
Pachysoma aesculapius (E1) 17.05 259 134 Endemic SW Cape - West Coast (Fynbos) 26 10 750
Pachysoma bennigseni (E1) 15.53 41 287 Endemic SW Cape - Namaqualand / Namib (Succ. Karoo) 24 8 500
SURICATA 6 (2020)

Pachysoma denticolle (E5) 16.11 45 578 Endemic SW Arid - Namib Desert 58 40 000
Pachysoma endroedyi (E1) 18.22 156 91 Endemic SW Cape - Namaqualand (Succulent Karoo) 12 130
Pachysoma fitzsimonsi (E5) 15.56 83 1 010 Endemic SW Arid - Namib Desert 13 8 250
Pachysoma gariepinum (E1) 15.63 57 397 Endemic SW Cape - Namaqualand / Namib (Succ. Karoo) 79 17 650
Pachysoma glentoni (E1) 17.91 191 136 Endemic SW Cape - Namaqualand (Succulent Karoo) 9 700
Pachysoma hippocrates (E1) 17.05 166 96 Endemic SW Cape - West Coast (Succ. Karoo, Fynbos) 64 8 500
Pachysoma rodriguesi (E5) 16.16 40 555 Endemic SW Arid - Namib Desert 44 24 000
Pachysoma rotundigena (E5) 16.06 89 1 021 Endemic SW Arid - Namib Desert 18 9 500
Pachysoma schinzi (E5) 14.02 89 1 397 Endemic SW Arid - S Namibia (Nama Karoo, Desert) 9 1 400
Pachysoma striatum (E1) 17.29 114 113 Endemic SW Cape - Namaqualand (Succulent Karoo) 85 6 000
Pachysoma valeflorae (E5) 15.20 30 388 Endemic SW Arid - Namib Desert (Succulent Karoo) 3 200
Scarabaeolus afronitidus (E8) 20.47 347 1 004 Endemic Kalahari (SW) 3 9 250
Scarabaeolus anderseni (E8) 19.81 321 1 070 Endemic Kalahari 55 627 900
Scarabaeolus andreaei (NE3) 23.58 835 83 NE Coast / E Savanna 6 34 050
Scarabaeolus bohemani (NE7) 17.98 332 1 141 Savanna (E & W) / Upper Karoo 290 1 063 280
Scarabaeolus canaliculatus (E5) 15.25 67 894 Endemic SW Arid - Namib Desert 2 910
Scarabaeolus carniphilus (E8) 19.40 334 1 135 Endemic Kalahari (SW) 3 4 285
Scarabaeolus clanceyi (NE3) 22.28 673 167 NE Coast / E Savanna 14 20 500
Scarabaeolus damarensis (E8) 19.65 297 1 078 Endemic Kalahari 113 952 000
Scarabaeolus flavicornis (NE4) 19.12 272 988 Kalahari / SW Cape - Namaqualand (Succ. Karoo) 178 844 500
Scarabaeolus fritschi (E9) 17.63 192 1 047 Endemic SW Arid (Nama Karoo) 63 110 000
Scarabaeolus gracai (NE3) 23.10 615 337 NE Coast / E Savanna 4 3 040
Scarabaeolus inoportunus (E8) 19.38 257 1 055 Endemic Kalahari (SW) 67 510 500
Scarabaeolus inquisitus (E4) 18.89 390 1 141 Endemic Kalahari / Nama Karoo 22 446 500
Scarabaeolus intricatus (E1) 17.09 233 224 Endemic SW Cape (Fynbos, Succulent Karoo) 69 48 000
Scarabaeolus karrooensis (E9) 18.46 210 1 019 Endemic SW Arid (Nama Karoo) 45 359 300
Scarabaeolus kochi (E8) 19.05 274 1 094 Endemic Kalahari (SW) 53 430 000
Scarabaeolus lucidulus (NE4) 20.90 559 1 020 Kalahari (N) / E Savanna 12 382 950

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
761

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe SCARABAEINI (Figs 6N, 6O) (continued)
Scarabaeolus megaparvulus (E9) 17.05 177 1 083 Endemic SW Arid (Nama Karoo) 40 59 650
Scarabaeolus namibensis (NE2) 16.69 60 577 SW Arid - Namib Desert 15 27 000
Scarabaeolus niemandi (E10) 19.63 521 1 006 Endemic E Savanna 5 4 395
762

Scarabaeolus obsoletepunctatus (NE2) 16.90 228 559 W Savanna - Arid NW Namibia 1 7 205
Scarabaeolus pabulator (E9) 18.79 168 941 Endemic SW Arid (Nama Karoo) 40 247 150
Scarabaeolus parvulus (NE5) 18.54 196 991 SW Arid (Nama Karoo, Kalahari (SW), Namib) 108 398 100
Scarabaeolus planipennis (NE3) 22.54 685 92 NE Coast / E Savanna 12 4 900
Scarabaeolus reichei (E1) 17.72 170 99 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 58 12 500
Scarabaeolus rubripennis (E5) 16.01 56 594 Endemic SW Arid - Namib Desert / Succulent Karoo 35 55 500
Scarabaeolus soutpansbergensis (E10) 21.18 434 741 Endemic E Savanna 10 5 100
Scarabaeus alienus (E1) 17.20 65 574 Endemic SW Cape - Namaqualand / Namib Desert 3 18 000
Scarabaeus ambiguus (E4) 18.02 509 1 323 Endemic Highveld (Grassland) / Savanna (E & W) 106 371 530
Scarabaeus basuto (E4) 14.81 455 1 460 Endemic Highveld (Grassland) / Upper Karoo 63 134 360
Scarabaeus bornemisszai (NE3) 22.01 875 12 NE Coast - KZN (Forest) 23 3 250
Scarabaeus caffer (E4) 16.74 891 1 214 Endemic Highveld (E Grassland) 29 70 500
Scarabaeus cognatus (E6) 17.66 77 531 Endemic W Savanna - Arid NW Namibia 5 18 345
Scarabaeus convexus / spretus (E4) 16.13 629 680 Endemic Highveld (E Grassland) / E Coast / S & SW Cape 9 45 980
Scarabaeus costatus (E9) 18.87 225 1 023 Endemic SW Arid (Kalahari (SW), Nama Karoo, Namib) 73 298 810
Scarabaeus deludens (E10) 20.44 610 795 Endemic E Savanna / N Namibia 40 186 900
Scarabaeus ebenus (NE3) 22.49 672 301 NE Coast - KZN / E Savanna 20 422 865
Scarabaeus funebris (E4) 18.93 538 1 135 Endemic Highveld (N Grassland) / Savanna (E & W) 22 104 390
Scarabaeus galenus (NE7) 20.59 630 750 Savanna (E & W) 699 090
Scarabaeus geminogalenus (NE3) 21.83 652 380 NE Coast - KZN / E Savanna 16 40 770
Scarabaeus goryi (NE4) 20.50 593 807 E Savanna 136 2 990 240
Scarabaeus heqvisti (E4) 18.75 652 1 173 Endemic Highveld (N Grassland) 3 305
Scarabaeus hottentorum (E1) 17.57 126 564 Endemic SW Cape - Namaqualand (Succulent Karoo) 7 3 335
Scarabaeus interstitialis (NE6) 18.75 584 1 151 E Savanna 43 119 975
Scarabaeus karae (E4) 16.90 676 1 447 Endemic Highveld (NW Grassland) 3 3080
Scarabaeus piliventris (E1) 17.76 181 295 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 6 3 305
Scarabaeus proboscideus (E8) 18.81 219 808 Endemic Kalahari (SW) / SW Cape - W Coast (S Karoo) 222 502 100
Scarabaeus proximus (E1) 15.85 62 274 Endemic SW Cape - Namaqualand / Namib (Succ. Karoo) 24 16 030
Scarabaeus rugosus (E1) 17.23 186 98 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 86 11 000
SURICATA 6 (2020)
Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Scarabaeus rusticus (E10) 18.25 681 1 228 Endemic E Savanna / Highveld (N Grassland) 156 105 335
Scarabaeus savignyi (E2) 16.76 455 388 Endemic S Cape (Fynbos, Albany Thicket) 7 31 775
Scarabaeus schulzae (E10) 19.41 564 1 016 Endemic E Savanna 13 2 500
Scarabaeus suri (E1) 16.69 276 214 Endemic SW Cape (Fynbos) 38 23 250
SURICATA 6 (2020)

Scarabaeus viator (E3) 17.04 252 1 135 Endemic Upper Karoo 169 288 635
Scarabaeus vicinus (E8) 17.91 309 1 180 Endemic Kalahari (SW) / Upper Karoo 74 791 785
Scarabaeus westwoodi (E4) 11.06 830 2 155 Endemic Highveld (NE Grassland) 8 18 000
Scarabaeus zambesianus (NE4) 20.71 445 947 E Savanna / Kalahari (NE) 100 579 415
Sceliages adamastor (E1) 17.32 337 339 Endemic S & SW Cape (Fynbos) / Upper Karoo 8 174 095
Sceliages brittoni (E1) 17.21 199 100 Endemic SW Cape - West Coast (Fynbos, Succ. Karoo) 17 6 180
Sceliages difficilis (NE6) 17.95 734 1 206 E Savanna / Highveld (NE Grassand) 41 214 130
Sceliages gagates (NE3) 22.43 778 31 NE Coast - KZN 11 13 335
Sceliages granulatus (E8) 18.91 352 1 167 Endemic Kalahari (SW) 7 109 200
Sceliages hippias (E10) 17.86 680 1 302 Endemic E Savanna 41 103 400
Tribe SISYPHINI (Fig. 6P)
Neosisyphus barbarossa (E4) 14.73 682 1 408 Endemic Highveld (E Grassland) / S Cape 23 27 150
Neosisyphus calcaratus (NE6) 21.02 612 744 E Savanna 86 712 200
Neosisyphus confrater (NE6) 19.68 892 687 E Savanna / NE Coast / Highveld (E Grassland) 50 118 955
Neosisyphus fortuitus (NE6) 19.81 746 748 E Savanna 56 154 350
Neosisyphus infuscatus (NE6) 21.18 687 431 E Savanna 56 900 785
Neosisyphus kuehni (E4) 14.11 783 1 646 Endemic Highveld (E Grassland) ) 8 30 620
Neosisyphus macrorubrus (E4) 16.82 366 1 295 Endemic Highveld (W) / Upper Karoo / N Namibia 129 239 965
Neosisyphus mirabilis (NE3) 21.74 825 42 E Coast / E Savanna 35 14 150
Neosisyphus quadricollis (E1) 15.73 298 552 Endemic SW Cape - West Coast (Fynbos) / Upper Karoo 8 25 800
Neosisyphus rubrus (NE6) 18.33 704 1 067 E Savanna / Highveld (Grassland) 299 720 705
Neosisyphus spinipes (NE6) 20.16 823 484 E Savanna / E Coast 126 1 156 170
Sisyphus alveatus (NE2) 22.60 541 919 W Savanna - N Botswana 1 1 801 180
Sisyphus caffer (E4) 16.53 674 1 260 Endemic Highveld (Grassland) 71 31 865
Sisyphus costatus (E4) 15.75 788 1 326 Endemic Highveld (Grassland) / S Cape 45 142 520
Sisyphus fasciculatus (NE3) 19.12 837 755 NE Coast - KZN / E Savanna (Forest) 31 50 820
Sisyphus goryi (NE7) 20.11 666 906 Savanna (E & W) 214 5 042 590
Sisyphus impressipennis (NE7) 20.04 693 987 Savanna (E & W) 59 2 175 705

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
763

***Does not include type locality.

Appendix 3: Summary of average climatic, topographical and distributional data for dung beetle species recorded in South Africa, Botswana and Namibia (continued)

Species/subspecies Average data* (see species accounts for SD) Endemic Main distribution patterns in RSA, Botswana, N 5×5 km2 EOO (km2)
(Generalised distribution patterns) Annual Annual rainfall Altitude status** Namibia (see Fig. 6, Summary Tables 2, 3, & species polygons*
(See summaries in Table 2) temperature (oC) (mm) (m) accounts)
Tribe SISYPHINI (Fig. 6P) (continued)
Sisyphus manni (E10) 17.95 749 1 126 Endemic E Savanna / Highveld (N Grassland) 45 124 410
Sisyphus muricatus (E4) 15.52 709 1 094 Endemic Highveld (E Grassland) / S Cape 28 24 500
Sisyphus nanniscus (NE6) 20.65 811 523 E Savanna / NE Coast - KZN 74 919 450
764

Sisyphus neobornemisszanus (NE3) 22.28 844 8 NE Coast - KZN 22 4 500

Sisyphus oralensis (NE3) 21.99 853 28 NE Coast - KZN 32 18 105
Sisyphus perissinottoi (E2) 16.44 353 608 Endemic S Cape (Albany Thicket, Fynbos) 4 9 685
Sisyphus sordidus (NE6) 21.99 803 225 E Savanna / NE Coast - KZN 59 326 650
Sisyphus splendidus (NE2) 18.67 563 1 495 W Savanna - N Namibia 3 18 600
Sisyphus umbraphilus (E10) 20.34 744 519 Endemic E Savanna 4 21 880

*Average data for total 5×5 km2 polygons occupied in Africa south of 15°S.
**Endemic to South Africa, Botswana and/or Namibia; absence of an entry = non-endemic species.
SURICATA 6 (2020)

***Does not include type locality.

SURICATA 6 (2020) 765

REFERENCES
Abdalla, I.H., Deschodt, C.M., Scholtz, C.H. & Sole, C.L. Balthasar, V. 1939a. Neue arten der coprophagen Scarabaeiden
2018. An update to the taxonomy of the genus Macroderes aus dem Museo Zoologico delle R. Universita di Firenze. Re-
Westwood 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) dia 25: 1–36.
with descriptions of new species from South Africa. Zootaxa Balthasar, V. 1939b. Neue Arten der palaearktischen und neot-
4504: 041–075. ropischen coprophagen Scarabaeiden. Entomologicke Listy.
Ádám, L. 2003. Faunisztikai adatok a Karpat-mendenceböl. Fo- Brno (Folia entomologica) 2: 41–47.
lia Historico Naturalia Musei Matraensis 27: 101–136. Balthasar, V. 1940. Einige neue Scarabaeiden-Arten. Casopis
Ancey, F. 1883. Contributions à la faune de l’Afrique orientale. Ceskoslovenske Spolecnosti Entomologicke 37: 67–76.
Descriptions de Coléoptères nouveaux. Il Naturalista Sicilia- Balthasar, V. 1941a. Eine reihe von neuen coprophagen
no 2: 68–72. Scarabaeiden. Mitteilungen der munchner entomologischen Ge-
Arrow, G.J. 1900. On a collection of insects and arachnids made sellschaft, Munchen 31: 164–184.
in 1895 and 1897, by Mr. C.V.A. Peels, F.Z.S., in Somali­ Balthasar, V. 1941b. Neue coprophage Scarabaeiden des Deutschen
land, with description of new species. Scarabaeidae. Proceed- Entomologischen Instituts (Coleoptera : Scarabaeidae). 71. Bei­
ings of the Zoological Society of London: 22–25. trag zur Kenntnis der Scarabaeidae. Arbeiten über morphologische
Arrow, G.J. 1909. Four new lamellicorn Coleoptera from the und taxonomische Entomologie aus Berlin-Dahlem 8 :181–184.
Oriental region. The Annals and Magazine of Natural History, Balthasar, V. 1942. Neue arten und gattungen der copropha-
including Zoology, Botany and Geology, London 8: 91–94. gen scarabaeiden. 75. Beitrag zur kenntnis der Scarabaeiden.
Arrow, G.J. 1931. The Fauna of British India, Including Ceylon Sbornik entomologickeho oddeleni narodniho Musea v Praze,
and Burma. Coleoptera Lamellicornia Part III (Coprinae). Tay- Praha 20: 188–206.
lor and Francis, London, 1–428. Balthasar, V. 1946. Les nouvelles espèces Aethiopiennes du genre
Arrow, G.J. 1933. Notes on Coprid Coleoptera with descrip- Onthophagus. Acta Societatis Entomologicae Cechosloveniae 43:
tions of a new genus and a few new species. The Annals and 15–19.
Magazine of Natural History, including Zoology, Botany and Balthasar, V. 1956. Einige neue laparosticte Scarabaeiden. 27.
Geology, London 10: 421–430. Beitrag zur Kenntnis der Scarabaeiden. Entomologische Blät-
Balthasar, V. 1937. Neue coprophage Scarabaeiden von der ter, Krefeld 52: 66.
Ukerewe-Insel in Ost-Afrika. Arbeiten uber morphologische Balthasar, V. 1958. Eine neue Untergattung und einige neue Ar-
und taxonomische Entomologie aus Berlin-Dahlem, Berlin- ten der Gattung Copris Geoffroy. Acta entomologica Musei na-
Dahlem 4: 192–203. tionalis Pragae 32: 471–480.
Balthasar, V. 1938. Neue gattungen und arten der Südameri- Balthasar, V. 1959. Beitrag zur Kenntnis der Gattung Ontho-
kanischen Coprophagen. Entomologische Blätter, Krefeld 34: phagus Latr. Acta Entomologica Musei Nationalis Pragae, 33:
210–223. 461–471.
766 SURICATA 6 (2020)

Balthasar, V. 1961a. Zwei neue Gattungen der Familie Scarabaei- Bates, H.W. 1888. Tropical african Coleoptera chiefly from the
dae (Col.). (107. Beitrag zur Kenntnis der Scarabaeidae.). En- Zanzibar mainland. Entomologist’s Monthly Magazine, Lon-
tomologische Arbeiten aus dem Museum G. Frey 12: 174–181. don 24: 237–242.
Balthasar, V. 1961b. Coléoptères Scarabaeidae nouveaux de l’Af- Bedel, L. 1892a. Synopsis des Chironitis, Onitis et Bubas de la
rique Tropicale. Revue de Zoologie et de Botanique africains, région méditerranéenne. L’Abeille, Paris 27: 250–280.
Bruxelles 63: 96–104. Bedel, L. 1892b. Revision des Scarabaeus paléarctiques. L’Abeille,
Balthasar, V. 1963a. Neue arten der Scarabaeidae und Aphodii- Paris 27: 281–288.
dae aus Asien und Afrika. Entomologische Blätter 59: 91–97. Bertolini, J. 1849. Illustratio rerum naturalium Mozambici.
Balthasar, V. 1963b. Eine neue Gattung und neue Arten der Dissertatio 1 de Coleopteris. Novi Commentarii. Academiae
Fam. Scarabaeidae. 114. Beitrag zur Kenntnis der Scarabae- Scientiarum Instituti Bononiensis 10: 381–434.
oidea. Revue de Zoologie et de Botanique Africains, Bruxelles Bertolini, J. 1855. Illustratio rerum naturalium Mozambi-
68: 86–98. co. Coleoptera. Memorie della Reale Accademia delle Scienze
Balthasar, V. 1963c. Neue arten der familie Scarabaeidae. Acta dell’Istituto di Bologna, Classe di Scienze Fisiche 6: 419–446.
Societatis Entomologicae Cechosloveniae 60: 284–295. Bezdek, A. & Sladecek, F.X.J. 2012. Platyonitis oberthueri Jans-
Balthasar, V. 1963d. Monographie der Scarabeidae und Apho- sens, 1942 and Epionitis tarsatus Balthasar, 1942 – synonymy
diidae der palaearktischen und orientalischen Region (Co- confirmed. Zootaxa 3267: 65–68.
leoptera: Lamellicornia). Band 2. Coprinae (Onitini, Oniti- Biswas, S. & Chatterjee, S.K. 1985. Insecta: Coleoptera:
cellini, Onthophagini). Verlag der Tschechoslowakischen Scarabaeidae: Coprinae. Records of the Zoological Survey of In-
Akademie der Wissenschaften, Prague: pp 627. dia 82: 147–177.
Balthasar, V. 1963e. Exploration du Parc National de la Garam- Blackburn, T. 1904. Revision of the Australian Aphodiides, and
ba. Oniticellini. Eine neue art von Drepanocerus. Institut des descriptions of three new species allied to them. Proceedings of
Parcs Nationaux du Congo et du Rwanda. Mission H. de Saeger the Royal Society of Victoria, Melbourne 17: 145–181.
(1949-1952) 40: 131–135. Boheman, C.H. 1857. Insecta Caffrarie annis 1838–1845 a J.A.
Balthasar, V. 1964a. Neue Oniticellinen-Arten. 119. Beitrag zür Wahlberg collecta amici auxilio suffultus descripsit. Coleop-
Kenntnis der Scarabaeoidea. Beitrage zur Entomologie, Berlin tera Officina Norstedtiana. Holmiae 2: 1–395. (Includes spe-
14: 619–624. cies described by Fåhraeus OJ. 1857).
Balthasar, V. 1964b. Caccobiomorphus, eine neue Gattung der Boheman, C.H. 1858. Coleoptera. In: Kongliga Svenska Fregat-
Pinotini. Entomologische Blätter, Krefeld 60: 31–33. ten Eugenies Resa Omkring Jorden under Befal af C.A.Vir-
Balthasar, V. 1965a. Eine neue undergattung und neue arten gin, Aren 1851–53. Vetenskapliga Iakttagelser Pa H.M.K.Os-
der familie Scarabaeidae (121. Beitrag zur Kenntnis der car den Forstes. Kongliga svenska Vetenskapsakademien
Scarabaeoidea). Acta entomologica bohemoslovaca 62: 14–23. Handlingar, Stockholm, 40–58.
Balthasar, V. 1965b. Monographie der untergattung Metacathar- Boheman, C.H. 1860. Coleoptera Caffraria annis 1838-
sius Paulian. Entomologische Abhandlungen Staatlichen Muse- 1845 a J.A. Wahlberg collecta. Öfversigt Kongliga Svenska
um für Tierkunde, Dresden 31: 1–48. Vetenskapsakademiens Förhandlingar 17: 107–120.
Balthasar, V. 1968. Neue Scarabaeiden Arten. Beitrage zur Ento- Bouchard, P., Bousquet, Y., Davies, A.E., Alonso-Zarazaga,
mologie, Berlin 18: 953–958. M.A., Lawrence, J.F., Lyal, C.H.C., Newton, A.F., Reid,
Balthasar, V. 1970. Neue Arten der Scarabaeoidea. Acta Entomo- C.A.M., Schmitt, M., Slipinski, S.A. & Smith, A.B.T. 2011.
logica Bohemoslovaca, Praha 67: 245–253. Family-group names in Coleoptera. Zookeys 88: 1–972.
Barbero, E., Palestrini, C. & Roggero, A. 1998. Phylogenetic Boucomont, A. 1914. Les Coprophages de l’Archipel Malais.
relationships in the genus Scarabaeus Linnaeus. Atti 1. Collo- Annales de la Société Entomologique de France, Paris 83: 238–
quio Nazionale di Sistematica Cladistica. Memorie del Mu- 350.
seo civico di storia naturale di Verona (2), Sezione Scienze Boucomont, A. 1920. Coprophaga Africana. Genre nouveau
della Vita 13: 87–96. et espèces nouvelles. Revue Zoologique Africaine, Bruxelles 8:
Barbero, E., Palestrini, C. & Roggero, A. 2003. Revision of the 261–269.
genus Phalops. Museo Regionale di Scienze Naturale Torino Boucomont, A. 1922. Les Coléoptères Coprophages du genre
Monografia 38: 1–378. Pedaria Cast. Bulletin du Musée Nationale d’Histoire naturelle,
Barbero, E., Palestrini, C. & Roggero, A. 2009a. Systematics Paris 28: 490–495.
and phylogeny of Eodrepanus, a new Drepanocerine genus, Boucomont, A. 1923a. Scarabéides recueillis par le Marquis S.
with comments on biogeographical data. Journal of Natural Patrizi dans l’Afrique Orientale Tropicale. Annali del Museo
History 43: 1835–1878. Civico di Storia Naturale “Giacomo Doria”, Genova (1923-
Barbero, E., Palestrini, C. & Roggero, A. 2009b. Taxonomic 1925) 51: 86–100.
study of Drepanellus gen. n., with the description of two new Boucomont, A. 1923b. Coprophaga africana, 2e note. Revue
species. Deutsche Entomologische Zeitschrift 56: 251–265. Zoologique Africaine 11: 40–58.
Barbero, E., Palestrini, C. & Zunino, M. 1991. Filogenesi, bio- Boucomont, A. 1932. Coprophaga Africana 6e Note. Revue de
geografia e sistematica generica dei Sisyphini (Coleoptera: Zoologie africaine, Tervuren, Bruxelles 22: 46–53.
Scarabaeidae). 16° Congresso Nazionale Italiano di Entomo- Boucomont, A. 1936. Coprophaga Africana 8e Note. Revue de
logia 1: 837–844. Zoologie Africaine, Tervuren, Bruxelles 28: 493–496.
Bates, H.W. 1868. Notes on genera and species of Copridae. Boucomont, A. & Gillet, J.J.E. 1927. Fam. Scarabaeidae. Sub-
Coleopterologische Hefte 4: 87–91. fam. Coprinae II. In Gillet, J.J.E. & Boucomont, Scarabaei-
Bates, H.W. 1887. Biologia Central Americana, Zoologia, Insec- dae : Coprinae, Termitotroginae. Coleopterorum Catalogus,
ta, Coleoptera, Pectinicornia, Lamellicornia (1887–1889). auspiciis et auxilio W. Junk, editus a S. Schenkling. (Partes
Godman F.D. & Salvin O. (Editors) 2: 1–432. 38 et 90). W. Junk, Berlin 19: 103–263.
SURICATA 6 (2020) 767

Branco, T. 1989. Étude de Cambefortius gen.n. et diagnose Cambefort, Y. 1991b. Biogeography and evolution. In: Dung
préliminaire de quatre nouveaux genres d’Onthophagini afro- beetle ecology; Hanski I, Cambefort Y. (eds); Princeton Uni-
tropicaux (Coleoptera, Scarabaeidae). Annales de la Société versity Press. Princeton, USA, 51–68.
Entomologique de France (N.S.) 25: 249–263. Cambefort, Y. 1992a. Révision des espèces afrotropicales du
Branco, T. 1991. Révision des genres du “groupe” Stiptopodius: genre Copris Müller, 1764. X. Espèces nouvelles ou peu
les genres Stiptopodius et Stiptotarsus. Annales de la Société En- connues. Revue française d’Entomologie (N.S.) 14: 179–183.
tomologique de France, Paris (N.S.) 27: 265–285. Cambefort, Y. 1992b. Revision des especes Afro-tropicales du
Branco, T. 2007. Scarabaeoidea (Coleoptera) of Portugal: genus- genre Copris Müller, 1764 IX. Le groupe urus Boheman (Co-
group names and their type species. Zootaxa 1453: 1–31. leoptera, Scarabaeidae). Bulletin de la Societe Entomologique
Branco, T. 2010. Revision of the genera Tiniocellus Péringuey, de France 97: 25–31.
1901 and Nitiocellus gen. n. Boletin de la Sociedad Entomolog- Cambefort, Y. 1993. Notes brèves. Une nouvelle espèce du
ica Aragonesa 47: 71–126. genre Xinidium Harold. Annales de la Société Entomologique
Branco, T. 2011. Scarabaeidae de l’Afrique de l’Ouest : les noms de France 29: 103–105.
du niveau genre et leurs espèces types. Catharsius, La Revue Cambefort, Y. 1996a. Une nouvelle espèce Afro-Tropicale d’Eu-
4: 9–25. oniticellus Janssens, 1953, avec une Phylogénie du genre. Re-
Branco, T. & Josso, J.-F. 2007. Un nouveau Pinacopodius Bran- vue française d’Entomologie (N.S.) Paris 18: 115–119.
co, 1992 et un nouvel Heteroclitopus Péringuey, 1901, du Cambefort, Y. 1996b. Phylogénie et biogéographie du genre
Malawi. Nouvelle Revue d’Entomologie 24: 3–12. Milichus Péringuey, avec la déscription de cinq espèces et
Branco, T. & Ziani, S. 2005. Cheironitis Lansberge, 1875 - its d›une sous-espèce nouvelles. Bulletin de la Société Ento-
correct spelling and validity. Boletin de la Sociedad Entomolog- mologique de France 101: 159–169.
ica Aragonesa 37: 267–272. Cambefort, Y., Gill, B.D. & Moretto, P. 2010. Une nouvelle
Brauns, H. 1914. Descriptions of some new species of myrme- espèce du genre Acanthonitis Janssens. Contribution à la con-
cophilous beetles from southern Rhodesia, with a description naissance des genres d’Onitini. Catharsius, La Revue 2: 1–10
of a new species of Acritus by Dr. H. Bickhardt (Cassel). Pro- Cambefort, Y. & Mauchamp, B. 1980. Coléoptères Scarabaein-
ceedings of the Rhodesian Science Association 13: 32–42. ae nouveaux ou peu connus du Malawi. Revue française d’En-
Brown, J., Scholtz, C.H., Janeau, J.-L., Grellier, S. & Podwojewski, tomologie (N.S.) 2: 177–181.
P. 2010. Dung beetles (Coleoptera: Scarabaeidae) can improve
Castelnau, F.L. de. 1832. Mémoire sur cinquante espèces nou-
soil hydrological properties. Applied Soil Ecology 46: 9–16.
velles ou peu connues d’insectes. Annales de la Société Ento-
Burmeister, H. 1846. III. Coleoptera. 7. Lamellicornia. 27. Co-
mologique de France 1: 386–415.
pris, 4 unnumbered pages. In: Genera Quaedam Insectorum.
Castelnau, F.L. de. 1840. Histoire Naturelle des Insectes
H. Burmeister (ed.). Privately published, Berlin, Germany.
Coléoptères. Avec une introduction renfermant L’Anatomie
Burmeister, H. 1873. Lamellicornia Argentina. Stettiner Ento-
et la Physiologie des Animaux Articulés, par M. Brullé. P.
mologische Zeitung 34: 403–417.
Duménil, Paris 2: 1–564.
Cambefort, Y. 1972. Scarabaeinae récoltés en Mauretanie par
Caveney, S., Scholtz, C.H. & McIntyre, P. 1989. Patterns of dai-
J.-L.Amiet. Nouvelle Revue d’Entomologie, Paris 2: 243–251.
ly flight activity in onitine dung beetles (Scarabaeinae: Oniti-
Cambefort, Y. 1975. Janssensellus, un nouveau genre d’Oniti-
ni). Oecologia 104: 444–452.
ni. Bulletin de la Societe Entomologique de France, Paris 80:
Chevrolat, A. 1830. In: Guérin-Méneville, F.E. 1844. Iconogra-
201–205.
Cambefort, Y. 1978. Canthonina nouveaux ou peu connus phie du Règne Animal de G.Cuvier. Part 7. Insectes. Ballière,
d’Afrique continentale. Journal of the Entomological Society of Paris (1829-1844) 7: 1–576.
Southern Africa 41: 199–213. Costa, A. 1853. Coleotteri, Parte 1a. Famiglia degli Scarabeidei-
Cambefort, Y. 1979. Données préliminaires sur la taxonomie des Scarabaeidea. Costa O.G.:Fauna del Regno di Napoli ossie
Caccobius. Nouvelle Revue d’Entomologie, Paris 9: 119–128. enumarazione di tutti gli animali che abitano le diverse re-
Cambefort, Y. 1980. Nouveaux Onthophagides du Kenya. Zool- gioni di questo Regno e le acque che le bagnano contenente
ogische Mededelingen Leyden 55: 195–200. la descrizione de’ nuovi o poco esattemente conosciuti con
Cambefort, Y. 1981. La nidification du genre Cyptochirus (Cole- figure ricavate da original viventi e dipinte al naturale. G.
optera, Scarabaeidae). Comptes Rendus de l’Academie des Sci- Sautto, Napoli, 1–44.
ences, Paris 292: 379–381. Creutzer, C. 1799. Entomologische Versuche (Kleine Beitrage
Cambefort, Y. 1982. Pedaria nouveaux ou peu connus de Cote zur nahren Berichtigungen einiger Kaferarten mit 3 aus-
d’Ivoire. Revue française d’Entomologie (N.S.) 4: 57–62. gemahften Kupfertafeln von Sturm J.). Wien, Austria.
Cambefort, Y. 1983. Nouveaux Scarabaeidae s.str. de Côte Daniel, G.M. 2019. The nomenclatural status of the generic
d’Ivoire et de pays voisins. Revue française d’Entomologie nomina Epirinus Dejean, 1833 and Epirhinus Agassiz, 1846
(N.S.) Paris 5: 7–9. (Coleoptera, Scarabaeidae, Scarabaeinae). Bionomina 15:
Cambefort, Y. 1984. Étude écologique des Coléoptères 63–65.
Scarabaeidae de Côte d’Ivoire. Annexe : Descriptions prélim- Daniel, G.M., Davis, A.L.V. & Scholtz, C.H. 2016. Three new
inaires d’un nouveau genre et de 36 nouvelles espèces, et nou- Sisyphus species from shaded vegetation in southern Africa.
velle synonymie. Travaux des chercheurs de la Station de Lamto Zootaxa 4147: 67–74.
(RCI) 3: 1–12. Daniel, G.M., Davis, A.L.V., Sole, C.L. & Scholtz, C.H. 2018.
Cambefort, Y. 1985a. Révision du genre Xinidium Harold. An- Taxonomic review of the tribe Sisyphini sensu stricto Mulsant,
nales de la Société Entomologique de France 21: 95–102. 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) in southern
Cambefort, Y. 1985b. Les espèces orientales du genre Cleptocac- Africa, including new species descriptions. Insect Systematics
cobius. Revue française d’Entomologie (N.S.) 7: 125–130. & Evolution 9: 1–61.
768 SURICATA 6 (2020)

Daniel, G.M., Sole, C.L., Davis, A.L.V., Strümpher, W.P. & Davis, A.L.V., Scholtz, C.H. & Harrison, J. du G. 1999b. New
Scholtz, C.H. 2020. Systematics of the dung beetle tribe Si- and threatened Afrotropical dung beetle taxa in the Gondwa-
syphini Mulsant (Scarabaeidae: Scarabaeinae) inferred from a naland tribe Canthonini. African Entomology 7: 77–84.
molecular phylogeny and biogeography of southern African Davis, A.L.V., Scholtz, C.H. & Sole, C.L. 2017. Biogeograph-
species. Systematic Entomology 45: 73–84. ical and co-evolutionary origins of scarabaeine dung beetles:
Davis, A.L.V. 1986. Three new Afrotropical dung beetles in the Mesozoic vicariance versus Cenozoic dispersal and dinosaur
genera Kheper Janssens, Gymnopleurus Illiger and Onitis Fa- versus mammal dung. Biological Journal of the Linnean Society
bricius, with notes on related species. Journal of the Entomo- 120: 258–273.
logical Society of Southern Africa 49: 373–387. Davis, A.L.V., Scholtz, C.H. & Swemmer, A.M. 2012. Effects of
Davis, A.L.V. 1993. Biogeographical groups in a southern Af- land usage on dung beetle assemblage structure: Kruger Na-
rican, winter rainfall, dung beetle assemblage (Coleoptera: tional Park versus adjacent farmland in South Africa. Journal
Scarabaeidae) - consequences of climatic history and habitat of Insect Conservation 16: 399–411.
fragmentation. African Journal of Ecology 31: 306–327. Davis, A.L.V., Stals, R. & Deschodt, C.M. 2015. New replace-
Davis, A.L.V. 1994. Associations of Afrotropical Coleoptera ment name for the species Scarabaeus (Scarabaeolus) nitidus
(Scarabaeidae, Aphodiidae, Staphylinidae, Hydrophilidae, Davis & Deschodt, 2015 – not an American pest. Zootaxa
Histeridae) with dung and decaying matter: implications for 4057: 582.
selection of fly-control agents for Australia. Journal of Natural Davis, A.L.V., Swemmer, A.M., Scholtz, C.H., Deschodt, C.M.
History 28: 383–399. & Tshikae, B.P. 2014. Roles of environmental variables and
Davis, A.L.V. 1995. Daily weather variation and temporal dy- land usage as drivers of dung beetle assemblage structure in
namics in an Afrotropical dung beetle community (Coleop- mopane woodland. Austral Ecology 39: 313–327.
tera: Scarabaeidae). Acta Oecologica 16: 641–656. Davis, A.L.V., Van Aarde, R.J., Scholtz, C.H. & Delport, J.H.
Davis, A.L.V. 1996a. Diel and seasonal community dynamics in 2002. Increasing representation of localized dung beetles
an assemblage of coprophagous, Afrotropical, dung beetles across a chronosequence of regenerating vegetation and nat-
(Coleoptera: Scarabaeidae s. str., Aphodiidae, Staphylinidae: ural dune forest in South Africa. Global Ecology and Biogeog-
raphy 11: 191–210.
Oxytelinae). Journal of African Zoology 110: 291–308.
Davis, A.L.V., Van Aarde, R.J., Scholtz, C.H., Guldemond,
Davis, A.L.V. 1996b. Habitat associations in a South Afri-
R.A.R., Fourie, J. & Deschodt, C.M. 2013. Is microclimate-
can, summer rainfall, dung beetle community (Coleoptera:
driven turnover of dung beetle assemblage structure in regen-
Scarabaeidae, Aphodiidae, Staphylinidae, Histeridae, Hydro-
erating coastal vegetation a precursor to re-establishment of
philidae). Pedobiologia 40: 260–280.
a forest fauna? Journal of Insect Conservation 17: 565–576.
Davis, A.L.V. 1997. Climatic and biogeographical associations
DeGeer, C. 1778. Mémoires pour servir à l’Histoire des Insectes
of southern African dung beetles (Coleoptera: Scarabaeidae s.
(tom. 1-7. 1752-1778). Hesselberg, Stockholm 7: 1–950.
str.). African Journal of Ecology 35: 10–38.
Dejean, P.F.M.A. 1821. Catalogue de la collection de Coléoptères
Davis, A.L.V. 2002. Dung beetle diversity in South Africa: in-
de M. le Baron Dejean. Crevot Libraire, Paris: 136.
fluential factors, conservation status, data inadequacies, and
Dejean, P.F.M.A. 1833. Catalogue des Coléoptères de la collec-
survey design. African Entomology 10: 53–65.
tion de M. le Comte Dejean (1833-1835). Deuxième Édi-
Davis, A.L.V. & Deschodt, C.M. 2018. Two new species of Gar- tion. Méquignon-Marvis Père et Fils, Paris: 1–443.
reta Janssens, 1940 (Coleoptera: Scarabaeidae: Scarabaeinae) Dejean, P.F.M.A. 1836. Catalogue des Coléoptères de la Collec-
from southern Africa. Zootaxa 4450: 242–248. tion de M. le Comte Dejean. Troisième Édition, Revue, Cor-
Davis, A.L.V., Deschodt, C.M. & Scholtz, C.H. 2019. Describ- rigée et Augmentée, Fascicles 1–4. Méquignon-Marvis Père
ing new tribes to resolve discrepancies between phylogeny et Fils, Paris, France.
and tribal classification in the subfamily Scarabaeinae (Cole- Deschodt, C.M. & Davis, A.L.V. 2017. Transfer of three spe-
optera: Scarabaeidae). Zootaxa 4608: 131–144. cies of Namakwanus Scholtz & Howden to Versicorpus De-
Davis, A.L.V., Frolov, A.V. & Scholtz, C.H. 2008. The African schodt, Davis & Scholtz or to Namaphilus gen. nov., with
dung beetle genera. Protea Book House, Pretoria: 1–272. descriptions of two new species (Coleoptera: Scarabaeidae:
Davis, A.L.V. & Scholtz, C.H. 2020. Dung beetle conservation Scarabaeinae). Annals of the Ditsong National Museum of Nat-
biogeography in southern Africa: current challenges and po- ural History 7: 109–114.
tential effects of climatic change. Biodiversity and Conserva- Deschodt, C.M. & Davis, A.L.V. 2018. New deltochiline (Co-
tion 29: 667–693. leoptera: Scarabaeidae: Scarabaeinae) taxa associated with
Davis, A.L.V., Scholtz, C.H. & Chown, S.L. 1999a. Species hyrax dung in arid south western Namibia. Zootaxa 4450:
turnover, community boundaries, and biogeographical com- 066–076.
position of dung beetle assemblages across an altitudinal gra- Deschodt, C.M. & Davis, A.L.V. 2019. New southern African
dient in South Africa. Journal of Biogeography 26: 1039–1055. species and a revalidation in the dung beetle genus Gyrono-
Davis, A.L.V., Scholtz, C.H. & Deschodt, C. 2005. A dung bee- tus van Lansberge, 1874 (Scarabaeidae: Scarabaeinae) with an
tle survey of selected Gauteng nature reserves: implications updated key. Zootaxa 4624: 275–282.
for conservation of the provincial scarabaeine fauna. African Deschodt, C.M., Davis, A.L.V. & Daniel, G.M. 2019. The en-
Entomology 13: 1–16. demic southern African genus Epirinus Dejean, 1833 (Cole-
Davis, A.L.V., Scholtz, C.H., Dooley, P.W., Bham, N. & Kry- optera: Scarabaeidae: Scarabaeinae) revisited: five new species
ger, U. 2004. Scarabaeine dung beetles as indicators of bio- and two new synonyms. Zootaxa 4603: 327–340.
diversity, habitat transformation and pest control chemicals Deschodt, C.M., Davis, A.L.V. & Scholtz, C.H. 2011. New
in agro-ecosystems. South African Journal of Science 100: dung beetle taxa from arid south-western Africa. Insect Sys-
415–424. tematics & Evolution 42: 277–294.
SURICATA 6 (2020) 769

Deschodt, C.M., Davis, A.L.V. & Scholtz, C.H. 2015a. A new productivity and fly control. Journal of Applied Ecology 24:
synonymy in the fidius group of Copris Müller 1764 and a 837–851.
new species from the highland grasslands of South Africa. Erichson, W.F. 1835. Neue Südamerikanische Käfergattungen
Zootaxa 3949: 431–438. aus der Familie der Blätterhörner. Archiv für Naturgeschichte
Deschodt, C.M., Davis, A.L.V. & Scholtz, C.H. 2015b. Status 1: 256–270.
changes, new synonymies, key and descriptions of seven new Erichson, W.F. 1843. Beitrag zur Insecten-Fauna von Angola, in
species in the subgenus Scarabaeus (Scarabaeolus) Balthasar besonderer Beziiehung zur geographischen Verbreitung der In-
1965. Zootaxa 3931: 505–527. secten in Africa. Archiv für Naturgeschichte, Berlin 9: 199–267.
Deschodt, C.M., Davis, A.L.V. & Scholtz, C.H. 2016. A new Erichson, W.F. 1847a. Conspectus Insectorum Coleopterum,
monotypic genus, a species synonymy and nomenclatural quae in Republica Peruana observata sunt. Archiv für Natur-
corrections in the arid-adapted Canthonini from the Suc- geschichte 13: 67–185.
culent Karoo Biome of south-western Africa. Zootaxa 4147: Erichson, W.F. 1847b. Naturgeschichte der Insecten Deutsch-
490–500. land. Erste Abteilung.Coleoptera. Dritter Band. Nicolaische
Deschodt, C.M., Davis, A.L.V. & Scholtz, C.H. 2017. Three Verlags-Buchhandlung, Berlin. Lieferung 5: 641–800.
new species, a synonymy and taxonomic notes on southern Erichson, W.F. 1848. Naturgeschichte des Insecten Deutsch-
African members of the dung beetle genera Scarabaeus s. str. land. I.Coleoptera, Scaphidilia, Scarabaeides. Nicolaischen
Linnaeus, 1758 and Escarabaeus Zídek & Pokorný, 2011. Buchhandlung, Berlin (1847–1848) 1: 1–968.
Journal of Natural History 51: 299–316. Eschscholtz, J.F. 1822. Entomographien. O. Reimer, Berlin.
Deschodt, C.M., Davis, A.L.V. & Sole, C.L. 2019. A beautiful Excoffier, L., Laval, G. & Schneider, S. 2006. Arlequin ver. 3.1.
new species of Hyalonthophagus Palestrini & Giacone, 1988 An integrated software package for population genetics data
(Coleoptera: Scarabaeidae: Scarabaeinae), from the Northern analysis. Available at: <http://cmpg.unibe.ch/software/arle-
Cape Province, South Africa. Zootaxa 4648: 194–200. quin3> (Minimum Spanning Tree module no longer avail-
Deschodt, C.M., Kryger, U. & Scholtz, C.H. 2008. New taxa of able as part of the Arlequin package).
relictual Canthonini dung beetles utilizing rock hyrax mid- Fabricius, J.C. 1775. Systema Entomologiae: sistens insectorum
dens as refuges in South-western Africa. Insect Systematics & classes, ordines, genera, species, adiectis synonymis, locis,
Evolution 38: 361–376. descriptionibus, observationibus. Officina Libraria Kortii.
Deschodt, C.M. & Scholtz, C.H. 2008. Systematics of South Flensburgi & Lipsiae, 1–832.
African forest-endemic dung beetles: new genera and species Fabricius, J.C. 1776. Genera Insectorum corumque characteres
of small Canthonini. African Entomology 16: 91–106. naturales secundum numerum, figuram, situm et propotio-
Deschodt, C.M. & Sole, C.L. 2019. A new species of Versicorpus nem omnium partium oris, adiecta mantissa specierum. Kil-
Deschodt, Davis & Scholtz, 2011 (Scarabaeidae: Scarabaein- iae: 1–310.
ae: Byrrhidiini) from the Brandberg Mountain, Namibia. Fabricius, J.C. 1781. Species insectorum, exhibentes eorum dif-
Zootaxa 4671: 139–144. ferentias specificas, synonyma auctorum, loca natalia, meta-
Dormont, L., Jay-Robert, P., Bessière, J.-M., Rapior, S. & Luma- morphosin, adiectis observationibus, descriptionibus. Tome
ret, J.-P. 2010. Innate olfactory preferences in dung beetles. I. Carol.Ernest.Bohnii, Hamburgi et Kilonii, VIII: 1–552.
Journal of Experimental Biology 213: 3177–3186. Fabricius, J.C. 1792. Entomologia systematica emendata et auc-
Doube, B.M. 1983. The habitat preference of some bovine dung ta. Secundum Classes, Ordines, Genera, Species adjectis Syn-
beetles (Coleoptera: Scarabaeidae) in Hluhluwe Game Reserve, onimis, Locis, Observationibus, Descriptionibus. Hafniae 1:
South Africa. Bulletin of Entomological Research 73: 357–371. 1–538, C. G. Proft & fils.
Doube, B.M. 1991. Dung beetles of southern Africa. In: Dung Fabricius, J.C. 1794. Entomologia systematica emendata et auc-
beetle ecology; Hanski I, Cambefort Y. (eds); Princeton Uni- ta. Secundum Classes, Ordines, Genera, Species adjectis syn-
versity Press, Princeton, USA, 133–155. onymis, locis, observationibus, descriptionibus. Hafniae 4:
Doube, B.M.1991. Appendix B.8. In: Dung beetle ecology; 1–472, CG Proft & fils.
Hanski I, Cambefort Y. (eds); Princeton University Press, Fabricius. J,C. 1798. Supplementum Entomologiae Systemati-
Princeton, USA, 382–387. cae. Hafniae: 1–572.
Doube, B.M., Giller, P.S. & Moola, F. 1988. Dung burial strate- Fabricius, J.C. 1801. Systema Eleutheratorum secundum or-
gies in some South African coprine and onitine dung beetles. dines, genera, species: adiectis synonymis, locis, observationi-
Ecological Entomology 13: 251–261. bus, descriptionibus. Tomus I. Impensis bibliopoli academici
Endrödi, S. 1971. The scientific Results of the Hungarian Zo- novi, Kiliae: 1–506.
ological Expeditions to Tanganyika. 16. Coleoptera Lamel- Fåhraeus, O.J. 1857. See Boheman, C.H. 1857.
licornia (partim). Folia Entomologica Hungarica, Budapest 24: Fairmaire, L. 1884a. Diagnoses des Coléoptères de l’Afrique ori-
289–311. entale (suite). Annales de la Société Entomologique de Belgique.
Endrödi, S. 1973. Entomological Explorations in Ghana by Dr. Comptes Rendus des seances, Bruxelles 28: 121–125.
S. Endrody-Younga 14, Lamellicornia I. Annales historico- Fairmaire, L. 1884b. Diagnoses des Coléoptères de l’Afrique.
naturales Musei nationalis hungarici, Budapest 65: 195–229. Annales de la Société Entomologique de Belgique. Comptes Ren-
Endrödi, S. 1976. Lamellicornia aus Ghana II. Annales historico- dus des seances Bruxelles 28: 142–149.
naturales Musei nationalis hungarici, Budapest 68: 155–164. Fairmaire, L. 1887. Coléoptères des voyages de M.G.Revoil chez
Edwards, P.B. 1988. Field ecology of a brood-caring dung bee- les Somalis et dans l’interieur du Zanguebar. Annales de la
tle Kheper nigroaeneus - habitat predictability and life history Société Entomologique de France, Bulletin 7: 102–134.
strategy. Oecologia 75: 527–534. Fairmaire, L. 1888. Enumération des Coléoptères recueillis par
Edwards, P.B., & Aschenborn, H. 1987. Patterns of nesting and M. le Dr.Hans Schinz dans le Sud de l’Afrique. Annales de la
dung burial in Onitis dung beetles: implications for pasture Société Entomologique de France, Paris 6: 173–202.
770 SURICATA 6 (2020)

Fairmaire, L. 1891. Coléoptères de l’Afrique Orientale. Annales du Rwanda. Mission H. De Saeger (1949–1952). Fascicule
de la Société Entomologique de Belgique. Compte Rendus des 30: 1–123.
Séances 35: 279–289. Ferreira, M.C. 1963. Contribuiçao para o conhecimento dos
Fairmaire, L. 1893. Note sur les Coléoptères de Choa. Annales Escarabídeos africanos. Novos Taxa Entomológicos 30: 3–39.
de la Société Entomologique de Belgique. Compte Rendus des Ferreira, M.C. 1964a. Contribuiçao para o estudo do género
Séances 37: 9–50. Epirinus Reiche. Novos Taxa Entomológicos 37: 3–13.
Felsche, C. 1907. Coprophage Scarabaeiden. Deutsche Entomol- Ferreira, M.C. 1964b. Contribuiçao para o estudo do subgénero
ogische Zeitschrift, 273–296. Metacatharsius Paulian. Descriçao de especies novas. Novos
Felsche, C. 1909. Neue und alte coprophage Scarabaeiden. Taxa Entomológicos 34: 3–32.
Deutsche Entomologische Zeitschrift, 751–765. Ferreira, M.C. 1965. Contribuiçao para o estudo dos Es-
Felsche, C. 1911. Coprophage Scarabaeiden. Deutsche Entomol- carabídeos de Moçambique. Descriçao de uma espécie nova
ogische Zeitschrift, 133–141. do subgénero Metacatharsius Paulian. Novos Taxa Entomológi-
Ferreira, M.C. 1952a. Duas espécies novas de Scarabaeus da cos 45: 3–10.
África do Sul. Boletim Sociedade de Estudos da Provincia de Ferreira, M.C. 1968. Contribuiçao para o estudo dos Es-
Moçambique 22: 19–24. carabídeos de Moçambique. Descriçao de uma espécie nova
Ferreira, M.C. 1952b. A new species of Scarabaeus. Annals of the do subgénero Scarabaeolus Balthasar. Novos Taxa Entomológi-
Transvaal Museum, Pretoria 22: 73–77. cos 58: 3–9.
Ferreira, M.C. 1953a. Monografia dos Escarabaeideos da Africa Ferreira, M.C. 1969. Contribuiçao para o conhecimento dos
do Sul. Tribo-Scarabaeini. I Parte Sub-tribo Pachysomides. Escarabídeos de África. Descriçao de uma espécie nova de
Boletim Sociedade de Estudos da Provincia de Moçambique 23: Scarabaeus s. str. Novos Taxa Entomológicos 68: 3–4.
1–80. Ferreira, M.C. 1971. Contribuiçao para o estudo dos Es-
Ferreira, M.C. 1953b. Contribuiçao para o estudo dos Es- carabídeos Africanos. Descriçao de uma espècie nova de Ca-
carabaeídeos da Africa do Sul. Aditamento à I parte da tharsius s.str. Novos Taxa Entomológicos 96: 5–6.
Monografia dos Escarabaeideos da Africa do Sul. Boletim So- Ferreira, M.C. 1972 (1968–1972). Os escarabídeos de África
ciedade de Estudos da Provincia de Moçambique 80: 1–9. (sul do Sáara). Revista Entomologia Moçambique 11: 1–1088.
Ferreira, M.C. 1953c. Contribuiçao para o estudo dos Es- Ferreira, M.C. 1976a. A. Some new species of Onitis F. (Coleop-
carabaeídeos da África do Sul. Memorias Museum Dr. Alvaro tera: Scarabaeidae). B. A new species of Chironitis Lansberge
de Castro. Lourenço Marques 2: 23–26. (Coleoptera : Scarabaeidae). Navorsinge van die Nasionale
Ferreira, M.C. 1954a. Monografia dos Escarabaeídeos da África Museum 3 (9): 181–198.
do Sul. V. Parte. Boletim Sociedade de Estudos da Provincia de Ferreira, M.C. 1976b. A new species of Platyonitis Janssens from
Moçambique 87: 1–15. Angola. Navorsinge van die Nasionale Museum 3 (8): 177–180.
Ferreira, M.C. 1954b. Contribuiçao para o estudo dos Es- Ferreira, M.C. 1977. Contribution to the knowledge of the Af-
carabídeos da África do Sul. Uma nova espécie do género rican Scarabaeidae. A new genus of Onitini and some new
Gymnopleurus Illiger. Boletim Sociedade de Estudos da Provin- species of Onitis F. Navorsinge van die Nasionale Museum 3
cia de Mocambique 84: 1–4. (10): 181–198.
Ferreira, M.C. 1954c. Algumas variedades novas de Gymno- Ferreira, M.C. 1978. The genus Onitis F. of Africa south of the
pleurineos e notas sinonimicas. Boletim Sociedade de Estudos Sahara. Memoirs van die Nasionale Museum 10: 1–410.
da Provincia de Mocambique 85: 1–3. Fincher, G.T. 1973. Nidification and reproduction of Phanaeus
Ferreira, M.C. 1954d. Some Scarabaeina in the Durban Mu- spp. in three textural classes of soil (Coleoptera: Scarabaei-
seum and Art Gallery. Durban Museum Novitates, Durban dae). Coleopterists Bulletin 27: 33–37.
4: 91–95. Forgie, S.A., Grebennikov, V.V. & Scholtz, C.H. 2002. Revi-
Ferreira, M.C. 1958a. Duas espécies novas do género Copris sion of Sceliages, a millipede-eating genus of southern African
Geoff. Novos Taxa Entomológicos 9: 3–5. dung beetles. Invertebrate Systematics 16: 931–955.
Ferreira, M.C. 1958b. Coleoptera Scarabaeidae. Coprinae. Re- Forgie, S.A., Philips, T.K. & Scholtz, C.H. 2005. Evolution of
sults of the Lund University Expedition in 1950-1951. In: B. the Scarabaeini (Scarabaeidae: Scarabaeinae). Systematic En-
Hanström, P. Brinck, G. Rudebeck eds. South African Animal tomology 30: 60–96.
Life 5: 479–516. Forgie, S.A., Kryger, U., Bloomer, P. & Scholtz, C.H. 2006. Evo-
Ferreira, M.C. 1958b. A new species of Scarabaeus from Angola. lutionary relationships among the Scarabaeini (Coleoptera:
Novos Taxa Entomológicos 1: 3–6. scarabaeidae) based on combined molecular and morpholog-
Ferreira, M.C. 1958c. A new species of Scarabaeus from South ical data. Molecular and Phylogenetic Evolution 40: 662–678.
West Africa. Novos Taxa Entomológicos 5: 3–4. Frey, G. 1955. Coprophagen-Studien über neue und wenig
Ferreira, M.C. 1960a. Revisao das espécies africanas de Cathar- bekannte Arten. Entomologische Arbeiten aus dem Museum G.
sius s.str. Do grupo adamastor e descricao de especies novas. Frey 6: 1057–1066.
Revista Entomologia Moçambique 3: 1–77. Frey, G. 1963. Neue Onthophagen aus Australien und aus Af-
Ferreira, M.C. 1960b. Revisao das espécies etiopicas do sub- rika. Entomologische Arbeiten aus dem Museum G. Frey 14:
genero Catharsius s.str. Revista Entomologia Moçambique 3: 538–549.
155–270. Frey, G. 1967. Zwei neue afrikanische Coprophagen sowie eine
Ferreira, M.C. 1962a. Contribuçao para o conhecimento dos neue Synonymie. Entomologische Arbeiten aus dem Museum
Escarabídeos de Angola. Publiçoes cultural Companhia Dia- G. Frey 18: 404–405.
mantes Angola 54: 121–190. Frey, G. 1975a. Neue Coprophagen aus dem Zentralafrikan-
Ferreira, M.C. 1962b. Exploration du Parc National de la Ga- ischen Museum in Tervuren und aus meine museum. Ento-
ramba. Coprinae. Institut des Parcs Nationaux du Congo et mologische Arbeiten aus dem Museum G.Frey 26: 292–296.
SURICATA 6 (2020) 771

Frey, G. 1975b. Neue Sudafrikanische Onthophagini gesammelt gli auspici della società geografica italiana. Risultati zoologici.
vom dung beetle research unit Pretoria. Entomologische Arbe- XVI. Coleotteri. Annali del Museo civico di Storia Naturale di
iten aus dem Museum G. Frey 26: 191–197. Genova, Genova (Series 2) 15: 309–341.
Frolov, A.V. 2004. New and little known species of the Afrotrop- Giller, P.S. & Doube, B.M. 1994. Spatial and temporal co-
ical dung beetle genus Sarophorus and a phylogenetic analysis occurrence of competitors in Southern African dung beetle
of the genus. Journal of Afrotropical Zoology 1: 95–100. communities. Journal of Animal Ecology 63: 629–643.
Frolov, A.V. 2014. Revision of the genus Delopleurus Boheman Gillet, J.J.E. 1907. Remarques sur quelques coprides du Museo
(Coleoptera: Scarabaeidae: Scarabaeinae) with description Civico de Gênes. Annali del Museo Civico di Storia Naturale
of new species from Africa. Journal of Natural History 49: “Giacomo Doria”, Genova 3: 587–603.
129–154. Gillet, J.J.E. 1908. Coprides d’Afrique tropicale. Descriptions
Frolov, A.V., Akhmetova L.A. & Scholtz, C.H. 2008. Revision d’espèces nouvelles et remarques sur quelques espèces con-
of the obligate mushroom-feeding African “dung beetle” ge- nues. Mémoires de la Société Entomologique de Belgique 16:
nus Coptorhina Hope (Coleoptera: Scarabaeidae: Scarabaein- 63–82.
ae). Journal of Natural History 42: 1477–1508. Gillet, J.J.E. 1909. Collections receuillis dans le Sahara et ré-
Frolov, A.V. & Scholtz, C.H. 2003a. Revision of the Afrotropi- gions voisines, par René Chudeau. Bulletin du Muséum na-
cal dung beetle genus Sarophorus Erichson. African Entomol- tionale d’Histoire Naturelle, Paris 15: 432–437.
ogy 11: 183–198. Gillet, J.J.E. 1911. Coprophaga Africana. Descriptions d’espèces
Frolov, A.V. & Scholtz, C.H. 2003b. A new genus and species nouvelles et remarques diverses. Annales de la Société Ento-
of dung beetle from southern Namibia. African Entomology mologique de Belgique 55: 308–312.
11: 297–299. Gillet, J.J.E. 1918. Coprides nouveaux de la région éthiopienne
Frolov, A.V. & Scholtz, C.H. 2004. Revision of the southern et remarques diverses. Zoologische Mededelingen, Leiden 4:
African genus Macroderes Westwood. Annales de la Société En- 146–158.
tomologique de France (N.S.) 40: 373–393. Gistel, J. 1857. Achthundert und zwanzig neue oder unbeschrie-
Frolov, A.V. & Scholtz, C.H. 2005. Revision of the southern bene wirbellose Thiere charakterisirt von Doctor Johannes
African genus Frankenbergerius Balthasar with description of Gistel. Schorner’sche Buchhandlung, Straubing. Vacuna 2:
513–606.
new taxa. Journal of Natural History 39: 2355–2377.
Goeze, J.A.E. 1777. Entomologische Beytrage zu des Ritter
Füessly, J.C. 1775. Verzeichnis der ihm bekannten Schweizeri-
Linne zwolften Ausgabe des Natursystems. Weidmanns, Er-
schen Insecten mit einer ausgemahlten Kupfertafel, nebst der
ben und Reich, Leipzig 1: 1–736.
Unkundigung eines neuen Insekten Werks. Winterthur &
Gomes Alves, M.-L. 1954. Alguns Escarabídeos de Moçambique
Füessly, Zürich: 1–62.
– II. Junta das Missões Geográficas e de Investigações do Ul-
Gebler, F.A. 1841. Characteristik mehrerer neuen Sibirischen
tramar, Lisboa. Anais, Vol. IX, Tomo III. Estudos de Zoolo-
Coleopteren. Bulletin scientific publié par Académie impériale
gia. Trabalhos da Missão Zoológica de Moçambique e do Centro
des Sciences de Saint-Petersbourg 8: 369–376.
de Zoologia (1957) 9: 55–61.
Génier, F. 2013. Transfert d’Onthophagus bubalus Harold, 1867,
Gory, M. 1833. Monographie du genre Sisyphe. Mequignon-
dans le genre Phalops Erichson, 1847, et notes sur sa position
Marvis pere et fils, Paris: 1–15.
phylogénétique. Catharsius, La Revue 7: 1–4.
Griffith, E. & Pidgeon, E. 1831. The Class Insecta arranged by
Génier, F. & Moretto, P. 2017. Digitonthophagus Balthasar, the Baron Cuvier, with Supplementary Additions to each Or-
1959: taxonomy, systematics, and morphological phylogeny der. Notices of New Genera and Species by G. Gray. Volume
of the genus revealing an African species complex (Coleop- the first. 446-537 In: The Animal Kingdom arranged in con-
tera: Scarabaeidae: Scarabaeinae). Zootaxa 4248: 1–110. formity with its Organisation by the Baron Cuvier, with Sup-
Génier, F. & Davis, A.L.V. 2017. Digitonthophagus gazella auc- plementary Additions to each Order; Griffith E. et al. (eds);
torum: an unfortunate case of mistaken identity for a widely Volume 14 (dated 1832). Whittaker, Treacher, & Co., Lon-
introduced species (Coleoptera: Scarabaeidae: Scarabaeinae: don: 1–570.
Onthophagini). Zootaxa 4221: 497–500. Guérin-Méneville, F.E. 1844. Iconographie du Règne Animal
Génier, F. & Josso, J.-P. 2016. Notes on Afrotropical Cathar- de G. Cuvier. Part 7. Insectes. Ballière, Paris (1829–1844)
sius species described by Edgar von Harold (Coleoptera: 7: 1–576.
Scarabaeidae, Scarabaeinae). Zootaxa 4072: 333–342. Haaf, E. 1955. Uber die gattung Sisyphus Latreille. Entomolo-
Geoffroy, E.L. 1762. Histoire Abrégée des Insectes, volume 1. Ca- gische Arbeiten aus dem Museum G. Frey 6: 341–381.
lixte-Volland and Rémont, Paris. Halffter, G. & Favila ME. 1993. The Scarabaeinae (Insecta: Co-
Gerstaecker, A. 1871. Beitrag zur Insektenfauna von Zanzibar. leoptera): an animal group for analyzing, inventorying and
III. Coleoptera. Archiv für Naturgeschichte, Berlin 37: 42– monitoring biodiversity in tropical rainforest and modified
349. landscapes. Biodiversity International 27: 15–21.
Gerstaecker, A. 1873. Die Gliedertier-Fauna des Sansibar- Hanski, I. & Cambefort, Y. 1991a. Competition in dung bee-
Gebietes. Decken C. C. Baron Carl Claus von der Decken’s tles. In: Dung beetle ecology; Hanski I, Cambefort Y. (eds);
Reisen in Ost Afrikas, Berlin: 1–542. Princeton University Press, Princeton: 305–329.
Gerstaecker, A. 1884. Bestimmungen der von Herrn Dr. G.A. Hanski, I. & Cambefort, Y. 1991b. Index of the genera in
Fischer wahrend seiner Reise nach dem Massai-Land gesa- Scarabaeidae. In: Dung beetle ecology; Hanski I, Cambefort Y.
mmelten Coleopteren. Jahresbuch der hamburgischen Wissen- (eds); Princeton University Press, Princeton: 465–473.
schaftlichen Anstalten, Hamburg (1883) 1: 43–63. Harold, E. von. 1859. Beitrage zur kenntnis einiger coprophagen
Gestro, R. 1895. Esplorazione del Giuba e dei suoi affluenti lamellicornien. (I. Stück). Berliner Entomologische Zeitschrift,
compiuta dal cap. V. Bottegno durante gli anii 1892-93 sotto Berlin 3: 193–224.
772 SURICATA 6 (2020)

Harold, E. von. 1862. Beiträge zur Kenntniss einiger copropha- Fortsetzung der von Büffonschen Naturgeschichte. Nach
gen Lamellicornien (Viertes Stuck). Berliner Entomologische dem System des Ritters von Linné und Fabricius. Ben Joa-
Zeitschrift, Berlin 6: 379–403. chim Pauli, Berlin 2: 1–330.
Harold, E. von. 1867a. Zur Kenntniss der Gattung Canthidium Hoffmannsegg, J.C. 1817. Entomologische Bemerkungen bei
und ihrer nächsten Verwandten. Coleopterologische Hefte 1: Gelegenheit der Abhandlungen über amerikanische Insecten.
1–61. Zoologisches Magazin 1: 8–56.
Harold, E. von. 1867b. Zur Synonymie zweier Gymnopleurus- Hope, F.W. 1833. Characters of several new genera and species
Arten. Coleopterologische Hefte 1: 73–75. of Coleopterous Insects. Proceedings of the Zoological Society,
Harold, E. von. 1867c. Beiträge zur Kenntniss der Gattung London 1: 61–64.
Onthophagus (I.Theil: Australier und Afrikaner). Coleopter- Hope, F.W. 1837. The Coleopterists Manual, Containing the
ologische Hefte 2: 23–59. Lamellicorn Insects of Linneus and Fabricius. Henry G.
Harold, E. von. 1868a. Diagnosen neuer Coprophagen. Coleop- Bohn, London: 121.
terologische Hefte 4: 79–86. Hope, F.W. 1838. Observations on the lamellicorns of Olivier.
Harold, E. von. 1868b. Abänderungen vergebener Namen. Co- The Entomological Magazine 5: 312–326.
leopterologische Hefte 4: 104. Hope, F.W. 1842. On some new insects from Western Africa.
Harold, E. von. 1868c. Diagnosen neuer Coprophagen. Coleop- The Annals and Magazine of natural History, including Zoolo-
terologische Hefte 3: 80–86. gy, Botany and Geology 1: 494–496.
Harold, E. von. 1869a. Diagnosen neuer Coprophagen. Coleop- Houston, W.W.K. 1983. Identification of the dung beetles Oni-
terologische Hefte 5: 95–104. tis crenatus Reiche from Ethiopia and O. crenatus sensu Bo-
Harold, E. von. 1869b. Ueber coprophage Lamellicornien mit heman from southern Africa. Journal of the Entomological So-
besonderer Berücksichtigungen der Pariser Sammlungen. ciety of Southern Africa 46: 249–256.
Coleopterologische Hefte 5: 46–70. Howden, H. & Scholtz, C.H. 1987. A revision of the African
Harold, E. von. 1870. Berichtigungen und Zusätze zum Catalo- genus Odontoloma. Journal of the Entomological Society of
gus Coleopterorum synonymicus et systematicus. Coleopter- Southern Africa 50: 155–192.
Howden, H. & Scholtz, C.H. 1988. A second species of the
ologische Hefte 6: 94–110.
African genus Peckolus. Journal of the Entomological Society of
Harold, E. von. 1871a. Diagnosen neuer Coprophagen. Coleop-
Southern Africa 51: 133–136.
terologische Hefte, Munchen 7: 112.
ICZN. 1993. Opinion 1708. Proagoderus Lansberge, 1883 (In-
Harold, E. von. 1871b. Verzeichniss der von Dr. Beccari in Bo-
secta, Coleoptera) : conserved. The Bulletin of Zoological No-
gos gesammelten coprophagen Lamellicornien. Coleopterolo-
menclature 50: 75–76.
gische Hefte, Munchen 8: 1–28.
Illiger, J.K.W. 1803. Verzeichniss der in Portugal einheimischen
Harold, E. von. 1871c. Diagnosen neuer Coprophagen. Coleop-
Käfer. Erste Lieferung. Magazin für Insektenkunde 2: 186–258.
terologische Hefte 8: 114–116.
Inouchi, J. & Shibuya, T. 1986. Odour responses of the olfacto-
Harold, E. von. 1873. Diagnosen neuer coprophagen. Coleopter-
ry receptor cells in the dung beetles. Chemical Senses 11: 286.
ologische Hefte 11: 102–105.
IUCN. 2001. IUCN Red List Categories and Criteria: Version
Harold, E. von. 1875. Diagnosen neuer Arten. Coleopterologische
3.1. IUCN Species Survival Commission. IUCN, Gland,
Hefte 14: 137–140.
Switzerland and Cambridge, UK.
Harold, E. von. 1877a. Enumeration des Lamellicorn co-
Jacobson, F.V.S. 1906. Revue critico-bibliographique. Kolbe.
prophages rapportes de l’Archipel Malais, de la Nouvelle H.J. Ueber die Lebensweise und die geographische Verbreit-
Guinée et de l’Australie boreale par M.M.J. Doria, O. Becca- ung der coprophagen Lamellicornien. Russkoe Entomologich-
ri et L.M. D’Albertis. Annali del Museo Civico di Storia Natu- eskoe Obozrenie 6: 312–315.
rale di Genova 10: 38–110. Jacobs, C.T. & Scholtz, C.H. 2015. A review on the effect of
Harold, E. von. 1877b. Coleopterorum species novae. Mit- macrocyclic lactones on dung-dwelling insects: Toxicity of
teilungen der munchner entomologischen Verein, Munchen 1: macrocyclic lactones to dung beetles. Onderstepoort Journal
97–111. of Veterinary Research 82: 8.
Harold, E. von. 1878. Ueber Coleopteren aus dem tropischen Jankielsohn, A., Scholtz, C.H. & Louw, S. v.d. M. 2001. Ef-
Afrika. Mittheilungen der munchner entomologischen Verein, fect of habitat transformation on dung beetle assemblages:
Munchen 2: 38–48. a comparison between a South African nature reserve and
Harold, E. von. 1880. Einige neue Coleopteren. Mitteilungen neighboring farms. Environmental Entomology 30: 474–483.
der munchner entomologischen Verein, Munchen 4: 148–171. Janssens, A. 1936. Un nouveau genre d’Onitide africain. Bulle-
Harold, E. von. 1886. Coprophage Lamellicornien beschrieben tin du Musée royal d’Histoire Naturelle de Belgique, Bruxelles
von E. von Harold. Berliner Entomologische Zeitschrift, Berlin 12: 1–3.
30: 141–149. Janssens, A. 1937. Révision des Onitides. Verhandelingen Konin-
Harrison, J. du G., Scholtz, C.H. & Chown, S.L. 2003. A re- klijk Natuurhistorisch Museum Belgie, Brussel 2: 1–200.
vision of the endemic south-western African dung beetle Janssens, A. 1938a. Exploration du Parc National Albert. Scarabaei-
subgenus Scarabaeus (Pachysoma), including notes on other ni. Institut des parcs nationaux du Congo Belge 21: 1–76.
flightless Scarabaeini. Journal of Natural History 37: 305–355. Janssens, A. 1938b. Notes sur les Onitides africains (Coleop-
Hausmann, J.F.L. 1807. Beiträge zur Insektenfauna des Vorge- tera Lamellicornia), Troisième Note. Bulletin du Musée royal
birges der guten Hoffnung. Magazin für Insektenkunde, K. d’Histoire naturelle de Belgique 14: 13.
Illiger, Braunschweig 6: 229–267. Janssens, A. 1939a. Exploration du Parc National Albert. Mis-
Herbst, J.F.W. 1789. Der Käfer. Jablonsky C.G.: Natursystem sion G.F. De Witte (1933-1935). Coprini. Institut des parcs
aller bekannten in- und auslandischen Insecten, als eine nationaux du Congo Belge 29: 1–104.
SURICATA 6 (2020) 773

Janssens, A. 1939b. Contribution à l’étude des Coléoptères Kabakov, O.N. 2006. Beetles of the subfamily Scarabaeinae of
Lamellicornes Coprophages. Bulletin du Musée Royal d’His- the fauna of Russia and adjacent countries. Moscow, Scientif-
toire Naturelle de Belgique, Bruxelles 15 :1–6. ic publications KMK: 1–374.
Janssens, A. 1940a. Monographie des Gymnopleurides (Coleop- Kirby, W. 1818. A century of insects, including several new gen-
tera Lamellicornia). Mémoires du Musée Royal d’Histoire Na- era described from his cabinet. Transactions of the Linnean
turelle de Belgique, 2e sér. 18: 1–74. Society, London 12: 375–453.
Janssens, A. 1940b. Monographie des Scarabaeus et genres vois- Kirby, W. 1828. A description of some Coleopterous insects in
ines. Verhandelingen Koninklijk Natuurhistorisch Museum Bel- the collection of the Rev. F.W. Hope. Zoological Journal 3:
gie, Brussel 2: 1–81. 520–525.
Janssens, A. 1942. Contribution à l’étude des Coléoptères Klug, J.C.F. 1835. V. Insekten (Coleoptera). In: Verzeichniss von
Lamellicornes Coprophages. V. Quelques Onitides nouveaux Thieren und Pflanzen, welche auf einer Reise um die Erde ge-
ou peu connus. Bulletin du Musée royal d’Histoire Naturelle de sammelt wurden von A. Erman. G. Reimer, Berlin, 27–50.
Belgique, Bruxelles 18:1–9. Klug, J.C.F. 1855. Diagnosen der neuer Coleopteren aus Mos-
Janssens, A. 1943. Contribution a l’étude des coléoptères Lamel- sambique. Berichten der Koniglich-Preussischen Akademie der
licornes coprophages. VII. Notes sur quelques Coprinae Wissenschaften, Berlin 20: 643–660.
d’Afrique. Bulletin du Musée royal d’Histoire Naturelle de Bel- Klug, J.C.F. 1862. Peters W.C.H.: Naturwissenschaftliche Reise
gique 19: 1–8. nach Mossambique auf Befehl Seiner Majestät des Königs
Janssens, A. 1946. Contribution à l’étude des coléoptères lamel- Friedrich Wilhelm IV in den Jahren 1842 bis 1848 aus-
licornes coprophages, XI. - Table synoptique et essai de clas- geführt. Zoologie V. Insecten und Myriapoden. Georg Rein-
sification pratique des Scarabaeidae Laparosticti. Bulletin du er, Berlin: 1–566.
Musée Royal d’Histoire Naturelle de Belgique 22: 1–13. Kolbe, H.J. 1887. Beiträge zur Zoogeographie Westafrikas nebst
Janssens, A. 1949. Un Scarabaeinae termitophile nouveau du einem Bericht über die während der Loango-Expedition von
Congo Belge. Revue de Zoologie et de Botanique africains, Herrn Dr. Falkenstein bei Chinchoxo gesammelten Coleop-
Bruxelles 42: 183–184. tera. Nova Acta Caesareae Leopoldina Carolinae germanicae
Janssens, A. 1950. Contribution à l’étude des Coléoptères naturae curiosorum Halle 50: 153–364.
Kolbe, H.J. 1893a. Ueber afrikanische Mistkäfer (Cathar-
Lamellicornes. XV. Un nouveaux et curieux genre d’Onitini.
sius, Diastellopalpus). Sitzungsbericht der Gesellschaft natur-
Bulletin de l’Institut royal des Sciences naturelle de Belgique,
forschender Freunde zu Berlin: 131–151.
Bruxelles 26: 1–4.
Kolbe, H.J. 1893b. Beiträge zur Kenntniss der Mistkafer, Lamel-
Janssens, A. 1953. Exploration du Parc National de l’Upemba.
licornia Onthophila. Stettiner entomologische Zeitung, Stettin
Mission G.F. De Witte (1946-1949). Oniticellini. Institut des
54: 188–202.
parcs nationaux du Congo Belge 11: 1–118.
Kolbe, H.J. 1895. Beiträge zur Kenntniss der Mistkafer, Lamel-
Josso, J.F. 2011. Description de quatre nouvelles espèces de
licornia Onthophila. Stettiner entomologische Zeitung, Stettin
Scarabaeidae de Zambie, appartenant aux genres Catharsius
56: 329–345.
Hope, 1837, Copris Müller, 1764, Pleuronitis Lansberge,
Kolbe, H.J. 1897. Käfer und Netzflüger Ost-Afrikas. Moebius
1875 et Onthophagus Latreille, 1802. Catharsius, La Revue
K.: Deutsch Ostafrika. Wissenschaftliche Forschungsresul-
3: 1–8.
tate über Land und Leute unseres ostafrikanischen Schutzge-
Josso, J.F. 2012. Les Onitis Fabricius, 1798 du Groupe XI de bietes und der Angrenzenden Länder. Band IV. Die Thierwelt
Janssens : redéfinition du groupe, mise en synonymie, de- Ost-Afrikas und der Nachbargebiete. Wirbellose Thiere, Co-
scription de six espèces nouvelles et clé de détermination. Ca- leopteren. Dietrich Reimer, Berlin 4: 1–367.
tharsius, La Revue 5: 6–15. Kolbe, H.J. 1905. Über die Lebensweise und die geographische
Josso, J.F. 2014. Description de quatre nouvelles espèces de Pro- Verbreitung der coprophagen Lamellicornier. Zoologische
agoderus Lansberge, 1883, de Zambie, du Mozambique et du Jahrbücher, Supplement 8: 475–594.
Malawi. Catharsius, La Revue 10: 1–8. Kolbe, H.J. 1908. Dynastidae, Cetoniidae und Scarabaeidae.
Josso, J.F. & Prévost, P. 2001. Contribution à la connaissance Schultze L.: Zoologische und Arthropologische ergebnisse
des Scarabaeidae africains I. Les Cahiers Magellanes Hors Série einer forschungsreise im westlichen und zentralen südafrika.
2: 1–14. Bd.1. Denkschrift Medizinisch-Naturwissenschaftlichen Ge-
Josso, J.F. & Prévost, P. 2003. Contribution à la connaissance sellschaft zu Jena 13: 121–132.
des Scarabaeidae africains II. Etude du genre Pedaria. Les Kolbe, H.J. 1914. Wissenschaftliche Ergebnisse der deutschen
Cahiers Magellanes Hors Série 7: 1–39. Zentral-Afrika-Expedition 1907-1908 unter Führung Adolf
Josso, J.F. & Prévost, P. 2006a. Contribution à la connaissance Friedrichs, Herzog zu Mecklemburg. Band V: Zoologie III.
des Scarabaeidae africains III, étude du genre Pedaria (2). Les Lieferung 3. Scarabaeiden. Klinkhardt & Biermann, Leipzig
Cahiers Magellanes Hors Série 14: 1–20. 5: 261–416.
Josso, J.F. & Prévost, P. 2006b. Description de cinq Ontho- Kraatz, G. 1895. Zwei neue westafrikanische Oniticellus. Deut-
phagini nouveaux pour l’Afrique de l’Ouest (Parc du W). Les sche Entomologische Zeitschrift, Berlin 34: 142–144.
Cahiers Magellanes Hors Série 21: 1–10. Krajcik, M. 2006. New species of Kheper Janssens from Namib-
Josso, J.F. & Prévost, P. 2009a. Contribution à la connaissance ia. Animma X 14: 19–23.
des Scarabaeidae africains IV. Etude du genre Pedaria (3). Les Král, D. 2014. Delopleurus mencli sp. nov. from Socotra Island.
Cahiers Magellanes Hors Série 30: 1–22. Acta Entomologica Musei Nationalis Pragae 54 (supplemen-
Josso, J.F. & Prévost, P. 2009b. Description de trois nouvelles tum), 133–138.
espèces de Catharsius de Zambie et Tanzanie. Les Cahiers Kràl, D. & Rejsek, J. 1999. The Liatongus species with moveable
Magellanes Hors Série 31: 1–8. clypeal horn. Folia Heyrovskyana Supplementum 5: 1–18.
774 SURICATA 6 (2020)

Krell, F.-T. 2004. East African dung beetles (Scarabaeidae) at- Lansberge, J.W. van. 1887. Note III. Cinq espèces nouvelles de
tracted by defensive secretions of millipedes. Journal of East Coléoptères exotiques appartenant au Musée de Leyde. Notes
African Natural History 93: 69–73. from the Leyden Museum, Leyden 9: 107–112.
Krell, F.-T., Branco, T. & Ziani, S. 2012. Case 3590 Scarabaeus Latreille, P.A. 1802. Histoire Naturelle, Générale et Particulière
Linnaeus, 1758, Dynastes MacLeay, 1819, SCARABAEIN- des Crustacés et des Insectes. Tome troisième. F. Dufart, Par-
AE Latreille, 1802, and DYNASTINAE MacLeay, 1819 is: 464.
(Insecta, Coleoptera, SCARABAEOIDEA): proposed con- Latreille, P.A. 1804. Histoire naturelle, générale et particuliere
servation of usage. Bulletin of Zoological Nomenclature 69: des crustacés et des insectes. Paris 10: 1–445.
182–190. Latreille, P.A. 1807. Genera Crustaceorum et Insectorum secun-
Krell, F.-T. & Philips, K. 2010. Formicdubius Philips & Scholtz dum Ordinem naturalem in Familias disposita, Iconibus Es-
from South Africa, a junior synonym of Haroldius Bouco- emplisque plurimis explicata (1806-1809). A. Koenig Parisis
mont, and a survey of the trichomes in the African species. et Argentorati Volume I–IV 2: 1–280.
Zookeys 34: 41–48. Latreille, P.A. 1810. Considérations générales sur l’ordre naturel
Krikken, J. 1974. A new species of Platyonitis Janssens from des animaux composant les classes des crustacés, des arach-
Kenya, with notes on the genus. Zoologische Mededelingen nides, et des insectes; avec un tableau méthodique de leurs
Leyden 48: 195–203. genres, disposés en familles. F. Schoell, Paris: 444.
Krikken, J. 1977. Notes on African Onitini mainly from South- Latreille. P,A. 1823. Description des Insectes d’Afrique receuillis
eastern Kenya. Zoologische Mededelingen Leyden 51: 141–170. par M. Cailliaud Voyage à Méroé. Debure, Paris 4: 279–287.
Krikken, J. 2009. Drepanocerine dung beetles: a group over- Latreille, P.A. 1825. Familles Naturelles du Règne Animal, Ex-
view, with description of new taxa. Haroldius 4: 3–30. posées Succinctement et Dans un Ordre Analytique avec l’in-
Kryger, U., Deschodt, C., Davis, A.L. & Scholtz, C.H. 2006. dication de Leurs Genres. Encyclopédie Méthodique. J.-B.
Effects of cattle treatment with a cypermethrin/cymiazol Baillière, Paris: 1–570.
spray on survival and reproduction of the dung beetle species Latreille, P.A. 1829. Le Règne Animal Distribué d’Après son
Euoniticellus intermedius (Coleoptera: Scarabaeidae). Bulletin Organisation, pour Servir de Base a` l’Histoire Naturelle
of Entomological Research 96: 597–603. des Animaux et d’Introduction à l’Anatomie Comparée. Par
Kryger, U., Deschodt, C., Davis, A.L. & Scholtz, C.H. 2007. M. Le Baron Cuvier, avec Figures Dessinées d’Après Nature.
Effects of cattle treatment with a fluazuron pour-on on sur-
Nouvelle Édition, Revue et Augmentée. Tome IV. Crustacés,
vival and reproduction of the dung beetle species Onthopha-
Arachnides et Partie des Insectes. Déterville, Paris, France.
gus gazella (Fabricius). Veterinary Parasitology 143: 380–384.
Leach, W.E. 1815. Entomology. In: Edinburgh Encyclopaedia,
Lacordaire, J.T. 1856. Histoire naturelle des insectes. Genera
volume 9, part 1; Brewster (ed.); Edinburgh: 57–172.
des Coléoptères ou exposé méthodique et critique de tous
LeConte, J.L. 1861. Article III. Classification of the Coleoptera
les genres proposés jusqu’ici dans cet ordre d’insectes. Tome
of North America, part 1. Smithsonian Miscellaneous Collec-
troisième contenant les familles des pectinicornes et lamel-
tions, Washington 3: 1–286.
licornes. Librairie Encyclopédique de Roret, Paris: 1–594.
Le Peletier, A.L.M. & Serville, J.G.A. 1828. Scarabé, Scarabaeus.
Lansberge, J.W. van. 1874a. Observations sur la classification
In: Encyclopédie Méthodique. Histoire Naturelle. Entomolo-
des Lamellicornes coprophages. Annales de la Société Ento-
gie, ou Histoire Naturelle des Crustacés, des Arachnides et
mologique de Belgique 17: 177–193.
des Insectes, volume 10, part 2; Latreille, P.A., Le Peletier,
Lansberge, J.W. van. 1874b. Description de quelques Lamel-
licornes Coprophages. Coleopterologische Hefte 12: 4–12. A.L.M., Serville, J.G.A., Guérin-Méneville, F.E. (eds); Mme
Lansberge, J.W. van. 1875a. Observations sur la classification Veuve Agasse, Paris: 346–382.
des Lamellicornes coprophages. Annales de la Société Ento- Lesne, P. 1900. Collections entomologiques recueillies au cours
mologique de Belgique 17: 177–193. de la mission Bonchamps et remise au Muséum par M. Ch.
Lansberge, J.W. van. 1875b. Monographie des Onitides. Annales Michel. Coléoptères. In: Mission de Bonchamps. Vers Facho-
de la Société Entomologique de Belgique 18: 1–148. da à la rencontre de la mission Marchand à travers l’Éthiopie.
Lansberge, J.W. van. 1882. Enumeration des Scarabaeides rap- Plon-Nourrit et Cie, Paris: 498–501.
portés du pays des Somalis (Afrique Equatoriale) par M. Lewis, G. 1895. On the Lamellicorn Coleoptera of Japan, and
Révoil. Avec diagnoses des espèces nouvelles. Annales de la notices of others. Annals and Magazine of Natural History (se-
Société Entomologique de Belgique 26: 21–31. ries 6) 16: 374–408.
Lansberge, J.W. van. 1883. Description de trois espèces nou- Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae,
velles d’Onthophagus appartenant au Musée Royal de Leyde. Secundum Classes, Ordines, Genera, Species, cum Charac-
Note VII. Notes from the Leyden Museum 5 :14–16. teribus, Differentiis, Synonymis, Locis, edition 10, volume
Lansberge, J.W. van. 1885a. Note III. Description de quatre es- 1. Laurentii Salvii, Stockholm.
pèces nouvelles de Coprophages appartenant au Musée de Löbl, I. 2006. New nomenclatural and taxonomic acts, and
Leyde. Notes from the Leyden Museum 7: 17–20. comments: Scarabaeidae: Scarabaeinae. In: Catalogue of Pa-
Lansberge, J.W. van 1885b. Descriptions d’espèces nouvelles laearctic Coleoptera. Volume 3: Scarabaeoidea – Scirtoidea –
de Coléoptères appartenant au Musée civique de Gênes. An- Dascilloidea – Buprestoidea – Byrrhoidea; Löbl I, Smetana
nali del Museo Civico di Storia Naturale Giacomo Doria 2: A. (eds.); Apollo Books, Stenstrup: 31–32.
375–400. Lumaret, J.-P. & Moretto, P. 1983. Contribution à l’étude des
Lansberge, J.W. van. 1886. Scarabaéides, Buprestides et Ceram- Oniticellini: nidification et morphologie larvaire d’Oniti-
bycides de l’Afrique occidentale, envoyés au Musée de Leyde cellus rhadamistus (F.) et considerations sur la position tax-
par MM. Veth et Van der Kellen (Note XIII). Notes from the onomique de cette espèce. Annales de la Société Entomologique
Leyden Museum, Leyden 8: 69–120. de France (N.S.) 19: 311–316.
SURICATA 6 (2020) 775

Macleay, W.S. 1821. Horae entomologica: or essays on the an- Moretto, P. 2011. Nouveaux Scarabaeidae africains appartenant
nulose animals. S.Bagster, London 1: 161–524. aux genres Lophodonitis Janssens, 1938 et Copris Müller,
Marchisio, R. & Zunino, M. 2012. Il genere Copris Müller Tas- 1764. Notes sur quelques Onitis Fabricius, 1798. Catharsius,
sonomia, filogenesi e note di zoogeografia. WBA Monographs La Revue 3: 9–19.
2, Verona: 1–176. Moretto, P. 2013a. Descriptions de nouveaux Onthophagus La-
Masumoto, K. 1987. Coprophagid beetles from Northwest treille, 1802 d’Afrique tropicale. Notes de taxonomie et de
Thailand I. The Entomological Review of Japan 42: 125–131 faunistique. Entomologia Africana 18: 48–55.
Maud, R.R. 2012. Macroscale geomorphic evolution. In: South- Moretto, P. 2013b. La validité du nom du niveau genre Pro-
ern African Geomorphology: Recent Trends and New Directions; agoderus. Descriptions de deux nouveaux Proagoderus Lans-
Holmes P, Meadows M. (eds); Sun Media, Bloemfontein: berge, 1883, d’Afrique tropicale, et de la femelle de Proag-
5–21. oderus cerathoterium Janssens, 1953. CATHARSIUS La Revue
McGeoch, M.A., Van Rensburg, B.J. & Botes, A. 2002. The 8: 1–11.
verification and application of bioindicators: a case study of Moretto, P. 2016a. Description d’un nouveau Canthonini de
dung beetles in a savanna ecosystem. Journal of Applied Ecol- Namibie appartenant au genre Dicranocara Frolov & Scholtz,
ogy 39: 661–672. 2003 (Coleoptera, Scarabaeidae). Entomologia Africana 21:
Medina, C.A. & Scholtz, C.H. 2005. Systematics of the south- 47–49.
ern African genus Epirinus Reiche: descriptions of new spe- Moretto, P. 2016b. Description d’une nouvelle espèce de
cies and phylogeny. Insect Systematics & Evolution 36: 1–16. Scarabaeolus Balthasar, 1965, d’Afrique de l’Ouest (Coleop-
Mittal, I.C. 2011. Synoptic key to Indian species of genus Garre- tera, Scarabaeidae). CATHARSIUS La Revue 13: 8–23.
ta Janssens with a new species from Haryana (India). Journal Moretto, P. & Nicolas, J.L. 2002. Description de cinq nouveaux
of Entomological Research 35: 295–298. Proagoderus d’Afrique. Entomologia Africana 7: 29–37.
Mlambo, S., Sole, C.L. & Scholtz, C.H. 2015. A molecular phy- Moretto, P. & Nicolas, J.L. 2004. Emmendata: un nouveau nom
logeny of the African Scarabaeinae (Coleoptera: Scarabaei- pour Proagoderus palestrinii Moretto & Nicolas, 2002. Ento-
dae). Arthropod Systematics & Phylogeny 73: 303–321. mologia Africana 9: 48.
Moczek, A.P. 2002. Allometric plasticity in a polyphenic beetle. Moretto, P. & Perissinotto, R. 2013. Description and ecology
Ecological Entomology 27: 58–67. of two new species of Gyronotus van Lansberge, 1874 from
Monaghan, M.T., Inward, D.J.G., Hunt, T. & Vogler, A.P. 2007.
southern Africa. ZooKeys 344: 73–82.
A molecular phylogenetic analysis of the Scarabaeinae (dung
Mostert, L.E. & Holm, E. 1982. Notes on the flightless
beetles). Molecular Phylogenetics and Evolution 45: 674–692.
Scarabaeina with a description of a new species. Cimbebasia
Montreuil, O. 1998. Analyse Phylogénétique et Paraphylie des
A 5: 273–284.
Coprini et Dichotomiini. Scénario Biogéographique. Annales
Motschulsky, V. 1860. Sur les Collections Coléoptèrologiques de
de la Société Entomologique de France (N.S.) 34: 135–148.
Linné et de Fabricius. (Continuation). XXII. Lamellicornes.
Montreuil, O. 2014. Deux nouveaux Sisyphus Latreille de Tan-
Études Entomologiques, 8: 147–162.
zanie. Bulletin de la Société Entomologique de France 119:
Mucina, L. & Rutherford, M.C. 2006. The vegetation of South
177–180.
Africa, Lesotho and Swaziland. Strelitzia 19: 1–807.
Montreuil, O. 2015a. Le genre Neosisyphus Müller en Afrique.
Müller, G. 1942. Nuovi coleotteri dell’Africa Orientale (2a se-
Catharsius, La Revue 12: 1–36.
rie). Atti del Museo Civico di Storia Naturale di Trieste 15:
Montreuil, O. 2015b. Nouveaux Sisyphus Latreille d’Afrique
Bulletin de la Société Entomologique de France 120: 91–102. 63–86.
Montreuil, O. 2015c. Premiers cas de brachyptérisme dans le Müller, G. 1947. Nuovi Coleotteri dell’Africa orientale (Quarta
genre Sisyphus Latreille. Annales de la Société Entomologique serie). Atti del Museo Civico di Storia Naturale di Trieste 16:
de France (N.S.) 51: 281–293. 81–93.
Montreuil, O. 2017. Nouveaux Sisyphus Latreille, 1807, du Müller, G. 1949. Nuovi coleotteri dell’Africa orientale: V serie.
groupe seminulum : le complexe arboreus. Bulletin de la So- Atti del Museo Civico di Storia Naturale di Trieste 17: 152–
ciété Entomologique de France 121: 167–174. 161.
Moretto, P. 1998. Kheper festivus (Harold): un complexe d’es- Mulsant, É. 1842. Histoire naturelle des Coléoptères de France.
pèces. Entomologia Africana 3: 27–38. Lamellicornes. [Tome 2]. Maison Libraire, Paris: 623.
Moretto, P. 2004. Description de nouveaux Proagoderus d’Af- Nguyen-Phung, T. 1987. Révision des Copris afro-tropicaux
rique et remarques sur quelqeus espèces peu ou mal connues V. Une nouvelle espèce du <groupe fidius Olivier>. [Col.
(2ème contribution à la révision des Proagoderus). Entomolo- Scarabaeidae]. Bulletin de la Société Entomologique de France
gia Africana 9: 20–47. 91: 271–273.
Moretto, P. 2007. Les Onitis Fabricius, 1798 du XIIIème groupe Nguyen-Phung, T. 1988a. Révision des espèces Afrotropicales
de Janssens, espèces nouvelles et espèces affines. Entomologia du genre Copris. IV. Le groupe fidius Müller, 1764 [Cole-
Africana 12: 2–14. optera, Scarabaeidae]. Revue française d’Entomologie, Paris
Moretto, P. 2008. Catharsius karlwerneri, nouvelle espèce d’Af- (N.S.) 10: 5–15.
rique de l’Est. Révision des Catharsius Hope, 1837 du groupe Nguyen-Phung, T. 1988b. Révision des espèces afro-tropicales
“heros”. Entomologia Africana 13: 38–44. du genre Copris Müller, 1764 VI. Le groupe bootes Klug (Co-
Moretto, P. 2009. Essai de classification des Onthophagus La- leoptera, Scarabaeidae). Nouvelle Revue d’Entomologie (N.S.)
treille, 1802 africains des 5ème et 6ème groupes de d’Orbig- 5: 247–258.
ny. Nouvelle Revue d’Entomologie (N.S.) 25: 145–178. Nguyen-Phung, T. 1988c. Révision des espèces afrotropicales du
Moretto, P. 2010. Les Scarabéides coprophages de Bayanga genre Copris Müller, 1764, VII. Le groupe corniger Sahlberg.
en République Centrafricaine. Bulletin de la Société Ento- [Col. Scarabaeidae]. Bulletin de la Société Entomologique de
mologique de France 115: 455–477. France 92: 209–219.
776 SURICATA 6 (2020)

Nguyen-Phung, T. 1989. Révision des espèces afrotropicales du Palestrini, C. & Giacone P. 1988. Filogenesi e zoogeografia del
genre Copris Müller, 1764 (Coleoptera: Scarabaeidae). VIII. sottogenere Hyalonthophagus nov. Atti XV Congresso Nazio-
Le groupe orion Klug. Annales de la Société Entomologique de nale Italiano di Entomologia L’Aquila: 319–327.
France (N.S.) 25: 199–208. Palestrini, C. & Giacone, P. 1989. Systematics Phylogeny and
Nguyen-Phung, T. & Cambefort, Y. 1986a. Révision des espèces biogeography of the subgenus Hyalonthophagus. Elytron 3:
afro-tropicales du genre Copris Müller, 1764. I. Le groupe in- 79–101.
teger Reiche (Coleoptera, Scarabaeidae). Nouvelle Revue d’En- Pallas, P.S. 1781. Icones Insectorum praesertim Rossiae Siberi-
tomologie (N.S.) 3: 401–416. aeque pecularium quae collegit et Descriptionibus illustravit.
Nguyen-Phung, T. & Cambefort, Y. 1986b. Révision des es- Walther, Erlangae: 1–96.
pèces afro-tropicales du genre Copris Müller, 1764. III. Le Partridge, T.C. & Maud, R.R. 1987. Geomorphic evolution of
groupe inhalatus Quedenfeldt [Coleoptera, Scarabaeidae]. southern Africa since the Mesozoic. South African Journal of
Revue française d’Entomologie, Paris (N.S.) 8: 100–106. Geology 90: 179–208.
Nguyen-Phung, T. & Cambefort, Y. 1987. Révision des espèces Paschalidis, K.M. 1974a. The genus Sisyphus Latr. (Coleoptera:
afro-tropicales du genre Copris Müller, 1764. II. Le groupe Scarabaeidae) in Southern Africa. MSc. Thesis, Rhodes Uni-
jacchus (Fabricius) (Coleoptera Scarabaeidae s. str.). Annales versity, Grahamstown, South Africa, pp 183.
de la Société Entomologique de France (N.S.) 23: 67–75. Paschalidis, K.M. 1974b. The identity of two species in the ge-
Ochi T., Kon M. & Bai M. 2009. Three new species of Copris nus Sisyphus (Coleoptera: Scarabaeidae). Journal of the Ento-
from China, with description of a new subgenus. The Ento- mological Society of Southern Africa 37: 299–303.
mological Review of Japan 64: 207–216. Paulian, R. 1932. Description d’un nouveau genre et d’une nou-
Ochi, T., Kon, M. & Masumoto, K. 2012. A new species of the velle espèce de coprophage du Tonkin. Bulletin de la Societe
genus Parascatonomus from the Malay Peninsula. Kogane, To- Entomologique de France, Paris 37: 205–206.
kyo 13: 139–142. Paulian, R. 1933. Révision des Coptodactylini. Bulletin de la So-
Olivier, A.G. 1789. Entomologie, ou histoire naturelle des in- ciété Entomologique de France, Paris 38: 67–74.
sectes, avec leurs caractères génériques et spécifiques, leur Paulian, R. 1937. Sur quelques nouveaux onthophagides afric-
description, leur synonymie, et leur figure enluminée. ains et australiens. Arbeiten uber morphologische und taxon-
Coléoptères. Baudouin, Paris 1: 1–190. omische Entomologie aus Berlin-Dahlem, Berlin-Dahlem 4:
Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess,
336–346.
N.D., Powell, G.V.N., Underwood, E.C., d’Amico, J.A.,
Paulian, R. 1939. Trois nouveaux coprophages de Mauritanie.
Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt,
Revue Française d’Entomologie 6: 10–16.
T.F., Ricketts, T.H., Kura, Y., Lamoreux, J.F., Wettengel,
Paulian, R. 1945. III Coléoptères Scarabéides de l’Indochine.
W.W., Hedao, P. & Kassem, K.R. 2001. Terrestrial ecore-
Première partie. Faune de l’Empire Français. Librairie Larose,
gions of the world: a new map of life on Earth. Bioscience
Paris: 225.
51: 933–938.
Paulian, R. 1953. Recherches sur les insectes d’importance bi-
Orbigny, H. d’. 1900. Supplement au synopsis des Ontho-
ologique à Madagascar. XX. Les Scarabaeini de Madagascar.
phagides paléarctique. L’Abeille 29: 289–300.
Mémoires de l’Institut scientifique de Madagascar (E) 3: 24–27.
Orbigny, H. d’. 1902. Mémoire sur les Onthophagides d’Af-
Paulian, R. 1975. Sur quelques Canthonina montagnards de
rique. Annales de la Société Entomologique de France 71:
Madagascar. Annales de la Société Entomologique de France,
1–324.
Orbigny, H. d’. 1904. Onthophagides africains de la collection Paris (N.S.) 11: 221–252.
du Musée Civique de Gênes. Annali del Museo Civico di Sto- Paulian, R. 1986. Quelques modifications dans la nomenclature
ria Naturale “Giacomo Doria”, Genova 41: 253–331. des coléoptères Scarabéides. Nouvelle Revue d’Entomologie
Orbigny, H. d’. 1905. Onthophagides provenant du voyage de (N.S.) 3: 214.
M. Alluaud dans l’Afrique orientale et descriptions d’au- Paulian, R., Cambefort, Y. & Mauchamp, B. 1982. Deux nou-
tres espèces africains nouvelles. Annales de la Société Ento- veaux Pedaria. Bulletin de la Société Entomologique de France,
mologique de France 74: 381–536. Paris 87: 13–15.
Orbigny, H. d’. 1908. Descriptions d’espèces nouvelles Pereira, F.S. 1954. O género Scatonomus Er. (Coleoptera,
d’Onthophagides africains et notes synonimiques. Annales de Scarabaeidae). Revista Brasileira de Biologia 1: 53–78.
la Société Entomologique de France 77: 65–208. Pereira, F.S. & d’Andretta, M.A.V. 1955. Novos escarabeidos e
Orbigny, H. d’. 1910. Collections receuillis par M. Maurice de novas sinonimias (Col. Scarabaeidae). Papéis Avulsos do De-
Rothschild en Abyssinie et dans l’Afrique orientale anglaise. partamento de Zoologia, Secretaria da Agricultura, S. Paulo,
Coléoptères: Onthophages. Bulletin du Musée Nationale Brasil 12: 247–264.
d’Histoire Naturelle, Paris 16: 308–310. Péringuey, L. 1885. First contribution to the South-African Co-
Orbigny, H. d’. 1913. Synopsis des Onthophagides d’Afrique. leopterous Fauna. Transactions of the South African Philosoph-
Annales de la Société Entomologique de France 82: 1–742. ical Society 3: 74–149.
Orbigny, H. d’. 1915. Synopsis d’un genre nouveau d’Oniticel- Péringuey, L. 1888. Second contribution to the South-African
lides special a Madagascar. Annales de la Société Entomologique Coleopterous Fauna. Transactions of the South African Philo-
de France, Paris 84: 402–434. sophical Society 4: 67–194.
Palestrini, C. 1982. Il “sottogenere” Pseudonthophagus Balth. Péringuey, L. 1892. Third contribution to the South-African
(Coleoptera : Scarabaeidae : Onthophagini). Bollettino della Coleopterous Fauna. Transactions of the South African Phil-
Società Entomologica Italiana 114: 97–102. osophical Society 6: 1–94.
Palestrini, C. 1992. Systematica é zoogeografica del genere Péringuey, L. 1896. Descriptions of new genera and species of
Onthophagus sottogenere Proagoderus Lansberge. Memoirie Coleoptera from South Africa, chiefly from Zambezia. Trans-
della Società Entomologica Italiana, Genova 71: 1–358. actions of the Entomological Society of London 2: 149–189.
SURICATA 6 (2020) 777

Péringuey, L. 1901. Descriptive catalogue of the Coleoptera of Roets, F., Crous, C. & Pryke J. 2017. Sarophorus diabolus sp.
South Africa. Transactions of the South African Philosophical n. and Sarophorus frolovi sp. n. (Coleoptera: Scarabaeidae:
Society 12: 1–563. Scarabaeinae) from South Africa. African Entomology 25:
Péringuey, L. 1908. Descriptive catalogue of the Coleoptera of 264–270.
South Africa. Additions and corrections. Transactions of the Roggero, A., Barbero, E. & Palestrini, C. 2015. Phylogenetic
South African Philosophical Society, Cape Town (1907) 13: and biogeographical review of the Drepanocerina. Arthropod
547–752. Systematics & Phylogeny 73: 153–174.
Perty, J.A.M. 1830. Insecta Brasiliensia. In: Spix J, Martius C: Roggero, A., Barbero, E. & Palestrini, C. 2016. Revised classifi-
Delectus animalium articulatorum, quae in itinere per Brasil- cation and phylogeny of an Afrotropical species group based
iam annis MDCCCXVII-MDCCCXX jussu et auspiciis on molecular and morphological data, with the descrip-
Maximilliani Josephi I. Bavariae regis augustissimi peracto. tion of a new genus. Organisms Diversity & Evolution DOI
Frid. Fleischer, Monachii [München]: 1–60. 10.1007/s13127-016-0297-z: 1–18.
Philips, T.K. 2016. Phylogeny of the Oniticellini and Ontho- Roggero, A., Dierkens, M., Barbero, E. & Palestrini, C. 2016.
phagini dung beetles from morphological evidence. ZooKeys Combined phylogenetic analysis of two new Afrotropical
579: 9–57. genera of Onthophagini. Zoological Journal of the Linnean
Philips, T.K. & Bell, K. 2008. Attavicinus, a new generic name Society doi: 10.1111/zoj.12498: 1–23.
for the myrmecophilous dung beetle Liatongus monstrosus. Roggero, A., Moretto, P., Palestrini, C. & Barbero, E. 2019. The
The Coleopterists Bulletin 62: 67–81. phylogenetic relationships of Tiaronthophagus n.gen. (Cole-
Philips, T.K. & Scholtz, C.H. 2000. A new genus and species optera, Scarabaeidae, Onthophagini) evaluated by phenotyp-
of trichome-bearing dung beetle from South Africa. African ic characters. Insects 10: 3390.
Entomology 8: 227–231. Rossi, P. 1792. Mantissa Insectorum, exhibens species nuper in
Pokorný, S. & Zídek, J. 2009. Review of the Gymnopleurini. Etruria collectas, adiactis Faunae Etruscae illustrationibus, ac
I. Introduction and the genus Allogymnopleurus. Folia Hey- amendationibus (1792–1794). Typographia Polloni. Pisis 1:
rovskiana Series A 17: 133–149. 1–154.
Pokorný, S. & Zídek, J. 2015. Checklist of Kheper Janssens and Roth, J.R. 1851. Diagnosen neuer Coleoptera aus Abyssinien.
description of a new species from northern Tanzania. Insecta Archiv für Naturgeschichte, Berlin 17: 115–133.
Mundi 0429: 1–7.
Rougon, D. & Rougon, C. 1982. Le comportement nidificateur
Pokorný, S., & Zídek, J. 2018. Review of the Gymnopleurini
des Coléoptères Scarabaeinae Onilicellini en zone sahélienne.
(Coleoptera: Scarabaeidae: Scarabaeinae). IV. African species
Bulletin de la Société Entomologique de France 81: 272–279.
of Garreta Janssens, with description of a new species from
Sahlberg, C.R. 1823. Periculi Entomographici, species Insecto-
Guinea. Folia Heyrovskiana Series A 26: 105–109.
rum nondum descriptas propositour. Fasciculus, Insecta Fen-
Pokorný, S., Zídek, J. & Werner, K. 2009. Giant dung beetles
nica, 1–82.
of the genus Heliocopris. Taita Publishers, Czech Republic:
Schäfer, P. & Fischer, E. 1992. Zur Scarabaeiden-Fauna Rwan-
1–135.
das, Zentralafrika. 1. Sisyphini. Senckenbergiana biologica 72:
Preudhomme de Borre, A. 1880. Note sur le genre Macroderes
119–137.
Westwood. Annales de la Societe Entomologique de Belgique
Schäfer, P. & Fischer, E. 2001. Zur Scarabaeidenfauna Rwan-
23: 7–11.
das, Zentralafrika 2, Gymnopleurini und nachtrag Sisyphini.
Quedenfeldt, G. 1884. Verzeichniss der von Herrn Major a D
von Mechow in Angola und am Quango-strom 1878–1881 Mitteilungen des internationalen entomologischen Vereins sup-
gesammelten Pectinicornen und Lamellicornen. Berliner en- plement 9: 1–93.
tomologische Zeitschrift 28: 265–340. Scheuern, J. 1988. Zur taxonomie der gattung Liatongus mit
Raffray, A. 1877. Coléoptères Lamellicornes rapportés par M.A. beschreibung einer neuen art aus Bhutan. Entomologica Basil-
Raffray d’Abyssinie et Zanzibar. Descriptions des espèces iensia 12: 305–317.
nouvelles. Revue et Magazine de Zoologie, Paris 5: 312–336. Scholtz, C.H. & Howden, H. 1987. A revision of the African
Reiche, L.J. 1841. Tableau d’une division systématique de la tri- Canthonina. Journal of the Entomological Society of Southern
bu des Coprophages dans la famille des Lamellicornes. Revue Africa 50: 75–119.
Zoologique 4: 211–213. Scholtz, C.H. & Howden, H. 1987. A revision of the southern
Reiche, L. 1848. Ferret A. & Galinier: Voyage en Abyssinie dans African genus Epirinus. Journal of the Entomological Society of
les provinces du Tigre, du Samen et de l’Amhara (1839- Southern Africa 50: 121–154.
1843). Tome I–III (1847–1848). Entomologie. Paris (1849) Schoolmeesters, P. 2017. Scarabs: World Scarabaeidae Database
3: 304–361. (version Jan 2017). In: Species 2000 & ITIS Catalogue of
Reitter, E. 1893. Bestimmungstabelle der Lucaniden und co- Life, 29th May 2017. Digital resource at www.catalogueofli-
prophagen Lamellicornen. XXIV. Heft, (Sonderabdruck aus fe.org/col. Species 2000; Roskov Y, Abucay L, Orrell T, Nic-
dem XXX. Bande der Verhandlungen des naturforscheden olson D, Bailly N, Kirk PM, Bourgoin T, DeWalt RE, De-
Vereins in Brünn). Verlag des Verfassers, Brünn: 3–230. cock W, De Wever A, Nieukerken E van, Zarucchi J, Penev
Ritsema, C. 1888. Lijst der entomologische Geschriften van Mr. L (eds) (2017); Naturalis, Leiden, the Netherlands. ISSN
J.W. van Lansberge, gevolgd door eenen opgawe der daarin 2405-8858.
beschreven nieuwe geslachten, ondergeslachten en soorten. Schreber, J.C.D. 1759. Novae species Insectorum. Halae,
Tijdschrift voor Entomologie 31: 201–234. Schneider: 1–16.
Roets, F. & Oberlander, K.C. 2010. Silvaphilus: a new relict Sharp, D. 1880. Note sur quelques espèces du genre Macroderes.
forest-endemic Canthonini dung beetle genus from the Annales de la Société Entomologique de Belgique 23: 36–39.
Western Cape Province of South Africa. African Entomology Shibuya, T. & Inouchi, J. 1982. Fine structure and odor re-
18: 369–373. sponses of olfactory sensillae of the Japanese dung beetle,
778 SURICATA 6 (2020)

Copris pecuarius (Coleoptera: Scarabaeidae). Japanese Journal formen. Entomologische Blätter für Biologie und Systematik der
of Applied Entomology and Zoology, 26: 194–195. Käfer 63: 129–173.
Shipp, J.W. 1895a. The coprophagous Lamellicornia: a list of Strassen, R. zur. 1980. A new species of Scarabaeus Linnaeus,
species belonging to the genera Circellium and Sceliages, with 1758 from coastal sand areas in Natal. Journal of the Entomo-
notes on Ateuchidae. The Entomologist 28: 37–40. logical Society of Southern Africa 43: 107–112
Shipp, J.W. 1895b. Gymnopleurus; correction of nomenclature. Tagliaferri, F., Moretto, P. & Tarasov S. 2012. Essai sur la sys-
The Entomologist 28: 2–3. tématique et la phylogénie des Onthophagus Latreille, 1802,
Shipp, J.W. 1895c. Notes on Onthophagus Latr., with correc- d’Afrique tropicale appartenant au septième groupe de d’Or-
tions of nomenclature, and a description of a new genus. The bigny. Description d’un sous-genre nouveau et de trois es-
Entomologist 28: 178–179. pèces nouvelles. Catharsius, La Revue 6: 1–31.
Shipp, J.W. 1895d. Onthophagus: correction in Nomenclature. Takano, H. 2018. A systematic revision of the Afrotropical
The Entomologist 28: 221–222. members of the dung beetle genus Catharsius Hope, 1837
Shipp, J.W. 1895e. A revised classification of the genus Ateuchus (Coleoptera: Scarabaeidae). D. Phil. thesis, University of Ox-
Weber. The Entomologist 28: 218–221. ford.
Shipp, J.W. 1896a. Irrorhotides: a new genus of Ateuchidae, with Tarasov, S.I. & Dimitrov, D. 2016. Multigene phylogenetic
a description of a new species. The Entomologist 29: 116– analysis redefines dung beetles relationships and classifica-
4117. tion. BMC Evolutionary Biology 16: 1–19.
Shipp, J.W. 1896b. On a new species of Ateuchus (Actinophorus) Thomson, C.G. 1859. Skandinaviens Coleoptera. Synoptiskt
from Australia. Novitates Zoologica 3: 72. bearbetade. I. Tom. Berlingska Boktryckeriet, Lund: 290.
Shipp, J.W. 1897a. I. Coleoptera by Dr Karl Jordan. Appendix Thunberg, C.P. 1818. Coleoptera Capensia antennis lamellatis,
1. Scarabaeidae. In: A. Donaldson Smith, Through unknown sive clava fissili instructa. Memoires de l’Academie imperiale
African countries: the first expedition from Somaliland to lake des Sciences de Saint Petersburg 6: 395–450.
Lamu. E. Arnold: London, New York: 448–452. Tshikae, B.P., Davis, A.L.V. & Scholtz, C.H. 2013a. Dung bee-
Shipp, J.W. 1897b. On the genus Gymnopleurus, Illiger; with a tle assemblage structure across the aridity and trophic re-
list of species and descriptions of two new genera. The Ento- source gradient of the Botswana Kalahari: patterns and driv-
mologist 30: 62–66, 131–135, 166–168. ers at regional and local scales. Journal of Insect Conservation
Silvestri, F. 1924. Description of a new myrmecophilous 17: 623–636.
Scarabaeidae of India. Records of Indian Museum 26: 583– Tshikae, B.P., Davis, A.L.V. & Scholtz, C.H. 2013b. Does an
586. aridity and trophic resource gradient drive patterns of dung
Simonis, A. & Zunino, M. 1980. Il genere Cyptochirus. Bollet- beetle food selection across the Botswana Kalahari? Ecological
tino del Museo di Zoologia dell’Università di Torino 4: 21–40. Entomology 38: 83–95.
Smith, A.G. 2006. A review of the family-group names for the Tyson, P.D. 1986. Climatic change and variability in southern Af-
superfamily Scarabaeoidea with corrections to nomenclature rica. Oxford University Press, Cape Town.
and a current classification. Coleopterists Society Monograph Vaz de Mello, F.Z. 2008. Synopsis of the new subtribe Scatimi-
5: 144–204. na, with descriptions of twelve new genera and review of Ge-
Smith, A.G. 2009. Checklist and nomenclatural authority file of nieridium, new genus. Zootaxa 1955: 3–75.
the Scarabaeoidea of the Nearctic realm. Version 4 – released Vinson, J. 1946. On Nesosisyphus, a new genus of coprine beetles
22 April 2009. Available at http://museum.unl.edu/research/ from Mauritius. Proceedings of the Royal Entomological Society
entomology/SSSA/Nearctic-Scarabs4.pdf. of London 15: 89–96.
Spector, S. 2006. Scarabaeine dung beetles (Coleoptera: Wall, R. & Beynon, S. 2012. Area-wide impact of macrocyclic
Scarabaeidae: Scarabaeinae): an invertebrate focal taxon for lactone parasiticides in cattle dung. Medical and Veterinary
biodiversity research and conservation. The Coleopterists Bul- Entomology 26: 1–8.
letin Monograph Number 5: 71–83. Wallengren, H.D.J. 1881. Coleoptera Transvaliensia. Bidrag till
Steven, C. von 1809. Descriptions de quelques insectes du Cau- kännedora où Transvaal-Landets I S. Africa Coleopter-fauna.
case et de la Russie méridionale. Mémoires de la Société Impe- Entomologisk Tidskrift, Stockholm 2: 9–22.
riale des Naturalistes de Moscow 2: 31–42 Walter, P.H. 1976. Un nouveau Metacatharsius du Zaire et de
Strassen, R. zur. 1961a. Eine neue Scarabaeus art aus Südafrika. République Populaire du Congo. Bulletin de la Société Ento-
Senckenbergiana Biologica 42: 437–441. mologique de France 81: 267–270.
Strassen, R. zur. 1961b. Einige seltene Sudafrikanische Scarabae- Walter, P.H. 1982. Contribution à la connaissance des
us arten im Mus. Frey mit einer neu-beschreibung. Entomolo- scarabaeides corprophages du Gabon. I. Trois espèces nou-
gische Arbeiten aus dem Museum G. Frey 12: 236–241. velles de la région de Makokou. Revue francaise d’Entomolo-
Strassen, R. zur. 1962. Beschreibung einiger neuer oder verkann­ gie, Paris (N.S.) 4: 167–171.
ter südafrikanischer Scarabaeus-Arten. Senckenbergiana Bio- Walter, P.H. 2014. Les Caccobius Thomson, 1859, appartenant
logica 43: 339–357. au couplet 65/12 du ‘Synopsis’ de d’Orbigny. CATHARSIUS
Strassen, R. zur. 1963. Scarabaeus andreaei, ein neuer Pillendre- La Revue 10: 9–34.
her aus Mozambique. Senckenbergiana Biologica 44: 107– Walter, H. & Lieth, H. 1964. Klimadiagramm-Weltatlas. Jena:
110. Gustav Fischer.
Strassen, R. zur. 1965. Zur Kenntnis Gattung Sceliages West- Wasmann, E. 1918. Myrmecophile und termitophile coleop-
wood, mit zwei Neubeschreibungen. Senckenbergiana Biolog- teren aus Ostindien. Wiener Entomologische Zeitung 37: 1–23.
ica 46: 219–232. Waterhouse, C.O. 1876. On various new genera and species of
Strassen, R. zur. 1967. Arten ubersicht der gattung Scarabaeus Coleoptera. Transactions of the Royal Entomological Society
Linnaeus mit besonderer berucksichtigung der Athiophischen Fourth Series: 11–25.
SURICATA 6 (2020) 779

Waterhouse, C.O. 1888. On some Coleoptera from eastern Wiedemann, C.R.W. 1823. Zweihundert neue Käfer von Java,
Equatorial Africa received from Emin Pasha. Proceedings of Bengalen, und dem Vorgebirge der guten Hoffnung. Akade-
the Zoological Society of London, 86–87. mische Buchhandlung: Kiel Zoologische Magazin Vol. 2, Part
Waterhouse, C.O. 1890. XLVII. New Scarabaeidae in the Brit- 1: 1–133.
ish Museum. The Annals and Magazine of natural History, Wiedemann, C.R.W. & Germar, E.F. 1821. Neue exotische
including Zoology, Botany and Geology, London 6: 365–373. Käfer. Magazin der Entomologie, herausgegeben von Dr. Ernst
Waterhouse, C.O. 1891. New Scarabaeidae in the British Mu- Germar. J. Chr. Hendel und Sohn, Halle 4: 107–183.
seum: a fifth contribution. The Annals and Magazine of nat- World Wildlife Fund. 2017. Terrestrial Ecoregions. Available
ural History, including Zoology, Botany and Geology, London at: https://www.worldwildlife.org/biome-categories/terrestri-
6: 507–522. al-ecoregions.
Weber, F. 1801. Observationes Entomologicae, Continentes Ziani, S. & Branco, T. 2011. The authorship and date of the
Novorum Quae Condidit Generum Characteres, et Nuper genus-group name Pachylomera. Zootaxa 2803: 63–68.
Detectarum Specierum Descriptiones. Bibliopolii Academici Ziani, S. & Gudenzi, I. 2012. Commenti sulla sistematica ge-
Novi, Kiel, Germany. nerica degli Scarabaeini del bacino del Mediterraneo con una
Westwood, J.O. 1837a. Partington C.F. British Cyclopaedia chiave dicotomica per il loro riconoscimento (Insecta Cole-
of Natural History: Combining a scientific classification of optera Scarabaeidae: Scarabaeinae). Quaderno Di Studi E No-
animals, plants and minerals; with a popular view of their tizie Di Storia Naturale Della Romagna 36: 115–158.
habits, economy, and structure. Orr & Smith London 2: 55. Zídek, J. & Pokorný, S. 2004. Checklist of the genus Scarabaeus
Westwood, J.O. 1837b. Characters and descriptions of some Linné. Animma. X 5: 1–30.
new coleopterous insects, belonging to the family of sacred Zídek, J. & Pokorný, S. 2008. Illustrated keys to Palearctic
beetles. Proceedings of the Zoological Society of London 5: 12– Scarabaeus Linné (Scarabaeidae). Animma. X 27: 1–27.
13. Zídek, J. & Pokorný, S. 2011. Replacement name for a subgenus
Westwood, J.O. 1839. Descriptions of insects figured in plate of Scarabaeus Linné, and remarks on Scarabaeus isidis. Klapa-
9 and 10. Royle J.F.: Illustration of the Botany and other lekiana 47: 89–90.
Branches of the Natural History of the Himalayan Moun- Zídek, J. & Pokorný, S. 2018. New species of the Scarabaeus
tains and of the Flora of Cashmere. I. London 7: 53–55. subgenus Scarabaeolus Balthasar, with a review of the subge-
Westwood, J.O. 1842. Descriptions of some new exotic genera nus (Scarabaeidae: Scarabaeinae: Scarabaeini). Insecta Mundi
belonging to the family of the sacred beetles. Proceedings of 0611: 1–35.
the Royal Entomological Society of London: 59. Zunino, M. 1977. Una nuova specie di Neopachysoma della Na-
Westwood, J.O. 1844. A memoir containing descriptions of two mibia. Bollettino del Museo di Zoologia dell’Università di To-
species of sacred beetles from southern Africa. Proceedings of rino 2: 15–18.
the Entomological Society of London (1841), 100. Zunino, M. 1979. Gruppi artificiali e gruppi naturali negli
Wiedemann, C.R.W. 1819. Neue Käfer aus Bengalen und Java. Onthophagus. Bollettino del Museo di Zoologia dell’Università
Zoologisches Magazin, Kiel 1: 157–183. di Torino 1: 1–18.
780 SURICATA 6 (2020)

Glossary of scientific terms

The terms cited below are defined only in the context of Allotype: See ‘Type specimens’; a secondary type often
the present book. Definitions may not be universally ap- created if males and females differ markedly in appearance.
plicable. They have been simplified as far as possible for
readers who have not received a training in scientific ter- Alluvial soils: Soils deposited in riverine (rivers) or lacus-
minology. trine (lake) environments.

Alpha taxonomy: The most basic level of taxonomy, i.e.

5×5 km polygon: The spatial resolution of information description of animals and plants at species level.
(size of squares) on the GIS base maps from which av-
erage altitude, temperature and rainfall data were ex- Area of Occurrence (AOO): The area actually occupied
tracted. by a species (also see ‘Extent of Occurrence’).

A Assemblage: Refers to a co-occurring group of species;

species assemblage structure differs geographically, be-
Abdomen: The hind portion of the beetle body. tween habitats and between dung types.

Aedeagus: Sclerotised (hardened) reproductive organ of Asymmetrical: An irregular pattern.

male dung beetles, which may be similar or quite different
in shape between species; sometimes aedeagi (plural) are Auctorum: ‘of authors’; conveys that a species name is
extracted by dissection to confirm the identity of species. used in the sense of subsequent authors, which differs to
that intended by the original author.
Afrotropical: Major biogeographical region comprising
Africa south of the central Sahara Desert; sometimes con- Australasia: Major biogeographical region comprising
sidered to include Madagascar and southwest Arabia. Australia, New Zealand, New Guinea and nearby islands.
SURICATA 6 (2020) 781

Avermectins: A group of pesticides used to control pests used as an inexact type locality for species described in the
of domestic livestock; they are derived as fermentation earlier part of the 19th century.
products from actinomycete bacteria.
Carnivore: In reference to mammals: meat-eating species
dropping small amounts of dung that contain high levels
B of nitrogenous compounds; little known on its relative at-
traction to dung beetles.
Ball roller: One of four major behavioural groups in
scarabaeine dung beetles; species that construct and roll Clade: A branch on an evolutionary tree; may comprise
balls of dung (also see ‘Endocoprid’, ‘Kleptocoprid’, ‘Tun- one or more evolutionary lineages.
neller’).
Cladistic: Evolutionary analysis; subdivision into clades
Basal dichotomies: Lower and older levels of branching (see ‘Clade’).
on an evolutionary tree.
Classification: A hierarchical system that classifies the
Basally derived: Lineages derived from nodes at the base Class Insecta (insects) into orders, suborders, superfami-
of an evolutionary tree (also see ‘Basal dichotomies’). lies, families, subfamilies, tribes, subtribes, genera, subge-
nera, species, subspecies, down to varieties.
Bioassay: Use of dung beetles to determine the relative
non-target toxicity of different pesticides used on domes- Clay: The finest-grained division in the United States
tic livestock; assessment based on percentage mortality or Department of Agriculture (USDA) classification of soil
percentage reduction in breeding success due to pesticide types (see ‘Soil classification’).
residues in dung.
Cline: A range of variation within a species that matches
Biological indicators: Differences in the occurrences of a geographical gradient, e.g. exoskeleton colour change
dung beetle species are widely used to indicate effects of across a gradient from warmer to cooler temperatures.
changes in vegetation cover (habitat fragmentation) and
effects of different pesticides in agricultural landscapes Coleoptera: The taxonomic Order to which beetles be-
(bioassay). long.

Biomass: The ecological impact of species varies accord- Collecting intensity: The relative effort expended on
ing to their different body sizes; therefore, assemblages of collecting insects, which frequently varies from region to
organisms may be compared by weight, which is termed region; such variation may introduce bias into recorded
biomass. distribution patterns and needs to be recognised.

Biome: Largest of three geographical scales used to define Collection artefact: Used in explanation when the dis-
floral distribution patterns in South Africa by Mucina and tribution of a species is poorly known and is suspected
Rutherford (2006) (also see ‘Bioregion’ and ‘Vegetation to represent under-collection rather than a true pattern
unit’). of rarity or restricted occurrence related to some environ-
mental factor.
Bioregion: Intermediate geographical scale out of three
used to define floral distribution patterns in South Africa Conspecifics: Individuals occurring in the same species.
by Mucina and Rutherford (2006) (also see ‘Biome’ and
‘Vegetation unit’). Convergence: See ‘Morphological convergence’.

Bootstrap: Statistical method for defining the strength of Correlation: See ‘Correlated’.
a predicted relationship between lineages diverging from
a node on an evolutionary tree; usually expressed as a per- Correlated: A relationship between different factors, e.g.
centage likelihood. between annual rainfall and the occurrence of a species.

Cover density: The degree by which vegetation covers the

C soil surface or by which the tree canopy shades the soil
surface; usually expressed as a percentage.
Caffraria: An old and inexact term considered to repre-
sent southeast southern Africa in its narrowest sense up to Critically Endangered (CR): One of the Red List cate-
the whole of southern Africa in its widest sense; frequently gories of threat designated by the IUCN; used to identify
782 SURICATA 6 (2020)

a species that is considered to face threats that are only Ectoparasites: External parasites of animals.
one level above Extinction (EX) in the wild (see IUCN
categories in Introduction). Elytra: The sclerotised (hardened) and modified fore
wings of beetles that cover the functional hind wings
when they are folded over the dorsal surface of the abdo-
D men; may be fused together in flightless species.

Data: General term used to refer to any information Endangered (EN): One of the Red List categories of
whether descriptive or quantitative. threat designated by the IUCN; used to identify a species
that is considered to face serious threats of extinction in
Data Deficient (DD): One of the Red List categories of the wild (see IUCN categories in Introduction).
threat designated by the IUCN; used to identify a species
for which information is insufficient to assess whether or Endemic: Found only within a specified region or situa-
not it faces any threats of extinction in the wild. tion and nowhere else.

Dentition: Tooth-like projections on the exoskeleton of Endocoprid: One of four major behavioural groups in
insects that is often used in taxonomy and classification. scarabaeine dung beetles; species that breed within or im-
mediately below droppings in situ (also see ‘Ball roller’,
Diel periodicity: Daily period of activity; either walking
‘Kleptocoprid’, ‘Tunneller’).
in flightless species or flying in winged species; may be
diurnal (daytime) or active in darkness (nocturnal: night Endoparasites: Internal parasites of animals.
time; crepuscular: dusk and dawn, dusk only, or, rarely,
dawn only). Exoskeleton: The often hard outer layer of the insect
body.
Digital databasing: Computerised listing of information
from labels attached to museum specimens is now wide- Extant: Existing at the present time, i.e. not extinct.
spread and forms a large part of the field of study termed:
Bioinformatics. Extent of Occurrence (EOO): The area enclosed by
outlining distribution records for a species; if species oc-
Disjunct: A geographical distribution characterised by cupancy is continuous, it will be equivalent to Area of
gaps between Areas of Occurrence (AOO). Occurrence (AOO); if species occupancy is disjunct or
non-continuous, EOO will be larger than AOO.
Diurnal: Activity during the hours of daylight.

Diversity: An often loosely used term referring to the F

scale of differences within a region or an assemblage of
organisms (high or low diversity – in numerical terms = Family: See ‘Classification’.
large numbers or low numbers).
Flight periodicity: See ‘Diel periodicity’.

E Fossorial: Associated with the soil.

Ecological associations: General term used to refer to Frons: From above: the middle part of the head between
associations with the environment at large (e.g. biogeog- the clypeus at the front, the vertex at the back and the
raphy) or small scales (e.g. habitat, food). genae at the sides.

Ecological Impact Assessments (EIA): Studies designed Fungi: Classified at Kingdom level along with plants and
to determine the potential effects of development; for in- animals; includes single up to multicellular organisms;
stance if development would be ecologically detrimental some dung beetles show specialist associations with mush-
or if an area earmarked for development contains threat- rooms and toadstools on which they feed and breed.
ened species.

Ecoregions: Geographical regions defined by Olson et al. G

(2001) from patterns of plant and animal distribution.
Used here only for areas outside the borders of South Africa. Generalisation: See ‘Generalist’.
SURICATA 6 (2020) 783

Generalist: A species that shows a wide range of ecologi- Incertae sedis: ‘of uncertain placement’. Used in this
cal associations without being specialised to any particular book when genera have been placed in a tribe, but proba-
factor e.g. found on a range of soil types from sand to bly do not belong there.
sandy loam to clay with no specialisation to any particular
one (see ‘Specialist’). Indument: A natural covering; in the context used here,
material covering the lateral edges of the elytra.
Genus: Singular of genera; a level of classification of
plants and animals (see ‘Classification’). Ingestables: In reference to pesticides administered to
domestic livestock orally; toxic pesticide residues are sub-
GIS base maps: Digital Geographical Information System sequently found in dung.
base maps divided into small squares (polygons) from which
computer information may be extracted; in this instance Injectables: In reference to pesticides administered to do-
maps for annual rainfall, annual temperature, altitude. mestic livestock by injection; toxic pesticide residues are
subsequently found in dung.
Gradsect: A study comprising data obtained from points
along a geographical gradient. Intraspecific: Variation within a species.

Grain size profiles: The proportional composition of a Iridescent: Often shiny metallic colouration shown by
soil in terms of three different grain sizes (see ‘Soil clas- many dung beetles, which is generated by reflectance of
sification’). different wavelenths of light from layers in the exoskel-
eton; different thicknesses of the reflective layers are re-
H sponsible for the different perceived blue, green or cupre-
ous colours.
Habitat: Characteristics of an area where an animal is
found, e.g. sandy desert, forest, grassland. IUCN Red List Categories: Categories to describe the
relative threat or lack of threat to survival of species des-
Habitat fragmentation: The process of subdivision of ignated by the International Union for Conservation of
natural habitat, primarily through urbanisation, agricul- Nature (IUCN).
tural or industrial development.

Habitat heterogeneity: The relative diversity of habitats K

in a geographical area.
Kleptocoprid: One of four major behavioural groups in
Habitat transformation: The modification of natural scarabaeine dung beetles; species that ‘steal’ dung buried
habitat, e.g. conversion of highland grassland to commer- by other dung beetle species for breeding purposes (also
cial tree plantation in South Africa. see ‘Ball roller’, ‘Endocoprid’, ‘Tunneller’).
Holotype: See ‘Type specimens’; A single type specimen
designated to represent the name allotted to a species; may
be supported by additional designated types with the sta-
L
tus of allotype or paratypes.
Later-derived: See ‘Terminally derived’.
Hydrological: Pertaining to water in soils.
Least Concern (LC): One of the Red List categories of
Hyrax middens: The toilet areas (middens) used by the threat designated by the IUCN; used to identify a species
rock hyrax, which contain a mass of dung pellets. that is considered to face no threats of extinction in the
wild.

I Lectotype: See ‘Type specimens’; a single specimen select-

ed from a type series containing only syntypes; thereafter,
Improved pasture: Agricultural term referring to the im- essentially has the status of a holotype.
provement of pasture so that it is more suitable for farm-
ing domestic livestock, particularly cattle; natural grass- Lineage: Refers to a single evolutionary line or collection
land is often ‘improved’ through replacement by Kikuyu of branched lines originating from a single point in a phy-
grass pasture. logeny.
784 SURICATA 6 (2020)

Localised: Refers to a small Area of Occurrence (AOO usually genetic material found in animal or plant cells
= actual presence) within a large Extent of Occurrence (DNA sequences).
(EOO = the overall range).
Monogastric herbivores: In reference to mammals: graz-
Localisation: see ‘Localised’. ing or browsing herbivores in which there is a single stom-
ach; dung usually coarse-fibred, e.g. elephants, horses.

M Monophyletic: Refers to a taxon derived from a single

ancestor; monophyletic ancestry may be supported by oc-
Macrohabitat: Large scale spatial associations, e.g. forest currence on a single evolutionary lineage.
or desert.
Monotypic: A genus which is represented by a single spe-
Major females: In some dung beetle species, females also cies.
show secondary sexual armature that varies in prominence
with body size so that it is greater in larger-bodied indi- Morphological phylogeny: Evolutionary tree based on
viduals than in smaller-bodied individuals; however, one analyses of similarity between characteristics of body
rarely sees the terms major or minor females (see ‘Sexual shape and body features.
dimorphism’).
Morphology: Relating to the shape and sculpturing of
Major males: In many dung beetle species, the promi- insect exoskeletons; morphological variation is used to
nence of secondary sexual armature is greater in larger- classify insects (see ‘Classification’).
bodied individuals (termed major males) than in smaller-
bodied individuals (termed minor males) (see ‘Sexual di- Morphological: See ‘Morphology’.
morphism’, ‘Secondary sexual armature’).
Morphological convergence: Sometimes seen in distant-
Means: Essentially average values, but termed means be- ly related species in separate geographical areas that have
cause they are cited together with a standard deviation independently evolved similar body shape due to similar
(SD); SDs merely represent a statistical range within habits.
which most data values occurred; the SD also conveys
whether there was little variation in data values (small Multilayer interference: See ‘Iridescent’.
plus or minus deviation in value away from the mean)
or a great deal of variation (large deviation in value away Mycetophage, Mycetophagous: Feeding on fungi.
from the mean).

Mesic: A relative term for moister as opposed to drier or N

arid conditions.
Nearctic: Major biogeographical region comprising
Microhabitat: Small scale spatial associations; e.g. in leaf North America southwards into higher altititude areas of
litter, under stones. Mexico.

Milbemycins: A group of pesticides used to control pests Near Threatened (NT): One of the Red List categories of
of domestic livestock that are chemically similar to aver- threat designated by the IUCN; used to identify a species
mectins; they are derived as fermentation products from that is considered to be close to facing threats of extinc-
Streptomyces bacteria. tion in the wild (see IUCN categories in Introduction).

Minimum spanning trees: Method used to statistically Neotropical: Major biogeographical region comprising
link species data points in the NMDS ordinations; these South America northwards through central America into
links are depicted by a network of lines joining data points lowland parts of Mexico.
for species that showed the most similar distribution pat-
terns to one another in southern Africa. Neotype: See ‘Type specimens’; a single specimen des-
ignated to represent a species name when no previously
Minor females: (see ‘Major females’). designated type can be located, often through loss or de-
struction.
Minor males: (see ‘Major males’).
Nest architecture: Immature dung beetles develop within
Molecular phylogeny: Evolutionary tree based on analy- modelled portions of dung, termed broods, that are con-
ses of similarity between molecular level characterisation; structed by the parent(s), mostly within nests in the soil;
SURICATA 6 (2020) 785

the manner of dung burial, brood and nest construction Phylogeny: Evolutionary tree defining relationships be-
varies between taxa; it can be simple or complex; the char- tween lineages; in this instance, tribes, genera or species
acteristics of a nest may be described as nest architecture. of dung beetles.

NMDS ordination: Non-metric Multidimensional Scal- Polyphyletic: Refers to a taxon with members that occur
ing ordination: a statistical method used, here, to sum- in different evolutionary lineages; indicates that the taxon
marise average similarities or dissimilarities between the is derived from more than one ancestor; also demonstrates
geographical distributions of species in southern Africa that as constituted, at least some included taxa have been
(represented by data points in a two-dimensional box); misclassified.
the distance between data points provides a rough indica-
tion of how close or distant the distribution patterns are Post-mining chronosequence: In reference to mining of
for each species or tribe. dunes for titanium at Richards Bay; refers to the succes-
sion of vegetation (grassland, shrubland, woodland) that is
Node: In relation to evolutionary trees, refers to the point restored in sequence following mining of the dune forest.
of branching between lineages.
Pour-ons: Refers to the administering of pesticides poured
Nomenclature: Refers to the names used in the classifi- onto the exterior of domestic livestock to control pests;
cation system. toxic pesticide residues are subsequently found in dung.

Nom. nov.: At species level, a new name to replace one Prothoracic disc: The visible upper part (= pronotal disc)
that represents duplication of the same name for a species of the thorax (middle section of a dung beetle between the
in the same genus described at an earlier date. head and hind part).

Psammophilous: Sand-associated.
O Pyrethroids: A group of manufactured pesticides used
Olfaction: The sense of smell by which dung beetles lo- to control pests of domestic livestock that are chemically
cate dung; the olfactory sensory cells are found on the an- similar to pyrethrins produced naturally by the plant ge-
tennae (feelers). nus, Pyrethrum; toxic pesticide residues are subsequently
found in dung.
Omnivores: In reference to mammals: a mixed animal
and plant diet; omnivore dung is often found to attract
more species and individuals than other dung types.
Q
Qualitative data: Descriptive data based on observations
Order: See ‘Classification’.
or on numerical data that are not supported by a stan-
Oriental: Major biogeographical region comprising the dardised method.
more tropical southern regions of southeast Asia, includ-
Quantitative data: Numerical data derived from a stan-
ing most of Indonesia.
dardised method of collection.

P R
Palaearctic: Major biogeographical region comprising Radiant heat: Heat projected directly from the sun.
Europe, Northern Asia, North Africa and the Middle East.
Range fragmentation: Fragmentation of a species range
Paratype: See ‘Type specimens’; additional designated through piecemeal habitat modification, or, in the case
specimens in a type series; often designed to show the of dung beetles, possibly through range contraction by
range of variation within a species or to act as alternative mammals dropping preferred dung types.
types if the holotype is lost or destroyed.
Raw data: Data (usually numerical) that has not been an-
Pars: One part of a type series that contains more than a alysed further from that first recorded.
single species in error.
Reference material: Stored museum specimens; informa-
Phylogenetic: Pertaining to phylogeny. tion on labels attached to validated reference material are
786 SURICATA 6 (2020)

used to determine species geographical distributions and are presence or absence of horns or projections on the head
ecological associations. or prothoracic disc, also, presence or absence of spines
on the legs; these features vary in prominence with body
Regression trees: A statistical predictive modelling tech- size from, so-called, major to minor features; graphs have
nique that was used to classify the South African flora into shown that the variation describes a sigmoid curve with
Vegetation Units, Bioregions and Biomes (see Mucina & change from major to minor features over a relatively small
Rutherford 2006). range in body size; the characters of major males decline in
prominence to minor males, which are close in appearance
Revalidation: Raising the status of taxa from synonymy to minor females, whose characters may increase in prom-
with other names back to fully valid genus or species status. inence with body size to those shown by major females.

Revision: Refers to a review of previously described taxa, Sigmoid curve: An S-shaped curve (see ‘Sexual dimor-
which may include correction of errors, synonymy or re- phism’).
validation of names and, possibly, the addition of newly
described taxa. Silt: A medium-sized soil grain (see ‘Soil classification’).

Ruminant herbivores: In reference to mammals; grazing Soil classification: Based on the system used by the
or browsing herbivores in which there is a four-chambered United States Department of Agriculture (USDA): sand,
stomach; dung usually fine-fibred, may be dropped as sandy loam, sandy clay loam, clay; each soil type com-
pads or pellets, e.g. cattle, sheep. prises different proportions of large (sand), medium (silt)
or small (clay) grains with a bias to either small or larger
grain size; soil grain size profiles are important for tun-
S nelling by dung beetles, some species are associated with
particular soil types.
Sand: The coarsest-grained division in the United States
Department of Agriculture’s (USDA) classification of soil Specialisation: See ‘Specialist’.
types (see ‘Soil classification’).
Specialist: A species that shows a narrow range of eco-
Sandy clay loam: The penultimate division towards finer- logical associations due to specialisation to one or more
grained soils in the United States Department of Agricul- particular factors (e.g. found only on deep sands, not in
ture’s (USDA) classification of soil types (see ‘Soil classi- finer-grained soil types, such as sandy loam or clay).
fication’).
Species: See ‘Classification’.
Sandy loam: The penultimate division towards coarser-
grained soils in the United States Department of Agricul- Species accounts: Refers to the sum total of information
ture’s (USDA) classification of soil types (see ‘Soil classi- provided for each species.
fication’).
Species complex: Refers to a cryptic (hidden) group of
Scarabaeidae: A family of beetles (Order Coleoptera) species that are very close in appearance to one another.
comprising 16 subfamilies that have diversified primarily
Species richness: In the sense used here: the number of
in association with living plants and dead or waste organic
recorded species.
matter.
Spectra of released volatiles: Refers to the range of chem-
Scarabaeinae: Subfamily of the Scarabaeidae to which the icals released into the air from dung, which are perceived
dung beetles featured in this atlas are allotted. by dung beetles by their sense of smell; only particular
chemicals within this range elicit a response, which is used
Secondary sexual armature: Variously characterised in
to locate the dropping.
different species by horns, projections and/or dentition.
Spp.: Species (plural).
Sensu: ‘In the sense of ’; used here to denote species that
were misidentified by an author and listed in the taxo- Standard deviation: A statistical range of variation
nomic literature under the wrong name. around a mean value (see ‘Means’).

Sexual dimorphism: Refers to the quite different sculptur- Subfamily: See ‘Classification’ and ‘Scarabaeinae’.
ing of the exoskeleton shown by males of many dung beetle
species compared to females; the most frequent examples Subsp.: Subspecies.
SURICATA 6 (2020) 787

Subspecies: See ‘Classification’. Themeda-dominated grassland: Many areas of natural

grassland on the Highveld of South Africa may be domi-
Succession: Refers to the replacement of different taxa nated by the genus, Themeda.
over time (= turn over), e.g. successive replacement of taxa
over time in response to the process of drying after depo- Topography: The physical surface relief of a geographical
sition of dung. area in terms of hills and mountains etc.

Superfamily: See ‘Classification’; dung beetles of the Sub- Topology: The pattern of clades and lineages on a evo-
family Scarabaeinae are allotted to the Family Scarabaeidae lutionary tree; a ladderised topology is one where each
in the Superfamily Scarabaeoidea. successive node gives rise to the remainder of the evolu-
tionary tree.
Survivorship: The proportion of individuals in a popula-
tion surviving over a defined timescale. Tribal: See ‘Tribe’.

Synonym: In taxonomy; a name or names provided for a Tribe: A level of classification of plants and animals (see
species, which had already been previously described un- ‘Classification’).
der a different name; under the rule of precedence, only
the first-described species name is valid. Trophic: Pertaining to food type and feeding.

Syntype: See ‘Type specimens’; two or more specimens in Tunneller: One of four major behavioural groups in
a type series for which no single specimen was designated scarabaeine dung beetles; species that construct tunnels
as a holotype, nor subsequently as a lectotype; all individ- from under dung into which dung is buried a piece at a
uals in the type series are considered to have equal rank; time (also see ‘Ball roller’, ‘Endocoprid’, ‘Kleptocoprid’).
species described a long time ago are often supported by
syntypes. Type localities: A single or several localities of origin de-
fined for type specimens of a species.
Systematics: Study of the evolutionary relationships be-
tween taxa (see ‘Phylogeny’). Type specimen(s): Designated specimen(s) that repre-
sent(s) a species name formally described in a publication.
Specimen(s) lodged in a museum or scientific collection
T where it/they may be viewed by other researchers; see ho-
lotype, allotype, lectotype, neotype, syntype, paratype.
Taxa: Plural of taxon.

Taxon: Term of reference for an organism (or organisms) V

without specifying the level it occupies in the hierarchy of
classification (see ‘Classification’). Validated: Confirmation that the name used for a partic-
ular species is valid; species identity has been validated for
Taxonomy: Allotment of taxa to levels in a classification all non-type material in this atlas, unless stated otherwise
system (see ‘Taxon’ and ‘Classification’). in the ‘Taxonomy’ sections of species accounts.

Temporal associations: Refers to time or period of activity Validation: See ‘Validated’.

at different scales in dung beetles; these may be predictable
in response to daily (diel) or seasonal cycles of light and tem- Varietal: See ‘Variety’.
perature or they may vary non-cyclically in time according
rainfall events or from year-to-year according to the cumu- Variety: Represents specified variation within a species,
lative effect of preceding ecologically important factors. e.g. colour varieties or exoskeleton sculptural varieties;
often used to distinguish variation between populations
Terminally derived: Lineage derived from a node on across a geographical range (see ‘Classification’).
higher branches of an evolutionary tree; in terms of se-
quence of derivation, represents later evolution along Vegetation classification: Two systems of vegetation
the lineage than those branches derived from more basal classification are used in the atlas, i.e., the hierarchi-
nodes along the same lineage (also see ‘Basally derived’). cal three-scale classification of Mucina and Rutherford
(2006) for South Africa (see ‘Biome’, ‘Bioregion’, ‘Veg-
Textural: Used here in terms of grain size structure of etation unit’) and the ecoregion classification of Olson
soils. et al. (2001) for the remainder of southern Africa; for
788 SURICATA 6 (2020)

habitat associations; physical structure of vegetation is by Mucina and Rutherford (2006) (also see ‘Biome’ and
classified according to increasing shade from grassland, ‘Bioregion’).
scrub, shrubland, open woodland, woodland and thick-
ets to forest. Vulnerable (VU): One of the Red List categories of threat
designated by the IUCN; used to identify a species that
Vegetation unit: Smallest of three geographical scales is considered to face threats of extinction in the wild (see
used to define floral distribution patterns in South Africa IUCN categories in Introduction).
SURICATA 6 (2020) 789

TAXONOMIC INDEX
of African taxa
The index only references Afrotropical taxa Anonychonitis . . . . . . . . . . 333–335, 711, 716, 727, 754
freyi . . . . . . . . . . . . . 334, 335, 711, 716, 727, 754
Subfamily/tribes: CAPITALS Aphengoecus . . . . . 58, 61–63, 701, 716, 724, 725, 748
Valid genera: bold italics clypeatus . . . . . . . . . . . . . 13, 61, 62, 716, 724, 748
Valid species: indented italics multiserratus . . . . . . . . . . . . . . . . . . . 63, 716, 748
Synonyms: roman text ATEUCHINI . . . . . . . . . . . . . 19, 20–55, 690, 716, 747
Ateuchus . . . . . . . . . . 94, 267, 271, 560, 561, 563, 564,
566-569, 573, 577, 594, 599, 607, 613,
Afrodrepanus . . . . . . . . . . 288, 290, 291, 710, 726, 754 616, 618, 620, 623, 625, 627, 629, 631,
impressicollis . . . . . . . . . . . . . . . 291, 710, 726, 754 632, 634, 640, 643-645, 647, 652, 653,
Afroharoldius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423 658, 673, 677
Aliuscanthoniola . . . . . . . . . . 57, 59, 60, 716, 719, 748 Bohepilissus . . . . . . 57, 64–66, 700, 710, 716, 722, 748
similaris . . . . . . . . . . . . . . . . 59, 60, 716, 719, 748 nitidus . . . . . . . . . . . . . . . . . . . . 65, 716, 722, 748
Allogymnopleurus . . . . . . 262–266, 726, 740, 753, 777 subtilis . . . . . . . . . . . . . . . . . . . . 66, 710, 722, 748
chloris . . . . . . . . . . . . . . . . . . . . . . . . . . . 264, 266 Byrrhidium . . . . . . . . . . 57, 67–69, 710, 716, 720, 748
consocius . . . . . . . . . . . . . . . . . . 264, 265, 726, 753 brevipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67, 69
coracinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 convexum . . . . . . . . . . . . . . . . . . . . . . 68, 720, 748
splendidus . . . . . . . . . . . . . . . . 266, 726, 740, 753 namaquensis . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
subcupratus . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 ovale . . . . . . . . . . . . . . 67, 69, 710, 716, 720, 748
thalassinus . . . . . . . . . . . . . . . . . . . . . . . . 264, 266 Caccobiomorphus . . . . . . 395, 396–398, 729, 756, 766
Anachalcos . . . . . . . . . . . . . . . . . . . . . . . . . . . 58, 70, 71 brevisetosus . . . . . . . . . . . . . . . . . . . . . . . 396, 397
790 SURICATA 6 (2020)

congolanus . . . . . . . . . . . . . . . . . . . . . . . . . . . 397 Cheironitis . . . . . . . . . . . 332, 333, 336–341, 344, 701,

megaponerae . . . . . . . . . . . . . . . 396, 397, 729, 756 727, 728, 741, 754, 755, 767
Caccobius . . . . . . . . 133, 141, 146, 147, 394–408, 438, audens . . . . . . . . . . . . . . . . . . . 337, 727, 741, 754
691, 729, 742, 756, 767, 778 hoplosternus . . . . . . . . . . . . . . . 338, 728, 741, 754
castaneus . . . . . . . . . . . . . . 398, 399, 729, 742, 756 imitator . . . . . . . . . . . . . . . . . . 339, 728, 741, 754
cavatus . . . . . . . . . . . . . . . . . . . 400, 729, 742, 756 indicus . . . . . . . . . . . . . . . 339, 340, 728, 741, 755
ferrugineus . . . . . . . . . . . . 246, 401, 729, 742, 756 scabrosus . . . . . . . . . . . . . . . . . . 341, 728, 741, 755
histerinus . . . . . . . . . . . . . . . . . 402, 729, 742, 756 Chironitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
inconspicuus . . . . . . . . . . . . . . . . . . . . . . . . . . . 691 Circellium . . . . . . . . . . . . . . . 57, 58, 72, 73, 700, 710,
litigiosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403 716, 720, 748, 778
mastrucatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 404 bacchus . . . . . . . . . . . . . . . 73, 710, 716, 720, 748
medioniger . . . . . . . . . . . . . . . . . . . . . . . . . . . 399 bacchus var. waterhousei . . . . . . . . . . . . . . . . . . 73
nigritulus . . . . . . . . . . . . . . . . . 403, 729, 742, 756 hemisphaericus . . . . . . . . . . . . . . . . . . . . . . . . . 73
obtusus . . . . . . . 103, 404, 710, 724, 729, 749, 756 lyceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
viridicollis var. picipennis . . . . . . . . . . . . . . . . 406 Cleptocaccobius . . . . . . . . . . . 394, 395, 398, 405, 406,
CANTHONINI . . . . . . . . . 19, 56–163, 716, 748, 750 407, 408, 729, 742, 756
Catharsius . . . . . . . 4, 8, 164–180, 182, 183, 246–249, convexifrons . . . . . . . . . . . . . . . 406, 729, 742, 756
251, 252, 254–257, 690, 701, 710, minimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 408
712, 722, 739, 750, 751, 767, 770, postlutatus . . . . . . . . . . . . . . . . . . . . 407, 729, 756
771, 773, 775, 778 pudens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 406
aegeus . . . . . . . . . . . . . . . . . . . 167, 722, 739, 750 viridicollis . . . . . . . . . . . . . . . . 408, 729, 742, 756
areolatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 var. semicoeruleus . . . . . . . . . . . . . . . . . . . . . . 408
bradshawi . . . . . . . . . . . . . . . . . . . . . . . . . . 8, 168 COPRINI . . . . . . . . . 19, 164–261, 690, 716, 750, 752
Coprioides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
calaharicus . . . . . . . . . . . . 169, 712, 722, 739, 750
Copris . . . . 4, 5, 13, 164–166, 178, 179, 184, 186–202,
camillus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
204–219, 225, 227–230, 250, 253, 256,
disseptus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
259, 324, 455, 477, 494, 525, 540, 700,
dubius . . . . . . . . . . . . . . . . . . . . . . . . . . . 180, 251
701, 710–712, 716, 720, 721, 738, 739,
gibbicollis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
751, 752, 765,767, 768, 770, 773, 775,
harpagus . . . . . . . . . . . . . . . . . . . . . . 170, 722, 750
776, 778
heros . . . . . . . . . . . . . . . . 171, 722, 739, 750, 775
achates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
insignis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
amyntor . . . . . . . . . . . . . . 186, 712, 720, 738, 751
laticeps . . . . . . . . . . . . . . . . . . . . . . . 172, 722, 750
anceus . . . . . . . . . . . . . . . . . . . . . . . 187, 720, 751
longiceps . . . . . . . . . . . . . . . . . . . . . . 173, 722, 750
antares . . . . . . . . . . . . . . . . . . . 188, 710, 720, 751
marcellus . . . . . . . . . . . . . . . . . . 174, 710, 722, 750 bihamatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
melancholicus . . . . . . . . . . . . . . 175, 722, 739, 750 bootes . . . . . . . . . . . . . . . . 189, 720, 738, 751, 775
melancholicus . . . . . . . . . . . . . . . . . . . . . . . . . 173 bornemisszai . . . . . . . . . . . . . . . . . . . . . . . . . . 189
mossambicanus . . . . . . . . . . . . . . . . . . . . . . . . 176 caelatus . . . . . . . . . . . . . . . . . . 190, 710, 720, 751
nemestrinus . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 cambeforti . . . . . . . . . . . . . . . . . . . . 191, 720, 751
obtusicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 capensis . . . . . . . . . . . . . . . . . . 192, 275, 720, 751
pandion . . . . . . . . . . . . . . . . . . . . . . 176, 722, 750 cassius . . . . . . . . . . . . . . . . . . . 193, 720, 738, 751
parafastidiosus . . . . . . . . . . . . . . . . . . . . . . . . . . 172 confusus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
pithecius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 contracta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
philus . . . . . . . . . . . . . . . . 177, 712, 722, 739, 750 contractus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
platycerus . . . . . . . . . . . . . . . . . . . . . 178, 722, 750 cornifrons . . . . . . . . . . . . . 194, 712, 720, 739, 751
princeps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178 corniger . . . . . . . . . . . . . . 195, 710, 721, 751, 775
pylades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 crassus . . . . . . . . . . . . . . . . . . . . . . . 196, 721, 751
sesostris . . . . . . . . . . . . . . . . . . . . . . . 179, 722, 750 curvicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
tricornutus . . . . . . . . . . . . . . . . 180, 722, 739, 750 denticulatus . . . . . . . . . . . . . . . 197, 721, 739, 751
ulysses . . . . . . . . . . . . . . . . . . . . 182, 722, 739, 751 elphenor . . . . . . . . . . . . 5, 198, 712, 721, 739, 751
vansoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 evanidus . . . . . . . . . . . . . . . . . . 199, 721, 739, 751
vitulus . . . . . . . . . . . . . . . . . . . 183, 710, 722, 751 excavata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Chalconotus . . . . . . . . . . . . . . 56–58, 70, 71, 115, 119, globulipennis . . . . . . . . . . . . . . . . . . . . . . . . . . 213
700, 723, 740, 748 gracilis . . . . . . . . . . . . . . . . . . . . . . . 201, 739, 751
convexus . . . . . . . . . . . . . . . . . . . 71, 723, 740, 748 fidius . . . . . . . . . . . . . . . . 200, 721, 751, 768, 775
sericeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 inhalatus . . . . . . . . . . . . . . 202, 711, 721, 739, 751
SURICATA 6 (2020) 791

jacchoides . . . . . . . . . . . . . . . . . . . . . 204, 721, 751 deschodti . . . . . . . . . . . . . . . . 74, 75, 716, 738, 748
jacchus . . . . . . . . . . . . . . . 205, 710, 721, 751, 776 inexpectata . . . . . . . . . . . . . . . . . . . . . 76, 738, 748
laioides . . . . . . . . . . . . . . . . . . . 206, 712, 739, 751 tatasensis . . . . . . . . . . . . . . . . . . . . . . . 77, 720, 748
laticornis . . . . . . . . . . . . . . . . . . . . . 195, 198, 205 vandersmisseni . . . . . . . . . . . . . . . . . . . 78, 738, 748
lineatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213 Digitonthophagus . . . . . . . . . . 394, 409, 410, 412, 413,
macer . . . . . . . . . . . . . . . . . . . . . . . . 207, 721, 751 442, 729, 742, 756, 771
mesacanthus . . . . . . . . . . . . . . . 208, 721, 739, 751 dorcas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 410
minator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 gazella . . . . . . . . . . . . . . . . . . . 410, 729, 742, 756
misellus . . . . . . . . . . . . . . . . . . . . . . . . . . 184, 209 namaquensis . . . . . . . . . . . . . . . 412, 729, 742, 756
modestus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 viridicollis . . . . . . . . . . . . . . . . 413, 729, 742, 756
obesus . . . . . . . . . . . . . . . . . . . . . . . . 209, 721, 751 Drepanellus . . . . . . . . . . . . . . . . . . . . . . . . . . . 312, 766
oedipus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205 Drepanocerus . . . . . . . . . 288, 290–292, 294–300, 302,
orion . . . . . . . . . . . . . . . . 210, 711, 721, 751, 776 310–313, 321–325, 726, 740, 754, 766
orphanus . . . . . . . . . . . . . . . . . . . . . . . . . 199, 201 arthuri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
plutus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 kirbyi . . . . . . . . . . . . . . . . 294, 295, 726, 740, 754
puncticollis . . . . . . . . . . . . . . . . 211, 721, 739, 751 patrizii . . . . . . . . . . . . . . . . . . . . . . 296, 726, 754
ritsemae . . . . . . . . . . . . . . . . . . . . . . 212, 721, 751 Drepanochirus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292
similis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208 Drepanopodus . . . . . . . . . . . . . . 556, 618, 624, 627, 641
sphaeropterus . . . . . . . . . . . 13, 214, 716, 721, 751 Drogo . . . . . . . . . . . . . . . . . . . . . . 58, 79, 80, 738, 748
subsidens . . . . . . . . . . . . . . . . . . 215, 712, 739, 752 stalsi . . . . . . . . . . . . . . . . . . . . . . 79, 80, 738, 748
urus . . . . . . . . . . . . . . . . . . . . . 216, 721, 752, 767 Dwesasilvasedis . . . . . . . . . . . . . . . 58, 81, 82, 716, 725
victorini . . . . . . . . . . . . . . . . . . 217, 716, 721, 752 medinae . . . . . . . . . . . . . . . . 81, 82, 716, 725, 748
victorini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212 Elassocanthon . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67, 69
vilhenai . . . . . . . . . . . . . . . . . . 218, 721, 739, 752 Endroedyantus . . . . . . . . . . . . . . . . . . . . . . . 85, 97, 111
vrydaghi . . . . . . . . . . . . . . . . . . . . . . 219, 721, 752 Endroedyolus . . . . . . . . . 57, 83, 84, 710, 716, 719, 748
Coptorhina . . . . . . . . . . . . . . . . 20–27, 31–34, 36, 700, paradoxus . . . . . . . . . . . 83, 84, 710, 716, 719, 748
710, 718, 738, 747, 771 Eodrepanus . . . . . . . 288, 289, 297–300, 727, 754, 766
africana . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 25 bechynei . . . . . . . . . . . . . . . . . . . . . . 298, 727, 754
auspicata . . . . . . . . . . . . . . . . . . . 23, 718, 738, 747 fastiditus . . . . . . . . . . . . . . . . . . . . . . 299, 727, 754
bicolor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 parallelus . . . . . . . . . . . . . . . . . . . . . 300, 727, 754
excavata . . . . . . . . . . . . . . . . . . . 24, 710, 718, 747 Ephillopus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
klugi . . . . . . . . . . . . . . . . . . . . . . . . . 25, 718, 747 Epidrepanus . . . . . . 289, 301, 302, 312, 313, 727, 754
nitidipennis . . . . . . . . . . . . . . . . 26, 718, 738, 747 caelatus . . . . . . . . . . . . . . . . . . . . . . 302, 727, 754
obtusicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 pulvinarius . . . . . . . . . . . . . . . . . . . . . . . . 302, 727
optata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Epionitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389, 766
punctata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Epirinus . . . . . 19, 56, 58, 85–114, 660, 701, 710, 712,
pygmaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 716, 723, 724, 748, 749, 767, 768,
saganicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 770, 775, 777
seminitida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 aeneus . . . . . . . . . . . . . . . . . . . . . . . . 86, 723, 748
subaenea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 aquilus . . . . . . . . . . . . . . . . . . . . 87, 716, 723, 748
vicina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 asper . . . . . . . . . . . . . . . . . . . . . 88, 710, 723, 748
Cyclotrogus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423 bentoi . . . . . . . . . . . . . . . . . . . . 89, 716, 723, 748
Cyptochirus . . . . . . . . . . . . . . 288, 292, 293, 296, 726, callosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
754, 767, 778 comosus . . . . . . . . . . . . . . . 90, 710, 716, 723, 748
ambiguus . . . . . . . . . . . . . . . . . 293, 710, 726, 754 convexus . . . . . . . . . . . . . . . . . . . 91, 716, 723, 748
impressus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293 davisi . . . . . . . . . . . . . . . . . . . . . 92, 716, 723, 748
Delopleurus . . . . . . . . . . . . . . 20, 21, 27–30, 700, 718, deplanatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
738, 747, 771, 773 drakomontanus . . . . . . . . . . . . . . . . . . 93, 723, 748
darrenmanni . . . . . . . . . . . . . . . 28, 712, 738, 747 flagellatus . . . . . . . . . . . . . . . . . . . 85, 94, 724, 748
gilleti . . . . . . . . . . . . . . . . . . . . . . . . . 29, 738, 747 granulatus . . . . . . . . . . . . . . . . . 95, 716, 724, 748
pullus . . . . . . . . . . . . . . . . . . 27, 30, 718, 738, 747 gratus . . . . . . . . . . . . . . . . . . . . . 96, 710, 724, 748
DELTOCHILINI . . . . . . . . . . . . . . . . . . . . 19, 56, 716 hilaris . . . . . . . . . . . . . . . . . . . . . . . . 97, 724, 748
Deronitis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329, 330 hluhluwensis . . . . . . . . . . . . . . . . 98, 716, 724, 748
Diaglyptus . . . . . . . . . . . . . . . . 133, 135, 141, 146, 147 minimus . . . . . . . . . . . . . . . . . . . 99, 716, 724, 748
Dicranocara . . . . . . . . . . . . . . 57, 58, 74–78, 716, 720, montanus . . . . . . . . . . . . . . . . . 100, 716, 724, 748
738, 748, 775 mucrodentatus . . . . . . . . . . . . . . . . . . 101, 724, 748
792 SURICATA 6 (2020)

ngomae . . . . . . . . . . . . . . . . . . 102, 716, 724, 749 delagorguei . . . . . . . . . . . . . . . . . . . . . . . . . . . 272

obtusus . . . . . . . . . . . . . . . . . . . 103, 710, 724, 749 hilaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
pseudorugosus . . . . . . . . . . . . . . 104, 716, 724, 749 insidiosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270
punctatus . . . . . . . . . . . . . 105, 710, 716, 724, 749 laetus . . . . . . . . . . . . . . . . . . . . . . . . 270, 726, 753
pygidialus . . . . . . . . . . . . . . . . . . . . . . . . . 106, 749 nitens . . . . . . . . . . . . . . . . . . . . . . . . 271, 740, 753
relictus . . . . . . . . . . . . . . . . . . . 107, 710, 724, 749 splendens . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271
rugosus . . . . . . . . . . . . . . . . . . . . . . . 108, 724, 749 unicolor . . . . . . . . . . . . . . . . . . . . . . 272, 726, 753
scabratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 wahlbergi . . . . . . . . . . . . . 273, 712, 726, 740, 753
scrobiculatus . . . . . . . . . . . . . . . 109, 710, 724, 749 zumpti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
sebastiani . . . . . . . . . . . . . . . . . 110, 716, 724, 749 Gilletellus . . . . . . . . 333, 342, 343, 350, 716, 741, 755
silvestris . . . . . . . . . . . . . . . . . . 111, 710, 724, 749 porculus . . . . . . . . . . 343, 350, 351, 716, 741, 755
striatus . . . . . . . . . . . . . . . . . . . 112, 712, 724, 749 GYMNOPLEURINI . . . . . . . . . . . . . 19, 262–286, 753
sulcipennis . . . . . . . . . . . . . . . . . . . . 113, 724, 749 Gymnopleurus . . . . . . . . 262–264, 266, 267, 269–286,
validus . . . . . . . . . . . . . . . . . . . 114, 710, 724, 749 712, 726, 740, 753, 768, 770,
vicinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 772, 778
youngae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 aenescens . . . . . . . . . . . . . . . . . . 275, 726, 740, 753
Escarabaeus . . . . . . . . . . . 10, 556–560, 618, 628, 701, var. capensis . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
712, 733, 744, 760, 769 andreaei . . . . . . . . . . . . . . 276, 726, 734, 740, 753
remii . . . . . . . . . . . . . . . . . 10, 559, 712, 744, 760 asperrimus . . . . . . . . . . . . 277, 712, 726, 740, 753
satyrus . . . . . . . . . . . 559, 560, 628, 733, 744, 760 bicolor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Euoniticellus . . . . . . . . . . . . . 288, 289, 303–309, 710, bufo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
727, 740, 741, 754, 774 caelatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
africanus . . . . . . . . . . . . . . . . . . . . . . 304, 727, 754 cupreus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
clavatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 gibbosus . . . . . . . . . . . . . . . . . . . . . . . . . . 283, 286
intermedius . . . . . . . . . . . 305, 306, 727, 740, 754 humanus . . . . . . . . . . . . . . 278, 712, 726, 740, 753
kawanus . . . . . . . . . . . . . . . . . . 307, 727, 740, 754 var. azureus . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
nasicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 humeralis . . . . . . . . . . . . . . . . . 279, 726, 740, 753
speciosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305 ignitus . . . . . . . . . . . . . . . . . . . . . . . 280, 740, 753
triangulatus . . . . . . . . . . . 308, 710, 727, 741, 754 var. laeviscula . . . . . . . . . . . . . . . . . . . . . . . . . . 280
zumpti . . . . . . . . . . . . . . . . . . . . . . . 309, 727, 754 var. nigrocupreus . . . . . . . . . . . . . . . . . . . . . . . 280
Euonthophagus . . . . . . . . . . . . 395, 414–418, 442, 712, imitator . . . . . . . . . . . . . . . . . . . . . . . . . . 281, 740
729, 742, 756 laeviuscula . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
carbonarius . . . . . . . . . . . 415, 418, 729, 742, 756 nigrocupreus . . . . . . . . . . . . . . . . . . . . . . . . . . 280
flavimargo . . . . . . . . . . . . 416, 712, 729, 742, 756 leei . . . . . . . . . . . . . . . . . . . . . . . . . . 282, 726, 753
jeanneli . . . . . . . . . . . . . . . . . . 417, 712, 729, 756 liechtensteini . . . . . . . . . . . . . . . . . . . . . . . . . . 282
vicarius . . . . . . . . . . . . . . 415, 418, 712, 729, 756 macleayi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Formicdubius . . . . . . . . . . . . . . . . . . . . . . 423, 424, 774 modestus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Frankenbergerius . . . . . . . . . . 20, 21, 31–38, 700, 710, peringueyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
712, 716, 718, 747, 771 pumilus . . . . . . . . . . . . . . . . . . 283, 726, 740, 753
armatus . . . . . . . . . . . . . . . . . . . 32, 710, 718, 747 pumilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 286
barratti . . . . . . . . . . . . . . . . . . . 33, 716, 718, 747 reichei . . . . . . . . . . . . . . . . . . . . . . . 284, 726, 753
forcipata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 sericatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
forcipatus . . . . . . . . . . . . . . . . . . . . . . 34, 718, 747 smaragdinus . . . . . . . . . . . . . . . . . . . . . . . . . . 280
gomesi . . . . . . . . . . . . . . . . . . . . 35, 716, 718, 747 thelwalli . . . . . . . . . . . . . . . . . . . . . . 285, 726, 753
granulifera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 vanderkelleni . . . . . . . . . . . . . . . . . . . . . . 275, 286
imitativa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 virens . . . . . . . . . . . . . . . . . . . . 286, 726, 740, 753
mirabilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31, 32 subsp. sternalis . . . . . . . . . . . . . . . . . . . . 283, 286
nanus . . . . . . . . . . . . . . . . . . . . . 36, 716, 718, 747 Gyronotus . . . . . . . . . . . . . . . . . 57, 115–120, 700, 711,
nitidus . . . . . . . . . . . . . . . . . . . . . . . . 37, 718, 747 716, 721, 722, 749, 775
opacus . . . . . . . . . . . . . . . . . . . . 38, 716, 718, 747 carinatus . . . . . . . . . . . . . . 116, 711, 716, 721, 749
Garreta . . . . . . . . . . . . . . . . . . 262, 263, 267–273, 711, glabrosus . . . . . . . . . . . . . . . . . . 117, 716, 722, 749
712, 726, 740, 753, 777 perissinottoi . . . . . . . . . . . . . . . 118, 716, 722, 749
australugens . . . . . . . . . . . . . . . . . . . 268, 726, 753 pumilus . . . . . . . . . . . . . . . . . . 119, 711, 722, 749
azureus . . . . . . . . . . . . . . . . . . . . . . . . . . 270, 271 schuelei . . . . . . . . . . . . . . . . . . . . . . . . . . 120, 749
caffer . . . . . . . . . . . . . . . . . . . . 269, 711, 726, 753 Hammondantus . . . . . . . . 58, 121, 122, 701, 740, 749
cupreovirens . . . . . . . . . . . . . . . . . . . . . . . . . . 270 psammophilus . . . . . . . . . . . . . . 122, 712, 740, 749
SURICATA 6 (2020) 793

Hamonthophagus . . . . . . . . . . 395, 419, 420, 421, 422, var. orientalis . . . . . . . . . . . . . . . . . . . . . . . . . . 569

442, 730, 742, 756 usurpator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 567
acutus . . . . . . . . . . . . . . . . . . . 420, 730, 742, 756 vethi . . . . . . . . . . . . . . . . . . . . 570, 712, 745, 760
carteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421 zurstrasseni . . . . . . . . . . . . . . . . 571, 716, 734, 760
depressus . . . . . . . . . . . . . . . . . . 421, 730, 742, 756 Kolbeellus . . . . . . . . . . . . 333, 346, 347, 716, 728, 755
var. marmoreus . . . . . . . . . . . . . . . . . . . . . . . . 421 ateuchoides . . . . . . . . . . . . . . . . 347, 716, 728, 755
fallax . . . . . . . . . . . . . . . . . . . . . . . . 422, 742, 756 Kurtops . . . . . . . . . . . . . . . . 6, 394, 430–434, 442, 446,
laceratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 421 712, 730, 742, 757
Haroldius . . . . . . . . 394, 395, 423, 424, 730, 756, 774 caffrarius . . . . . . . . . . . . . . . . . . . . . . . . 6, 431, 730
convexus . . . . . . . . . . . . . . . . . . . . . . 424, 730, 756 junodi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433
Heliocantharus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 618 quadraticeps . . . . . . . . . . . 432, 712, 730, 742, 757
Heliocopris . . . . . . 4, 39, 164, 165, 220–227, 442, 699, signatus . . . . . . . . . . . . . . 433, 434, 730, 742, 757
701, 710, 723, 740, 752, 777 Latodrepanus . . . . . . 289, 301, 312, 313, 727, 741, 754
andersoni . . . . . . . . . . . . . . . . . 221, 723, 740, 752 laticollis . . . . . . . . . . . . . . . . . . 313, 727, 741, 754
atropos . . . . . . . . . . . . . . . . . . . 222, 723, 740, 752 Liatongus . . . . . . . . . . . . 288, 289, 314–316, 318, 320,
bicarinulatus . . . . . . . . . . . . . . . . . . . . . . . . . . 225 710, 727, 741, 754, 773, 777
coriaceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 militaris . . . . . . . . . . . . . . 315, 710, 727, 741, 754
eryx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226 Litocopris . . . . . . . . . . . . 164, 165, 228–230, 721, 752
faunus . . . . . . . . . . . . . . . 223, 710, 723, 740, 752 muticus . . . . . . . . . . . . . . . . . . . . . . 229, 721, 752
hamadryas . . . . . . . . . . . . 224, 710, 723, 740, 752 simplex . . . . . . . . . . . . . . . . . . . . . . . 230, 721, 752
isidis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 Macroderes . . . . . . . . 58, 164, 165, 231–245, 701, 710,
japetus . . . . . . . . . . . . . . . . . . . 225, 723, 740, 752 716, 725, 726, 752, 765, 771, 777
neptunus . . . . . . . . . . . . . . . . . . 226, 723, 740, 752 amplior . . . . . . . . . . . . . . . . . . 232, 716, 725, 752
operosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223 arrowi . . . . . . . . . . . . . . . 233, 651, 716, 725, 752
pirmal . . . . . . . . . . . . . . . . . . . . . . . 227, 723, 752 bias . . . . . . . . . . . . . . . . . . . . . . . . . . 234, 725, 752
satyrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223 cornutus . . . . . . . . . . . . . . . . . . 235, 716, 725, 752
Heteroclitopus . . . . . . . . . 395, 425, 426, 730, 756, 767 endroedyi . . . . . . . . . . . . . . . . . 236, 716, 725, 752
remipes . . . . . . . . . . . . . . . . . . . 425, 426, 730, 756 fornicatus . . . . . . . . . . . . . . . . . 237, 716, 725, 752
Heteronitis . . . . . . . 333, 336, 344, 345, 728, 741, 755 foveatus . . . . . . . . . . . . . . . . . . 238, 716, 725, 752
castelnaui . . . . . . . . . . . . . . . . . 345, 728, 741, 755 greeni . . . . . . . . . . . . . . . . . . . . 239, 716, 725, 752
Histeridium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398 minutus . . . . . . . . . . . . . . . . . . 240, 716, 725, 752
Hyalonthophagus . . . . . . . . . . . . . . 395, 427–429, 730, mutilans . . . . . . . . . . . . . . . . . . 241, 710, 725, 752
742, 757, 769, 776 namakwanus . . . . . . . . . . . . . . 242, 716, 725, 752
alcyon . . . . . . . . . . . . . . . . . . . 428, 730, 742, 757 nitidus . . . . . . . . . . . . . . . . . . . . . . . 243, 725, 752
alcyonides . . . . . . . . . . . . . . . . . 429, 730, 742, 757 pilula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234
var. virens . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429 politulus . . . . . . . . . . . . . . . . . . 244, 716, 725, 752
var. viridiceps . . . . . . . . . . . . . . . . . . . . . . . . . . 429 undulatus . . . . . . . . . . . . . . . . . 245, 716, 726, 752
Irrorhotides . . . . . . . . . . . . . . . . . . . . . . . . . . . 575, 588 westwoodi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Ixodina . . . . . . . . . . 288, 289, 310, 311, 322, 727, 754 Megalonitis . . . . . . . . . . . 333, 348, 349, 728, 741, 755
abyssinica . . . . . . . . . . . . . . . . . 310, 311, 727, 754 bohemani . . . . . . . . . . . . . . . . . 349, 728, 741, 755
Kheper . . . . . . . . . . 556, 557, 561–571, 590, 701, 711, Megaponerophilus . . . . . . . . . . . . . . . . . . . . . . . . . . 396
712, 716, 734, 745, 760, 768, 769, Melinocerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 540
773, 775, 777 Mesoscarabaeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 558
bonellii . . . . . . . . . . . . . . . . . . . . . . 562, 734, 760 Metacatharsius . . . . . . . . 164, 165, 246–257, 690, 691,
clericus . . . . . . . . . . . . . . . 563, 711, 716, 734, 760 701, 710, 722, 723, 739, 752,
cupreus . . . . . . . . . . . . . . . . . . . 564, 734, 745, 760 753, 770, 778
infernalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 566 anderseni . . . . . . . . . . . . . 247, 712, 722, 739, 752
kalaharicus . . . . . . . . . . . . . . . . 565, 734, 745, 760 cephalotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 255
lamarcki . . . . . . . . . . . . . . . . . . 566, 734, 745, 760 dentinum . . . . . . . . . . . . . . . . . . . . . 248, 739, 752
namibicus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 564 dubius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
nigroaeneus . . . . . . . . . . . . . . . 567, 734, 745, 760 exiguiformis . . . . . . . . . . . . . . . 249, 722, 739, 752
pubiventris . . . . . . . . . . . . . . . . . . . . . . . . . . . 568 exiguiformis . . . . . . . . . . . . . . . . . . . . . . . . . . 247
prodigiosus . . . . . . . . . . . . . . . . 568, 734, 745, 760 exiguus . . . . . . . . . . . . . . . . . . . 250, 722, 739, 752
subaeneus . . . . . . . . . . . . . . . . . 569, 734, 745, 760 ferreirae . . . . . . . . . . . . . . . . . . 251, 722, 739, 753
subsp. angolensis . . . . . . . . . . . . . . . . . . . . . . . 569 freyi . . . . . . . . . . . . . . . . . . . . . 252, 722, 739, 753
var. atratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 569 kochi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
794 SURICATA 6 (2020)

latifrons . . . . . . . . . . . . . . . . . . 253, 722, 739, 753 quadricollis . . . . . . . . . . . 662, 671, 673, 737, 763
marani . . . . . . . . . . . . . . . 254, 712, 723, 739, 753 rubrus . . . . . . . . . . . . . . . 662, 672, 737, 746, 763
opacus . . . . . . . . . . . . . . . . . . . 255, 723, 739, 753 rugosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 664
pseudoopacus . . . . . . . . . . . . . . . . . . . . . . . . . . 255 spinipes . . . . . . . . . . . . . . 662, 667, 673, 737, 763
pumilioniformis . . . . . . . . . . . . . . . . 256, 739, 753 spinipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 667
pumilionis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256 Odontoloma . . . . . . . . . . . . 56–58, 133–148, 700, 710,
pusio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690 719, 738, 749, 750, 772
rugosipennis . . . . . . . . . . . . . . . . . . . . . . . 251, 690 apiculum . . . . . . . . . . . . . . . . . . . . . . 134, 719, 749
transvaalensis . . . . . . . . . . . . . . . . . . . . . . 246, 691 dentinum . . . . . . . . . . . . . . . . . 135, 710, 719, 749
troglodytes . . . . . . . . . . . . . . . . 257, 723, 739, 753 disalatum . . . . . . . . . . . . . . . . . . . . . 136, 719, 749
troglodytes . . . . . . . . . . . . . . . . . . . . . . . . . . . 247 doubei . . . . . . . . . . . . . . . . . . . . . . . 137, 719, 749
zuluanus . . . . . . . . . . . . . . . . . . . . . . . . . . 248, 753 endroedyi . . . . . . . . . . . . . . . . . . . . . 138, 719, 749
Milichus . . . . . . . . . . 395, 435, 436, 730, 742, 757, 767 louwi . . . . . . . . . . . . . . . . . . . . 139, 719, 738, 749
apicalis . . . . . . . . . . . . . . . . . . . 436, 730, 742, 757 multifidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Mimonthophagus . . . . . . . . . . 395, 437–440, 442, 711, obscurum . . . . . . . . . . . . . . . . . . . . . 140, 719, 750
716, 730, 743, 757 pauxillum . . . . . . . . . . . . . . . . 133, 141, 719, 750
ambiguus . . . . . . . . . . . . . . . . . 438, 711, 730, 757 peckorum . . . . . . . . . . . . . . . . . . . . . 142, 719, 750
anomalus . . . . . . . . . . . . . . . . . 439, 730, 743, 757 planatum . . . . . . . . . . . . . . . . . . . . . 143, 719, 750
bicornutus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438 pusillum . . . . . . . . . . . . . . . . . . 144, 145, 719, 750
calvus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 pygidiale . . . . . . . . . . . . . . . . . . 145, 710, 719, 750
hinnulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438 quadridens . . . . . . . . . . . . . . . . . . . . 146, 719, 750
julianae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 sculpturatum . . . . . . . . . . . . . . . . . . 147, 719, 750
limbibasis . . . . . . . . . . . . . . . . . 440, 716, 743, 757 serraticeps . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
moestus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 spinicaudum . . . . . . . . . . . . . . . . . . 148, 719, 750
muripedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439 ONITICELLINI . . . . . . . . . . . . . . . . 19, 288–330, 754
Monapus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441 Oniticellus . . . . . . . 288, 289, 293, 295, 301–305, 308,
Namakwanus . . . . . . . . . . . . . . 57, 123–126, 128, 129, 314–320, 325, 326, 328, 429, 710,
161, 163, 738, 749, 768 712, 727, 741, 754, 773
irishi . . . . . . . . . . . . . . . . . . . . 123, 124, 738, 749 egregius . . . . . . . . . . . . . . 317, 712, 727, 741, 754
irishi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 formosus . . . . . . . . . . . . . . . . . . 318, 727, 741, 754
scholtzi . . . . . . . . . . . . . . . . . . . 125, 590, 738, 749 parapictus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 318
Namaphilus . . . . . . . . 58, 123, 126–129, 738, 749, 768 pictus . . . . . . . . . . . . . . . . . . . . 319, 710, 727, 754
ameibensis . . . . . . . . . . . . . . . . . . . . 127, 738, 749 pictus . . . . . . . . . . . . . . . . . . . . . . . . . . . 318, 319
davisi . . . . . . . . . . . . . . . . . . . . . . . . 128, 738, 749 subsp. orientalis . . . . . . . . . . . . . . . . . . . . . . . . 319
endroedyi . . . . . . . . . . . . . . . . . . . . . 129, 738, 749 planatus . . . . . . . . . . . . . . 316, 320, 727, 741, 754
Neateuchus . . . . . . . . . . . . . . . . . . . 556, 618, 640, 642 ONITINI . . . . . . . . . . . . . 19, 332–392, 716, 754, 756
Nebulasilvius . . . . . . . 58, 130–132, 151, 716, 719, 749 Onitis . . . . . . . . . . . . . 3, 292, 293, 332, 333, 337, 338,
insularis . . . . . . . . . . . . . . 130, 131, 716, 719, 749 342–345, 348, 349, 351, 352,
johani . . . . . . . . . . . . . . . . . . . 132, 716, 719, 749 354–366, 368–388, 391, 392, 701,
Neonitis . . . . . . . . . . . . . . . . . . 333, 350, 351, 728, 755 710, 712, 728, 729, 741, 742, 755,
nigritiae . . . . . . . . . . . . . . . . . . 350, 351, 728, 755 756, 766, 768–770, 773, 775
porculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351 aerarius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366
rhodesiae . . . . . . . . . . . . . . . . . . . . . . . . . 350, 351 aeruginosus . . . . . . . . . . . . . . . . 354, 728, 741, 755
Neopachysoma . . . . . . . . . . . . . . . . . 575, 579, 585, 779 aeruscator . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368
Neosisyphus . . . . . . . . . . . . . . 660–674, 710, 736, 737, africanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355
746, 763, 775 var. tuberculatus . . . . . . . . . . . . . . . . . . . . . . . 355
appendiculatus . . . . . . . . . . . . . . . . . . . . . . . . 673 alexis . . . . . . . . . . . . 355, 356, 710, 728, 741, 755
barbarossa . . . . . . . . . . . . 662, 663, 710, 736, 763 subsp. septentrionalis . . . . . . . . . . . . . . . . . . . . 355
bornemisszai . . . . . . . . . . . . . . . . . . . . . . . . . . 667 aygulus . . . . . . . . . . . . . . . . . . . 357, 728, 741, 755
calcaratus . . . . . . . . . . . . . . . . . 664, 736, 746, 763 aygulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355
confrater . . . . . . . . . . . . . . . . . . . . . . 665, 736, 763 caffer . . . . . . . . . . . . 359, 360, 710, 728, 741, 755
fortuitus . . . . . . . . . . . . . . . . . . . . . . 666, 736, 763 cerutii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
infuscatus . . . . . . . . . . . . . . . . . . . . . 667, 736, 763 confusus . . . . . . . . . . . . . . . . . . 361, 728, 741, 755
kuehni . . . . . . . . . . . . . . . . . . . . . . . 668, 737, 763 consanguineus . . . . . . . . . . . . . . . . . . . . . . . . . 387
macrorubrus . . . . . . . . . . . 669, 710, 737, 746, 763 cribratus . . . . . . . . . . . . . . . . . . . . . . 362, 728, 755
mirabilis . . . . . . . . . . . . . . . . . . . . . . 670, 737, 763 cupreus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366
SURICATA 6 (2020) 795

curvipes . . . . . . . . . . . . . . . . . . . . . . 363, 728, 755 apiciosus . . . . . . . . . . . . . . . . . . 449, 730, 743, 757

deceptor . . . . . . . . . . . . . . . . . . 364, 728, 741, 755 aschenborni . . . . . . . . . . . . . . . . . . . . . . . . . . . 691
fabricii . . . . . . . . . . . . . . . . . . . . . . . 365, 728, 755 asimilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691
fodiens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386 aspericeps . . . . . . . . . . . . . . . . . . . . . . . . . 460, 691
fulgidus . . . . . . . . . . . . . . . . . . 366, 728, 741, 755 asperrimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691
granulisetosus . . . . . . . . . . . . . . . . . . . . . . . . . . 354 asperulus . . . . . . . . . . . . . . . . . . . . . . 450, 730, 757
insulanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 387 axillaris . . . . . . . . . . . . . . . . . . . . . . 451, 743, 757
inuus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 355 bayeri . . . . . . . . . . . . . . . . . . . . . . . . 452, 743, 757
inversidens . . . . . . . . . . . . . . . . 368, 728, 741, 755 beiranus . . . . . . . . . . . . . . . . . . 453, 693, 730, 757
inversus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368 betschuanus . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
janssenii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371 bicavifrons . . . . . . . . . . . . . . . . 454, 730, 743, 757
klugii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366 bicolor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
licitus . . . . . . . . . . . . . . . . . . . . . . . . 369, 728, 755 biconifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
longitibialis . . . . . . . . . . . . . . . . . . . 370, 728, 755 binodis . . . . . . . . . . . . . . . . . . . 455, 710, 730, 757
lunaris . . . . . . . . . . . . . . . . . . . . . . . . . . . 184, 383 biplagiatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502
mendax . . . . . . . . . . . . . . 371, 712, 728, 741, 755 bovinus . . . . . . . . . . . . . . . . . . 456, 730, 743, 757
minutus . . . . . . . . . . . . . . . . . . . . . . 372, 728, 755 brevicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 509
mniszechi . . . . . . . . . . . . . . . . . . . . . 373, 741, 755 breviculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
nicanor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359 burchelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
obenbergeri . . . . . . . . . . . . . . . . 374, 728, 741, 755 cameloides . . . . . . . . . . . . . . . . 457, 710, 730, 757
obscuratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365 caucanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507
obscurus . . . . . . . . . . . . . . . . . . . . . . 375, 741, 755 chalcostomus . . . . . . . . . . . . . . . . . . . . . . . . . . 445
orthopus . . . . . . . . . . . . . . . . . . . . . . 376, 742, 755 cinctipennis . . . . . . . . . . . . . . . . . . . 458, 730, 757
var. tanganyikensis . . . . . . . . . . . . . . . . . . . . . . 376 cineraceus . . . . . . . . . . . . . . . . . . . . . 459, 730, 757
paraconfusus . . . . . . . . . . . . . . . . . . . 377, 728, 755 columella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455
parainflaticollis . . . . . . . . . . . . . 378, 728, 742, 755 confertus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
pecuarius . . . . . . . . . . . . . . 379, 710, 729, 755, 778 convexus . . . . . . . . . . . . . . . . . . 460, 730, 743, 757
perplexus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365 coptorhinodes . . . . . . . . . . . . . . . . . . . . . . . . . . 692
perpunctatus . . . . . . . . . . . . . . . . . . . 380, 729, 755 corniculatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 520
picticollis . . . . . . . . . . . . . . . . . . 381, 729, 742, 755 corniculiger . . . . . . . . . . . . . . . . . . . 461, 730, 757
pseudosetosus . . . . . . . . . . . . . . . . . . 382, 729, 756 costipennis . . . . . . . . . . . . . . . . . . . . . . . . . . . 503
robustus . . . . . . . . . . . . . . . . . . 383, 729, 742, 756 cretus . . . . . . . . . . . . . . . . . . . . 462, 711, 731, 757
setosus . . . . . . . . . . . . . . . . . . . . . . . 384, 742, 756 cretus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
sphinx . . . . . . . . . . . . . . . . . . . . . . . . . . . 352, 355 cribripennis . . . . . . . . . . . . . . . 463, 710, 731, 757
tortuosus . . . . . . . . . . . . . . . . . . . . . . 385, 729, 756 criniger . . . . . . . . . . . . . . . . . . . . . . . . . . 451, 692
uncinatus . . . . . . . . . . . . . . . . . 386, 729, 742, 756 crinitus. . . . . . . . . . . . . . . . . . . . . . . . . . . 451, 692
vethi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365 croesulus . . . . . . . . . . . . . . . . . . . . . . 464, 731, 757
viridulus . . . . . . . . . . . . . . 387, 712, 729, 742, 756 crucifer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515
viridulus . . . . . . . . . . . . . . . . . . . . . . . . . . 379, 387 cupricollis . . . . . . . . . . . . . . . . . 465, 712, 731, 757
westermanni . . . . . . . . . . . . . . . 388, 729, 742, 756 cyaneoniger . . . . . . . . . . . . . . . 466, 710, 731, 757
zambesianus . . . . . . . . . . . . . . . . . . . . . . . . . . 383 decipiens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694
ONTHOPHAGINI . . . . . . . . . . . . . 19, 394–554, 691, declivicollis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
716, 756, 758, 760 dedecor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693
Onthophagus . . . . . . . . . . 3, 4, 19, 231, 239, 314, 394, deterrens . . . . . . . . . . . . . . . . . . 467, 468, 731, 757
395, 398, 399, 401–404, 406, discretus . . . . . . . . . . . . . . . . . . . . . . 469, 731, 758
408, 414–422, 427–433, 435–445, dispar . . . . . . . . . . . . . . . . . . . . . . . . . . . 325, 691
447–472, 474–529, 531, 533–538, dubius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691
540–554, 691–697, 700, 701, 710–712, ebenicolor . . . . . . . . . . . . . . . . . 470, 731, 743, 758
730–733, 743, 744, 757–759, 765, ebenus . . . . . . . . . . . . . . . . . . . 471, 731, 743, 758
772, 773, 775, 776, 778, 779 exiguus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
absyrtus . . . . . . . . . . . . . . . . . . . . . . 443, 730, 757 fimetarius . . . . . . . . . . . . . . . . . 472, 731, 743, 758
aequepubens . . . . . . . . . . . . . . . 444, 730, 743, 757 flavolimbatus . . . . . . . . . . . . . . 474, 731, 743, 758
aeruginosus . . . . . . . . 445, 446, 710, 730, 743, 757 fritschi . . . . . . . . . . . . . . . . . . . . . . . 475, 731, 758
var. janthinus . . . . . . . . . . . . . . . . . . . . . . . . . . 445 fugitivus . . . . . . . . . . . . . . . . . . . . . . 476, 731, 758
albipennis . . . . . . . . . . . . . . . . . . . . 447, 730, 757 giraffa . . . . . . . . . . . . . . . . . . . 477, 710, 731, 758
albipodex . . . . . . . . . . . . . . . . . 448, 730, 743, 757 giuseppecarpanetoi . . . . . . . . . . . . . . . . . . 478, 758
796 SURICATA 6 (2020)

glaber . . . . . . . . . . . . . . . . . . . . . . . . . . . 415, 485 opimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 492

gnu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 otjivarongus . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
gonopygus . . . . . . . . . . . . . . . . . . . . . 479, 743, 758 ovigranosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
gracilicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . 696 pallidipennis . . . . . . . . . . 499, 512, 732, 743, 758
granulifer . . . . . . . . . . . . . 480, 712, 731, 743, 758 pallidipennis . . . . . . . . . . . . . . . . . . . . . . . . . . 512
granulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 parumnotatus . . . . . . . . . . . . . . . . . . 500, 732, 758
granum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 var. conjunctus . . . . . . . . . . . . . . . . . . . . . . . . 500
graphicus . . . . . . . . . . . . . . 442, 481, 731, 743, 758 patricius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 458
guttatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 pauxillus . . . . . . . . . . . . . . . . . . . . . . 501, 732, 758
haroldi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520 pectoralis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477
hector . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520 pedestris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
herus . . . . . . . . . . . . . . . . . . . . . . . . 482, 731, 758 pellax . . . . . . . . . . . . . . . . . . . . . . . . 503, 732, 759
hyaena . . . . . . . . . . . . . . . . . . . . . . . 483, 731, 758 phalopsides . . . . . . . . . . . . . . . . . . . . . . . . . . . 479
hybridus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696 piceus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 472
imitativus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497 pilosus . . . . . . . . . . . . . . . . . . . . . . . 503, 732, 759
immundus . . . . . . . . . . . . . . . . . . . . 484, 731, 758 plebejus . . . . . . . . . . . . . . . . . . 504, 732, 743, 759
impictus . . . . . . . . . . . . . . . . . . . . . . . . . . 450, 483 probus . . . . . . . . . . . . . . . . . . . 505, 732, 743, 759
importunus . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 politissimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 499
incertus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 691 producticollis . . . . . . . . . . . . . . . . . . 506, 732, 759
infuscatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 509 pugionatus . . . . . . . . . . . . . . . . 507, 508, 732, 759
insulsus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 var. latefulvus . . . . . . . . . . . . . . . . . . . . . . . . . . 507
intermedius . . . . . . . . . . . . . . . . . . . . . . . . . . . 696 var. quadraticornis . . . . . . . . . . . . . . . . . . . . . . 507
interstitialis . . . . . . . . . . . . . . . . . . . . 485, 731, 758 var. rufus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507
interstitialis . . . . . . . . . . . . . . . . . . . . . . . . . . . 415 pusillus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694
juvencus . . . . . . . . . . . . . . . . . . 486, 731, 743, 758 pusio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
kalaharicus . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 pusio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 692
kochi . . . . . . . . . . . . . . . . . . . . 487, 695, 743, 758 quadricallosus . . . . . . . . . . . . . . . . . . . . . . . . . . 457
lacustris . . . . . . . . . . . . . . . . . . . . . . 488, 731, 758 quadrimaculatus . . . . . . . . . . . . 510, 732, 743, 759
lamelliger . . . . . . . . . . . . . 489, 712, 731, 743, 758 quadrinodosus . . . . . . . . . . . . . . 511, 732, 743, 759
lamnifer . . . . . . . . . . . . . . . . . . . . . . 490, 731, 758 quadrinotatus . . . . . . . . . . . . . . . . . . . . . . . . . . 510
leroyi . . . . . . . . . . . . . . . . . . . . . . . . 491, 731, 758 rasipennis . . . . . . . . . . . . . . . . . 512, 732, 743, 759
leucopygus . . . . . . . . . . . . 492, 692, 731, 743, 758 rubens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
lobigena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456 rufaticollis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 507
lugens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 rufovirens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
lugens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 472 rusticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494
lugubris . . . . . . . . . . . . . . . . . . . . . . 493, 732, 758 scabrosus . . . . . . . . . . . . . . . . . . . . . . . . . 483, 696
lutulentus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493 scapularis . . . . . . . . . . . . . . . . . . . . . 513, 732, 759
macrothorax . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 schimbanus . . . . . . . . . . . . . . . . . . . . . . . . . . . 509
mactatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696 semiflavus . . . . . . . . . . . . . 514, 712, 732, 743, 759
marshalli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 semiflavus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474
merus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 semigraniger . . . . . . . . . . . . . . . . . . . . . . . . . . . 695
minutulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 senescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
minutus . . . . . . . . . . . . . . . . . . . . . . 494, 732, 758 seniculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
modestus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 setosus . . . . . . . . . . . . . . . . . . . . . . . . . . . 462, 696
monodon . . . . . . . . . . . . . . . . . . . . . . 495, 732, 758 simoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
nanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 694 stenocerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
naso . . . . . . . . . . . . . . . . . . . . . . . . . 496, 732, 758 stercorarius . . . . . . . . . . . . . . . . . . . . . . . . . . . 504
natalicus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 471 suffusus . . . . . . . . . . . . . . 515, 712, 732, 744, 759
nigrescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . 693 sugillatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
nitidulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 523 suturalis . . . . . . . . . . . . . . . . . . . . . . . . . . 499, 502
nudus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 695 talpa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 696
obesus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 522 temporalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697
obtusicollis . . . . . . . . . . . . . . . . . . . . . . . . . . . 464 tenuicornis . . . . . . . . . . . . . . . . . . . . . . . . . . . 445
obtusicornis . . . . . . . . . . . . . . . 497, 732, 743, 758 tricorniger . . . . . . . . . . . . . . . . . . . . 516, 744, 759
obtutus . . . . . . . . . . . . . . . . . . . 498, 710, 732, 758 trinodosus . . . . . . . . . . . . . . . . . . . . . 517, 732, 759
opacus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 494 trucidatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511
SURICATA 6 (2020) 797

truncaticornis . . . . . . . . . . . . . . . . . . . . . . . . . . 511 poenskopius . . . . . . . . . . . . . . . 156, 716, 720, 750

ursinus . . . . . . . . . . . . . . . . . . . . . . . 518, 732, 759 Pedaria . . . . . . . . 20, 21, 39–45, 55, 58, 690, 700, 710,
urus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 457 720, 738, 747, 766, 767, 773, 776
variolosus . . . . . . . . . . . . . . . . . . . . . . . . . 467, 759 alternans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690
ventrosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697 aspera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690
venustulus . . . . . . . . . . . . . . . . 519, 732, 744, 759 barrei . . . . . . . . . . . . . . . . . . . . . . . . . 40, 720, 747
verticalis . . . . . . . . . . . . . . . . . . 520, 732, 744, 759 brancoi . . . . . . . . . . . . . . . . . . . . . . . 41, 738, 747
vigens . . . . . . . . . . . . . . . . . . . . . . . . 521, 732, 759 conformis . . . . . . . . . . . . . . . . . . . . . . . . . . 39, 690
vinctus . . . . . . . . . . . . . . . . . . . 522, 732, 744, 759 cuprascens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
virescens . . . . . . . . . . . . . . . . . . 523, 732, 744, 759 cylindrica . . . . . . . . . . . . . . . . . . 42, 720, 738, 747
vitulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 484 picea . . . . . . . . . . . . . . . . . . . . . 43, 720, 738, 747
vylderi . . . . . . . . . . . . . . . . . . . 524, 733, 744, 759 segregis . . . . . . . . . . . . . . . . . . . . 44, 710, 720, 747
zumpti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697 sobrina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Onthotrogus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 540 somalica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 690
Outenikwanus . . . . . . . . . . . . . . 57, 149, 150, 719, 750 Phalops . . . . . . . . 4, 394, 441, 442, 525–539, 710, 712,
tomentosus . . . . . . . . . . . . . . . . 149, 150, 719, 750 733, 744, 759, 760, 766, 771
Pachylomera . . . . . . . . . . . . . . 556, 557, 572–574, 701, adspersipennis . . . . . . . . . . . . . . . . . . . 4, 529, 536
734, 745, 761, 779 ardea . . . . . . . . . . . . . . . . . . 4, 526, 733, 744, 759
femoralis . . . . . . . . . . . . . . . . . . 573, 734, 745, 761 var. chloritus . . . . . . . . . . . . . . . . . . . . . . . . 4, 526
horridus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 573 boschas . . . . . . . . . . . . . . . 527, 539, 733, 744, 759
opaca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 574 var. chloronotus . . . . . . . . . . . . . . . . . . . . . . . . 527
Pachylomerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 572 boschimanus . . . . . . . . . . . . . . . . . . . . . . . . . . 538
Pachysoma . . . . . 3, 556, 557, 575, 577–589, 595, 701, bubalus . . . . . . . . . . . . . . 528, 710, 733, 759, 771
710, 712, 716, 734, 745, 761, 772 congenitus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 529
densegranosus . . . . . . . . . . . . . . . . . . . . . . . . . 533
aesculapius . . . . . . . . . . . . 576, 577, 716, 734, 761
dregei . . . . . . . . . . . . . . . . 529, 530, 733, 744, 759
barbatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 577
euplynes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536
bennigseni . . . . . . . . . . . . . . . . 578, 734, 745, 761
flavocinctus . . . . . . . . . . . 531, 710, 733, 744, 759
denticolle . . . . . . . . . . . . . . 575, 579, 712, 745, 761
var. nigrostriatus . . . . . . . . . . . . . . . . . . . . . . . 531
endroedyi . . . . . . . . . . . . . . . . . 580, 716, 734, 761
lansbergei . . . . . . . . . . . . . . . . . . . . . . . . . . . . 536
fitzsimonsi . . . . . . . . . . . . . . . . . . . . 581, 745, 761
pauliani . . . . . . . . . . . . . . . . . . 532, 712, 744, 759
gariepinum . . . . . . . . . . . . . . . 582, 734, 745, 761
plancus . . . . . . . . . . . . . . . . . . . 533, 712, 744, 759
glentoni . . . . . . . . . . . . . . . . . . 583, 716, 734, 761
praeustus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 531
granulatum . . . . . . . . . . . . . . . . . . . . . . . . . . . 578
prasinus . . . . . . . . . . . . . . . . . . 525, 712, 744, 759
hessei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 584
var. aeraneus . . . . . . . . . . . . . . . . . . . . . . . . . . 534
hippocrates . . . . . . . . . . . . 575, 584, 710, 734, 761 pyroides . . . . . . . . . . . . . . . . . . . . . . . . . . 535, 760
marginatum . . . . . . . . . . . . . . . . . . . . . . . . . . 588 rufatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 538
macleayi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 584 rufosignatus . . . . . . . . . . . . . 4, 536, 733, 744, 760
penrithae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579 rufosignatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 529
rodriguesi . . . . . . . . . . . . . . . . . 585, 712, 745, 761 smaragdinus . . . . . . . . . . . . . . . 537, 733, 744, 760
rotundigena . . . . . . . . . . . . . . . . . . . 586, 745, 761 var. coerulosus . . . . . . . . . . . . . . . . . . . . . . . . . 537
schinzi . . . . . . . . . . . . . . . 587, 595, 716, 745, 761 var. cuprinus . . . . . . . . . . . . . . . . . . . . . . . . . . 537
striatum . . . . . . . . . . . . . . . . . . 588, 710, 734, 761 n. f. idiomelas . . . . . . . . . . . . . . . . . . . . . . . . . . 537
valeflorae . . . . . . . . . . . . . . . . . . . . . . 589, 745, 761 wittei . . . . . . . . . . . . . . . . . . . . 538, 733, 744, 760
validum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 577 zuninoi . . . . . . . . . . . . . . . . . . . . . . 539, 744, 760
Paraixodina . . . . . . 289, 310, 321–323, 727, 741, 754 Platyonitis . . . . . . . . . . . . . . . 333, 376, 389, 390, 716,
freyi . . . . . . . . . . . . . . . . . . . . . . . . . 322, 727, 754 742, 756, 766, 770, 774
saegeri . . . . . . . . . . . . . . . . . . . 323, 727, 741, 754 bicuariensis . . . . . . . . . . . 376, 390, 716, 742, 756
Parapinotis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 258 Ponerotrogus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 423
Parascarabaeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 652 Proagoderus . . . . . . . . 4, 394, 427, 429, 441, 540–554,
Parvuhowdenius . . . . . . . . . . . 58, 130, 131, 151, 152, 712, 733, 744, 760, 772, 775, 776
716, 719, 750 aciculatus . . . . . . . . . . . . . . . . . . . . . 541, 733, 760
harrisoni . . . . . . . . . . 131, 151, 152, 716, 719, 750 var. ahenus . . . . . . . . . . . . . . . . . . . . . . . . . . . 541
Peckolus . . . . . . . 57, 153–156, 710, 716, 720, 750, 772 aulicus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547
alpinus . . . . . . . . . . . . . . . 154, 710, 716, 720, 750 aureiceps . . . . . . . . . . . . . . . . . . . . . . 542, 733, 760
parvus . . . . . . . . . . . . . . . 153, 155, 716, 720, 750 bicallosus . . . . . . . . . . . . . . . . . . 543, 733, 744, 760
798 SURICATA 6 (2020)

var. olivicolor . . . . . . . . . . . . . . . . . . . . . . . . . . 543 fritschi . . . . . . . . . . . . . . . . . . . 600, 734, 745, 761

collaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 548 gracai . . . . . . . . . . . . . . . . . . . . . . . . 601, 734, 761
chalcostolus . . . . . . . . . . . . . . . . . . . . 544, 733, 760 inoportunus . . . . . . . . . . . 602, 712, 734, 745, 761
var. panchlorus . . . . . . . . . . . . . . . . . . . . . . . . 544 inquisitus . . . . . . . . . . . . . . . . . 603, 734, 745, 761
dives . . . . . . . . . . . . . . . . . . . . . . . . 545, 733, 760 intricatus . . . . . . . . . . . . . . . . . 604, 710, 735, 761
var. funereus . . . . . . . . . . . . . . . . . . . . . . . . . . 545 karrooensis . . . . . . . . . . . . . . . . 605, 734, 745, 761
var. semicuprinus . . . . . . . . . . . . . . . . . . . . . . . 545 kochi . . . . . . . . . . . . . . . . . . . . 606, 735, 745, 761
furcifer . . . . . . . . . . . . . . . . . . . 546, 733, 744, 760 lucidulus . . . . . . . . . . . . . . . . . . 607, 735, 745, 761
harpax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 548 megaparvulus . . . . . . . . . . . . . . 608, 735, 745, 762
laniger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 547 namibensis . . . . . . . . . . . . . . . . 609, 712, 745, 762
lanista . . . . . . . . . . . . . . . 547, 710, 733, 744, 760 niemandi . . . . . . . . . . . . . . . . . . . . . 610, 735, 762
loricatus . . . . . . . . . . . . . . . . . . 548, 733, 744, 760 nitidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 591
optivus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 545 obsoletepunctatus . . . . . . . . . . . . . . . 611, 746, 762
plato . . . . . . . . . . . . . . . . . . . . . . . . 549, 744, 760 pabulator . . . . . . . . . . . . . . . . . 612, 735, 746, 762
porrectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 552 parvulus . . . . . . . . . . . . . . . . . . 613, 735, 746, 762
quadrituber . . . . . . . . . . . . . . . . . . . 550, 733, 760 planipennis . . . . . . . . . . . . . . . . . . . 614, 735, 762
rangifer . . . . . . . . . . . . . . 551, 712, 733, 744, 760 reichei . . . . . . . . . . . . . . . . . . . 615, 710, 735, 762
rectefurcatus . . . . . . . . . . . . . . . 552, 712, 733, 760 rubripennis . . . . . . . . . . . 616, 712, 735, 746, 762
var. metallarius . . . . . . . . . . . . . . . . . . . . . . . . 552 soutpansbergensis . . . . . . . . . . . . . . . 617, 735, 762
sapphirinus . . . . . . . . . . . . 553, 712, 733, 744, 760 xavieri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 593
sapphyrinus . . . . . . . . . . . . . . . . . . . . . . . . . . . 542 Scarabaeus . . . . . . 4, 70, 72, 73, 85, 94, 166, 180, 184,
tersidorsis . . . . . . . . . . . . . . . . . . . . . 554, 733, 760 187, 190, 200, 205, 220, 224, 234, 246,
tubericollis . . . . . . . . . . . . . . . . . . . . . . . . . . . 547 247, 271, 274, 275, 282, 303, 314, 316,
Pseudocoptorhina . . . . . . . . . . . . . . . . . . . . . . 31, 34, 35 336, 341, 352, 357, 361, 398, 409, 410,
Pseudoniticellus . . . . . . . . . . . . . . . . . . . . . . . . 316, 320 441, 483, 525, 556, 557, 558, 560, 561,
Psilax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441 562, 564, 565, 566, 567, 568, 569, 570,
Pycnopanelus . . . . . . . . . . . . . . . 56–58, 157, 158, 701, 572, 573, 575, 580, 583, 590–593,
712, 725, 740, 750 595–598, 600, 602, 604–612, 614, 615,
krikkeni . . . . . . . . . . . . . . 158, 712, 725, 740, 750 617–652, 655, 656, 682, 684, 701,
Sarophorus . . . . . . . . . . . 20, 21, 45–55, 700, 710, 716, 710–712, 716, 735, 736, 746, 762, 763,
718, 738, 747, 771, 777 766, 768–770, 772–774, 778, 779
angolensis . . . . . . . . . . . . . . . . . . . . . . 46, 738, 747 adelphus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 632
bidentatus . . . . . . . . . . . . . . . . . 47, 716, 718, 747 alienus . . . . . . . . . . . . . . . . . . . 619, 735, 746, 762
carinatus . . . . . . . . . . . . . . . . . . . 48, 716, 718, 747 ambiguus . . . . . . . . . 620, 648, 710, 735, 746, 762
costatus . . . . . . . . . . . . . . . . 49, 52, 718, 738, 747 ambiguus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 621
diabolus . . . . . . . . . . . 50, 710, 716, 718, 747, 777 arrowi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 651
frolovi . . . . . . . . . . . . . . . . 51, 716, 718, 747, 777 basuto . . . . . . . . . . . . . . . . . . . 621, 710, 735, 762
latus . . . . . . . . . . . . . . . . . . . . . . 52, 716, 718, 747 bornemisszai . . . . . . . 622, 684, 711, 716, 735, 762
punctatus . . . . . . . . . . . . . . . . . . 53, 716, 718, 747 caffer . . . . . . . . . . . . . . . . . . . . . . . . 623, 735, 762
striatus . . . . . . . . . . . . . . . . . . . . 54, 710, 718, 747 cognatus . . . . . . . . . . . . . . . . . . 624, 712, 746, 762
tuberculatus . . . . . . . . . . . . . . . . 55, 710, 718, 747 convalescens . . . . . . . . . . . . . . . . . . . . . . . . . . 643
SCARABAEINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 convexus . . . . . . . . . . . . . . . . . . . . . . 625, 626, 762
SCARABAEINI . . . . . . . . . 19, 556–658, 716, 760, 762 costatus . . . . . . . . . . . . . . . . . . 627, 735, 746, 762
Scarabaeolus . . . . . 247, 556, 557, 590–618, 637, 701, deludens . . . . . . . . . . . . . . . . . . 628, 735, 746, 762
710, 712, 734, 735, 745, 746, 761, ebenus . . . . . . . . . . . . . . . . . . . 629, 630, 735, 762
762, 768–770, 775, 779 funebris . . . . . . . . . . . . . . 631, 637, 735, 746, 762
afronitidus . . . . . . . . . . . . . . . . . . . . 591, 745, 761 subsp. pretoriensis . . . . . . . . . . . . . . . . . . . . . . 631
anderseni . . . . . . . . . . . . . . . . . 592, 734, 745, 761 galenus . . . . . . . . . . . . . . . . . . . 632, 735, 746, 762
andreaei . . . . . . . . . . . . . . . . . . . . . . 593, 734, 761 geminogalenus . . . . . . . . . . . . . . . . 4, 633, 735, 762
bohemani . . . . . . . . . 590, 594, 710, 734, 745, 761 glabratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629
canaliculatus . . . . . . . . . . 590, 595, 615, 745, 761 goryi . . . . . . . . . . . . . . . . . . . . 634, 735, 746, 762
carniphilus . . . . . . . . . . . . . . . . 596, 734, 745, 761 heqvisti . . . . . . . . . . . . . . . . . . 635, 716, 735, 762
cicatricosus . . . . . . . . . . . . . . . . . . . . . . . . . . . 594 hottentorum . . . . . . . . . . . . . . . 636, 710, 735, 762
clanceyi . . . . . . . . . . . . . . . . . . . . . . 597, 734, 761 hottentotus . . . . . . . . . . . . . . . . . . . . . . . . . . . 647
damarensis . . . . . . . . . . . . . . . . 598, 734, 745, 761 interstitialis . . . . . . . . . . . . . . . . . . . 629, 736, 762
flavicornis . . . . . . . . . . . . 599, 710, 734, 745, 761 karae . . . . . . . . . . . . . . . . . . . . 637, 716, 736, 762
SURICATA 6 (2020) 799

laticeps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 650 caffer . . . . . . . . . . . . . . . . . . . . . . . . 676, 737, 763

leei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282 callosipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 680
natalensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 621 costatus . . . . . . . . . . . . . . 677, 683, 710, 737, 763
nepos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 634 fasciculatus . . . . . . . . . . . . . . . . 678, 710, 737, 763
pacatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 634 goryi . . . . . . . . . . . . . . . . . . . . 679, 737, 746, 763
piliventris . . . . . . . . . . . . . . . . . 638, 716, 736, 762 hirtus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 679
plausibilis . . . . . . . . . . . . . . . . . . . . . . . . . 639, 746 impressipennis . . . . . . . . . . . . . . 680, 737, 746, 763
politifrons . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629 manni . . . . . . . . . . . . . . . . . . . 675, 681, 737, 764
proboscideus . . . . . . . . . . . 640, 642, 736, 746, 762 muricatus . . . . . . . . . . . . . . . . . 682, 710, 737, 764
procles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 636 nanniscus . . . . . . . . . . . . . . . . . . 10, 683, 737, 764
profanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 634 natalensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 687
proximus . . . . . . . . . . . . . . . . . . 641, 736, 746, 762 neobornemisszanus . . . . . . 684, 711, 716, 737, 764
rixosus . . . . . . . . . . . . . . . . . . . . . . . . . . . 642, 736 ocellatus subsp. nanniscus . . . . . . . . . . . . . . . . 683
rostratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 640 oralensis . . . . . . . . . . . . . . . . . . . . . . 685, 737, 764
rugosus . . . . . . . . . . . . . . . . . . . 643, 710, 736, 762 perissinottoi . . . . . . . . . . . 686, 710, 716, 737, 764
rusticus . . . . . . . . . . . . . . . . . . . . . . 644, 736, 763 rugosus . . . . . . . . . . . . . . . . . . . . . . . . . . 663, 677
sacer var. laevigatus . . . . . . . . . . . . . . . . . . . . . . 634 sordidus . . . . . . . . . . . . . . 684, 687, 710, 737, 764
savignyi . . . . . . . . . . . . . . . . . . 645, 710, 736, 763 splendidus . . . . . . . . . . . . . . . . . . . . 688, 746, 764
schulzeae . . . . . . . . . . . . . . . . . . . . . . 646, 712, 716 transvaalensis . . . . . . . . . . . . . . . . . . . . . . . . . . 680
sericeipennis . . . . . . . . . . . . . . . . . . . . . . . . . . 629 umbraphilus . . . . . . . . . . . . . . . . . . . 689, 737, 764
spretus . . . . . . . . . . . . . . . . . . . . . . . . . . . 625, 735 Spinigymnopleurus . . . . . . . . . . . . . . . . . . . . . . . . . . 274
suri . . . . . . . . . . . . . . . . . . . . . . 647, 710, 736, 763 Sulcodrepanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
transversus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 645 Symnopleurus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
viator . . . . . . . . . . . . . . . . . . . . 648, 712, 736, 763 Tapeinopterus . . . . . . . . . . . . . . . . . . . . . . . . . . 346, 347
viator subsp. deceptor . . . . . . . . . . . . . . . . . . . 620 Tauronthophagus . . . . . . . . . . . . . . . . . . . . . . . 441, 540
vicinus . . . . . . . . . . . . . . . . . . . 649, 736, 746, 763 Tibiodrepanus . . . . . . . . . 288, 289, 324, 325, 727, 754
westwoodi . . . . . . . . . . . . . . . . 633, 650, 736, 763 sulcicollis . . . . . . . . . . . . . . . . . . . . . . 325, 727, 754
zambesianus . . . . . . . . . . . . . . . 651, 736, 746, 763 Tiniocellus . . . . 288, 289, 326–328, 727, 741, 754, 767
Sceliages . . . . . . . . . . . . . . . . . 556, 557, 652–658, 701, asmarensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
736, 746, 763, 770, 778 eurypygus . . . . . . . . . . . . . . . . . . . . . 327, 727, 754
adamastor . . . . . . . . . . . . . . . . . . . . 653, 736, 763 humilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
brittoni . . . . . . . . . . . . . . . . . . . . . . 654, 736, 763 spinipes . . . . . . . . . . . . . . . . . . 328, 727, 741, 754
curvipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653 variegatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 328
difficilis . . . . . . . . . . . . . . . . . . . . . . 655, 736, 763 Tragiscus . . . . . . . . . . . . . . 289, 329, 330, 727, 741, 754
gagates . . . . . . . . . . . . . . . . . . . . . . . 656, 736, 763 dimidiatus . . . . . . . . . . . . 329, 330, 727, 741, 754
granulatus . . . . . . . . . . . . . . . . 657, 736, 746, 763 ephippiatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 330
hippias . . . . . . . . . . . . . . . . . . . . . . . 658, 736, 763 Tropidonitis . . . . . . 333, 391, 392, 711, 716, 729, 756
iopas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653 paradoxus . . . . . . . . . . . . . 392, 711, 716, 729, 756
jopas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 653 Uposlotus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 336
microcephalus . . . . . . . . . . . . . . . . . . . . . . . . . 658 Versicorpus . . . . 58, 123, 161–163, 738, 750, 768, 769
Sebasteos . . . . . . . . . . . . . . . . . . . . 4, 618, 632, 633, 636 erongoensis . . . . . . . . . . . . . . . . 161, 162, 738, 750
Silvaphilus . . . . . . . . 57, 159, 160, 716, 720, 750, 777 streyi . . . . . . . . . . . . . . . . . . . . . . . . 163, 738, 750
oubosiensis . . . . . . . . . . . . 159, 160, 716, 720, 750 Xinidium . . . . . . . . . . . . . . . . . 58, 164, 165, 258–261,
SISYPHINI . . . . . . . . . . . . 19, 660–689, 716, 763, 764 710, 716, 726, 753, 767
Sisyphus . . . . . . . . . . 10, 660–689, 710, 711, 716, 737, curvifrons . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259
. . . . . . . . . . . . 746, 763, 764, 767, 771, 775, 776 dentilabris . . . . . . . . . . . . . . . . 258, 259, 726, 753
alveatus . . . . . . . . . . . . . . . . . . 675, 681, 746, 763 dewitzi . . . . . . . . . . . . . . . 260, 710, 716, 726, 753
bornemisszanus . . . . . . . . . . . . . . . . . . . . 684, 687 howdeni . . . . . . . . . . . . . . . . . . . . . . 261, 726, 753
 SURICATA
1. Atlas and Red List of the Reptiles of South Africa, Lesotho and Swaziland. 2013. M.F. Bates, W.R. Branch, A.M.
Bauer, M. Burger, J. Marais, G.J. Alexander and M.S. de Villiers (eds). ISBN 978-1-919976-84-6.
2. Manual of Freshwater Assessment for South Africa: Dragonfly Biotic Index. 2016. M.J. Samways & J.P. Simaika.
ISBN 978-1-928224-05-1.
3. A Bilingual Field Guide to the Frogs of Zululand. 2017. Fortunate M. Phaka, Edward C. Netherlands, Donnavan
J.D. Kruger, & Louis H. du Preez. ISBN 978-1-928224-19-8.
4. Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. 2017. A.H. Kirk-
Spriggs & B.J. Sinclair (eds). ISBN 978-1-928224-11-2.
5. Manual of Afrotropical Diptera. Volume 2. Nematocerous Diptera and lower Brachycera. 2017. A.H. Kirk-Spriggs
& B.J. Sinclair (eds). ISBN 978-1-928224-11-2.
6. Conservation assessment of Scarabaeine dung beetles in South Africa, Botswana and Namibia: IUCN Red List
categories, atlas and ecological notes. 2020. Adrian L.V. Davis, Christian M. Deschodt & Clarke H. Scholtz. ISBN
978-1-928224-39-6.

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