Chapter 19

Beetles (Insecta: Coleoptera)

Frank-T. Krell and Wolfgang Schawaller

Abstract Nine fossil beetles and seven fossil brood balls et al. 1998), Miocene Barstow Formation in California (Park
made by dung beetles are described from Laetoli (Pliocene). and Downing 2001), Eocene London Clay of Bognor Regis
Seven beetles are Tenebrionidae, tribes Tentyriini and Molurini, (Britton 1960), and particularly the Miocene of Rusinga and
one is a June beetle of the tribe Schizonychini (Scarabaeidae: Mfangano Island in Lake Victoria, Kenya (Leakey 1952;
Melolonthinae) and one a rhinoceros beetle (Scarabaeidae: Paulian 1976). These fossils give unique insights in the actual
Dynastinae) described as Calcitoryctes magnificus sp.n. Seven shape of Tertiary insects. The beetle fossils from Laetoli
fossil dung beetle brood balls are described as Coprinisphaera belong to only two families, Tenebrionidae (darkling beetles)
laetoliensis ichnosp. n. and C. ndolanyanus ichnosp. n., the and Scarabaeidae (scarab beetles). They are mineralized,
first formally described scarab ichnofossils from Africa. Two filled replicas of the exoskeleton without preservation of the
specimens of C. laetoliensis show the largest known traces of original cuticle. Despite the fairly detailed preservation, they
kleptoparasites described as Lazaichnus amplus ichnosp. n. lack crucial specific and generic characters, particularly legs,
The fossil beetles and brood balls of Laetoli weakly indicate a preventing us from formally describing most of them as new
grassland, rather than a dense woodland habitat. species, but classification at tribal level is possible.
Additionally, mineralized dung beetle brood balls are pres-
Keywords Tenebrionidae • Scarabaeidae • Ichnofossils ent, the spherical dung portions covered by a soil layer that
• Coprinisphaera coprophagous scarab beetles form in their underground nests
as food provision for their larvae. All specimens are depos-
ited in the National Museum of Tanzania, Dar es Salaam.
Introduction

With more than 350,000 described species, the beetles Tenebrionidae (W. Schawaller)
(Coleoptera) form the largest order of organisms with the
oldest stem-group representatives recorded from the Early
Permian (Ponomarenko 2002; Grimaldi and Engel 2005). As The beetle family Tenebrionidae (Darkling Beetles) is one of
hard-shelled insects they are well-represented in the fossil the largest families of Coleoptera worldwide with about
record, often, however, as single elytra or flattened sand- 20,000 recent species. The family displays high morphologi-
wiches of several layers of dark cuticle, lacking or hiding cal and ecological plasticity with species inhabiting the sea-
crucial characters (Krell 2000). Apart from amber inclusions, shore, sandy and rocky deserts, and woodland habitats up to
three-dimensional, undistorted beetle fossils are rare and the alpine zone above the timberline. In spite of this recent
known from only a few lagerstätten (Krell 2006), mainly diversity, fossil records of tenebrionids are poor. The serrate
Oligocene-Miocene Riversleigh in Queensland (Duncan antenna of one fossil from the Mesozoic Crato Formation in
Brazil might point to the family Tenebrionidae, although
definite tenebrionid family characters cannot be seen. This
F.-T. Krell (*) would be the only fossil record of a tenebrionid beetle from
Denver Museum of Nature & Science, the Mesozoic (Wolf-Schwenninger and Schawaller 2007).
2001 Colorado Blvd, Denver, CO 80205, USA
e-mail: [email protected]
Younger fossil tenebrionids are known from Tertiary depos-
its, for example from the Florissant Fossil Beds in Colorado
W. Schawaller
Staatliches Museum für Naturkunde, Rosenstein 1,
(Wickham 1914) and from the Messel deposits in Germany
D-70191, Stuttgart, Germany (Hörnschemeyer 1994). Tenebrionids from Baltic Amber are
e-mail: [email protected] listed by Spahr (1981), and tenebrionids from Dominican

T. Harrison (ed.), Paleontology and Geology of Laetoli: Human Evolution in Context. Volume 2: Fossil Hominins 535
and the Associated Fauna, Vertebrate Paleobiology and Paleoanthropology, DOI 10.1007/978-90-481-9962-4_19,
© Springer Science+Business Media B.V. 2011
536 F.-T. Krell and W. Schawaller

Amber have been described by Kaszab and Schawaller Description. Joint elytra, pronotum, dorsal part of head, and
(1984) and Doyen and Poinar (1994). venter of anterior thorax preserved. Combined elytra of oval
From Laetoli, seven remnants of beetles have been found shape, widest before the middle, length of elytra 13.5 mm, max-
that probably belong to the family Tenebrionidae. They can imal width of combined elytra 9.0 mm. Elytral surface without
be assigned to four species and are described and figured, but recognizable punctural rows or striae and without recognizable
not named. Although the fossils are well preserved, they can surface structure. Pronotum as wide as long, widest just before
be classified only by the general external morphology, but the middle, surface convex, with dense but not confluent puncta-
not by distinct generic and/or specific characters. Thus, they tion. Dorsal side of head with similar punctation as on prono-
are not formally described as new taxa. tum, clypeus without tooth or other modification, eyes somewhat
prominent, kidney-like and only slightly excavated by the genae.
Without prominent prosternal process. Anterior and middle
Tentyriini Species a (? Genus Tentyria) coxal cavities widely separated, posterior coxal cavities not pre-
served. Last abdominal ventrites not distinguishable.
EP 734/05 (Fig. 19.1) EP 1598/04 (Fig. 19.2)
Laetoli Locality 3 [localities described by Harrison and Laetoli Locality 12: Upper Laetolil Beds, between Tuffs 5
Kweka 2011]: Upper Laetolil Beds, 60–70 cm above Tuff 7. and 8.

Fig. 19.1 Tentyriini species A

(?genus Tentyria, Tenebrionidae),
Laetoli. EP 734/05; (a) right side;
(b) ventral; (c) left; (d) dorsal.
Scale in mm

Fig. 19.2 Tentyriini species A

(?genus Tentyria, Tenebrionidae),
Laetoli. EP 1598/04; (a) right
side; (b) ventral; (c) frontal; (d)
left; (e) dorsal. Scale in mm
19 Beetles 537

Fig. 19.3 Tentyriini species A (?genus Tentyria, Tenebrionidae), Laetoli. EP 1670/00; (a) right side; (b) ventral; (c) left; (d) dorsal. Scale in mm

Description. Joint elytra without tip, pronotum, head,

venter of thorax and abdominal ventrites preserved.
Combined elytra of oval shape, widest before the middle,
length of elytra 13.0 mm, maximal width of combined elytra
8.0 mm. Elytral surface without recognizable punctural rows
or striae or other recognizable surface structure. Pronotum as
wide as long, widest just before the middle, surface convex,
surface with dense but not confluent punctation. Dorsal side
of head with similar punctation as on pronotum. Clypeus
without tooth or other modification, eyes somewhat promi-
nent and kidney-shaped, only slightly excavated by the
Fig. 19.4 Tentyriini species A (?genus Tentyria, Tenebrionidae), Laetoli.
genae. Without prominent prosternal process. Anterior, mid- EP 669/04; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm
dle and posterior coxal cavities widely separated. Five visible
abdominal ventrites, ventrites 3 and 4 not distinctly shorter
than ventrite 2, last visible ventrite 5 shorter than ventrites 3 surface structure. Anterior coxal cavities not preserved, mid-
and 4 combined. dle and posterior coxal cavities widely separated. Five visible
abdominal ventrites, ventrites 3 and 4 of similar length and
EP 1670/00 (Fig. 19.3)
slightly shorter than ventrite 2, last visible ventrite 5 longer
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above
than ventrites 3 and 4 combined.
Tuff 7.
Taxonomy. These four fossils represent the same species
Description. Joint elytra without tip, venter of posterior
because of similar characters. The large body size and the
thorax and basal abdominal ventrites 1–3 preserved. Combined
general shape of pronotum and elytra, the elytra without
elytra of oval shape, widest before the middle, length of elytra
punctural rows or striae, the structure of the head with slightly
15.0 mm, maximal width of combined elytra 9.8 mm. Elytral
prominent eyes and the shape of the eyes, the wide distance
surface without recognizable punctural rows or striae and
of all coxal cavities, and the shape of the abdominal ventrites
without recognizable surface structure. Anterior coxal cavi-
coincide with recent species of the genus Tentyria Latreille,
ties not preserved, middle and posterior coxal cavities widely
1802 (tribe Tentyriini Eschscholtz, 1831, subfamily
separated. Only three basal abdominal ventrites preserved.
Pimeliinae Latreille, 1802). Quite similar is the closely
EP 669/04 (Fig. 19.4) related genus Rhytinota Eschscholtz, 1831. However, the
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above recent congeners possess a narrower pronotum, narrower and
Tuff 7. longer elytra and non-prominent eyes.
Description. Joint elytra, venter of posterior thorax and all Zoogeography. The numerous extant species of the genus
abdominal ventrites preserved. Combined elytra of oval shape, Tentyria are distributed mainly in the Mediterranean region,
widest before the middle, length of elytra 14.0 mm, maximal Arabia, and Central Asia. Species of the genus Rhytinota
width of combined elytra 8.5 mm. Elytral surface without occur today in eastern Africa and the Indian subcontinent
recognizable punctural rows or striae or other recognizable (Gebien 1937).
538 F.-T. Krell and W. Schawaller

Fig. 19.5 ?Tentyriini species B (Tenebrionidae), Laetoli, EP 351/03; (a) from left; (b) ventral; (c) right; (d) dorsal. Scale in mm

Fig. 19.6 ?Tentyriini species C (Tenebrionidae), Laetoli, EP 2777/00; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm

? Tentyriini Species B (? Genus) Taxonomy. From the general shape of the joint elytra, this
fossil might also belong to the tribe Tentyriini. However,
EP 351/03 (Fig. 19.5) because of the nearly confluent anterior coxal cavities and
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above the structure of the elytra with rows of fine punctures or striae
Tuffs 7. and with slightly convex elytral intervals, I feel unable to
Description. Joint elytra, venter of complete thorax and assign this and the following fossil specimen to any tenebrionid
all abdominal ventrites preserved. Combined elytra of oval genus, but they differ on the species level because of distinctly
shape, widest at anterior third, length of elytra 6.5 mm, max- different shape of the elytra.
imal width of combined elytra 4.0 mm. Elytral surface with
an uncertain number of rows of fine punctures or striae, ely-
tral intervals slightly convex without recognizable surface ? Tentyriini Species C (? Genus)
structure. Without prominent prosternal process. Anterior
coxal cavities touching each other, middle coxal cavities EP 2777/00 (Fig. 19.6)
slightly separated, posterior coxal cavities widely separated. Laetoli Locality 3: Upper Laetoli Beds, 60–70 cm above
5 visible abdominal ventrites, ventrites 3 and 4 of similar Tuff 7.
length and distinctly shorter than ventrite 2, last visible ven- Description. Joint elytra without tip, pronotum, venter of
trite 5 longer than ventrites 3 and 4 combined. complete thorax and all abdominal ventrites preserved.
19 Beetles 539

Combined elytra of longitudinal shape, widest at anterior Zoogeography. The genus Arturium, with few recent
third, length of elytra 11.5 mm, maximal width of combined species, is restricted to eastern Africa.
elytra 3.5 mm. Elytral surface with at least 6 rows of distinct
punctures without striae, elytral intervals nearly flat without
recognizable surface structure. Pronotum as wide as long, Biology
widest shortly before the middle, surface convex, with dense
but not confluent punctation; without prominent prosternal
process. Anterior coxal cavities touching each other, middle Recent tenebrionids belong roughly to two ecological groups.
coxal cavities slightly separated, posterior coxal cavities One group includes characteristic dwellers of dry and even
widely separated. Five visible abdominal ventrites, basal ven- arid habitats. The second, probably larger, group populates
trite 1 long, nearly as long as ventrites 2, 3 and 4 combined. decayed wood and fungi and could be considered as an indi-
Taxonomy. The general shape of the joint elytra might cator of mature forests. A compact body with short legs are
point to the tribe Tentyriini. However, as in the previous fos- characters of the tenebrionid forest dwellers, whereas slen-
sil, the nearly confluent anterior coxal cavities and the struc- der bodies with longer legs are connected with running
ture of the elytra with rows of distinct punctures prevent a behavior in open habitats, including deserts. Unfortunately,
definitive assignment to any genus. body appendages are not preserved in the Laetoli tenebrion-
ids, so no conclusions can be deduced about their former
habitat based on morphology alone.
The taxonomic assignment of some fossil tenebrionids to
Molurini Species a (? Genus Arturium) the tribes Tentyriini and Molurini clearly points to an open
habitat during the Pliocene. Nearly all recent species of these
EP 668/04 (Fig. 19.7) tribes are soil dwellers in steppes, savannahs and deserts, and
Laetoli Locality 3: Upper Laetolil Beds, 60–70 cm above today eastern Africa is populated with abundant elements of
Tuff 7. these tenebrionid tribes. The fossil tenebrionids from Laetoli
Description. Joint elytra, venter of posterior thorax pre- give no hint of woodland or forested habitats.
served. Combined elytra oval shaped, widest at anterior third,
length of elytra 13.0 mm, maximal width of combined elytra
9.0 mm. Elytral disc with two high undulated keels and with
a somewhat lower humeral keel, medial part of elytral Scarabaeidae (F.-T. Krell)
between the humeral keels flat, lateral parts of elytra besides
the humeral keels vertical and not to be seen in dorsal view, With about 31,000 described extant species (Jameson and
surface between the keels slightly uneven und densely punc- Ratcliffe 2002) the Scarabaeoidea are one of the largest super-
tured. Middle and posterior coxal cavities widely separated. families in the Coleoptera. They are distributed world-wide
Abdominal ventrites not preserved. and comprise such varied groups as dung beetles, stag beetles,
Taxonomy. From the larger body size and the dorsal struc- and chafers, ranging from just over 1 mm to 170 mm body
ture of the elytra the fossil might be assigned to the genus length. In the fossil record they are fairly well represented
Arturium Koch, 1951 (tribe Molurini Latreille, 1829, sub- with about 230 species described from the Upper Jurassic to
family Pimeliinae Latreille, 1802), although further characters the Pleistocene (Krell 2007). However, apart from fossil
cannot be compared. brood balls made by dung beetles, Pliocene scarab fossils are

Fig. 19.7 Molurini species A (?genus Arturium, Tenebrionidae), Laetoli, EP 668/04; (a) from right; (b) ventral; (c) left; (d) dorsal. Scale in mm
540 F.-T. Krell and W. Schawaller

rare (Krell 2007) with some extant species recorded, but only Dynastinae: Oryctini/Pentodontini
two extinct species described: the dung beetle Copris kartli-
nus Kabakov, 1988, from the Kisatibi Formation in Georgia, Calcitoryctes Krell, gen.n.
and the dubious Melolonthites laterosinuatus Piton and
Derivatio nominis. From calcite (calcium carbonate) of which
Théobald 1935, from the Mio/Pliocene cinerites of Varennes,
the specimen consists, and Oryctes, the type genus of Oryctini
France, represented by only one elytron.
Mulsant, 1842, to which it might belong. Gender masculine.
Scarabaeoidea is a monophyletic group diagnosed by two
Type species. Calcitoryctes magnificus sp.n.
autapomorphies: antenna with a lamellate club and anterior bor-
Diagnosis. Outer side of mandibles entire or slightly den-
der of hind wings with sclerotized field proximal to a pinch, as
ticulate. Clypeus truncated with rounded angles. Head with
part of a spring folding mechanism (Krell 2006). Both charac-
tubercle. Pronotum without sculpture. Ventrites 1–4 much
ters are rarely preserved in fossils and are missing in the Laetoli
shorter than 5 and 6. Pygidium ca. three times as broad as
scarabs. Most fossil scarab beetles, including the two specimens
long, transversally bulged with strong apical impressions on
described below, were identified on the basis of other characters
both sides. Some extant Oryctes Illiger, 1798, and Cyphonistes
typical for Scarabaeoidea, such as enlarged prothorax adapted
Burmeister, 1847, species have a similar pygidium, but
for digging with short and powerful legs with tibiae toothed
Oryctes is much larger and has a deeply emarginated clypeus
along outer edge, large and narrowly separated to contiguous
and Cyphonistes has never one tubercle on the head (Endrődi
pro- and mesocoxae, and transverse and narrowly separated to
1985). Within Pentodontini Mulsant, 1842, it resembles the
contiguous metacoxae. The body fossils from Laetoli are well-
South/East African genus Pentodontoschema Péringuey,
preserved three-dimensionally, but are lacking finer surface
1901 (Péringuey 1901; Ferreira 1966) from which it differs
structures like punctation. In the melolonthine, even all sutures
by the margined base of the pronotum, the concave apical
are blurred. Moreover, the legs apart from some femora are
part of the pygidium and the probably less denticulated man-
missing, hardly leaving any genus- or species-diagnostic char-
dibles. It differs from Heteroligus Kolbe, 1900, by the miss-
acters. Both specimens are described, but only the exceptionally
ing pronotal tubercles and the concave apical part of the
preserved rhinoceros beetle is named.
pygidium and from Phyllognathus Eschscholtz, 1830, by the
Apart from the two body fossils, seven fossil dung beetle
more slender mandibles (Endrődi 1985).
brood balls were found at Laetoli. Fossil brood balls were
recorded from Laetoli previously (Sands 1987) and also from
Calcitoryctes magnificus Krell, sp.n.
other Pliocene sites in Africa, such as paleo-lake Chad
(Duringer et al. 2000) and Makapansgat Limeworks in South Derivatio nominis. Magnificus (adj.) (post-classical
Africa (Kitching 1980), as well as from the Pleistocene of Latin) = magnificent.
Rutana in Burundi (Basilewsky 1951). However, none of the Holotype. EP 2704/00 (Fig. 19.8), Laetoli Locality 2:
African specimens has been formally described and named Laetolil Beds, upper unit between Tuffs 5 and 7, deposited in
as ichnospecies. the National Museum of Tanzania, Dar es Salaam.

Fig. 19.8 Calcitoryctes magnificus sp.n. (Scarabaeidae, Dynastinae), holotype, Laetoli, EP 2709/00; (a) from frontal; (b) right; (c) dorsal; (d) left;
(e) ventral; (f) caudal. Scale in mm
19 Beetles 541

Description. Complete body with all femora apart from Classification. The only character separating the tribes
the right profemur; length: 18.4 mm; maximum width of Pentodontini and Oryctini in the current typological classifi-
pronotum: 9.5 mm; maximum width of elytra: 10.5 mm. cation is the tibial apex. Tibiae are not preserved in this spec-
Body with left profemur and all other femora present. imen. However, since the tibial apex is variable within tribes
Mandibles protruding beyond clypeus laterally and anteri- and the tribes themselves are only typologically defined
orly; outer side of mandibles entire or slightly denticulate, (Ratcliffe 2003:249), both groups might not survive a phylo-
basally broad, slightly emarginated to a relatively sharp tip, genetic analysis as equally ranked taxa as one might become
possibly with a denticle in the middle. Labium basally a subgroup of the other. Therefore, it is not a serious short-
broadly rectangular, apically with a deep emargination before coming that Calcitoryctes cannot be unequivocally assigned
the double-convex anterior border, incised in the middle to one of these groups.
(Fig. 19.9). Only scape and pedicel of the left antenna pres-
ent, both short as typical for Dynastinae. Clypeus truncated
with rounded angles, anterior margin bluntly triangular, lat-
eral margins slightly emarginated. Head with median conical Melolonthinae: Schizonychini, Species a
tubercle, steeper declined posteriorly than anteriorly; from
the tubercle, a slight bulge extends to the sides. Short ocular EP 2156/03 (Fig. 19.9)
canthus present. Pronotum regularly convex without any Laetoli Locality 7: Laetolil Beds, upper unit between
sculpture; lateral margins regularly convex, broadest in the Tuffs 5 and 7 [body, one femur]
middle; laterally and basally margined, lateral margin broader Description. Length: 12.4 mm; maximum width of pro-
and sharp. Anterior angles protruding and posterior angles notum: 5.0 mm; maximum width of elytra: 6.1 mm; height
obtuse. Anterior margin without tubercle. Basal margin con- of specimen: 5.1 mm. No microsculpture or punctation vis-
vex. Scutellum rounded. Elytra broadest just behind the mid- ible due to preservation. Labrum medianly incised,
dle. Sutural interval basally broad, apically tapered and Prementum bilobate, with a median furrow extending to
slightly elevated. Shallow traces of probably three discal mentum. Clypeus seems to be separated from the frons by a
stripes visible. Humeral callus and anteapical callus present. slight transversal bulge. Pronotum: Sides slightly diverging
Epipleura narrow (maximum 0.65 mm), with sharp border, in basal third, then straight strongly converging to head.
extending to pygidium. Ventrites 1–4 short, ventrite 5 longer Elytra: Lateral border behind the humeral callus slightly
than 3 + 4 together, ventrite 6 as long as 3–5 together. All emarginate, then elytra broadened, widest just behind the
coxae touching. Hind femora much thicker than middle and middle. All coxae adjacent.
slightly thicker than fore femora (ratio length/width f: 2.1; Classification. The elevated frontoclypeal suture and the
m: 2.6; h: 1.9). Propydidium not fully covered by elytra, not strongly incised labrum are characters of Schizonychini
produced posteriorly; no stridulatory area visible due to pres- Burmeister, 1855, and, together with the shape of the elytra,
ervation. Pygidium short (3.1 × as broad as long) as in most might even indicate that the specimen belongs to Schizonycha
Oryctini. Basal half of pygidium forms a strongly convex (Pope 1960; Lacroix 1989), which is a speciose genus
transversal bulge (visible from ventral), apical half impressed containing abundant African species. However, two crucial
and convex, with strong impressions on both sides, margin of characters for Schizonychini, the enlarged ventrite 6 and the
the tip of the pygidium slightly protruding. metepimeron (Lacroix 1989), are not sufficiently preserved

Fig. 19.9 Melolonthinae, Schizonychini species A (Scarabaeidae), Laetoli, EP 2156/03; (a) from right; (b) ventral; (c) frontal; (d) left; (e) dorsal.
Scale in mm
542 F.-T. Krell and W. Schawaller

to allow a reliable tribal classification. Preservation of antennae, thickness of the outer layer is not only an indication of the
claws, mouthparts and setation would be necessary for thickness of a possible soil cover, but also of the amount of
generic classification. dung still in existence at the time when the dung consump-
tion by the larva ended. The longer the larva feeds, the thin-
ner the walls become (Lengerken 1954). Successful
development of the larva and hatching of the beetle results in
Ichnofossils thin walls of the brood ball. Thick walls might indicate that
the development was disturbed or unsuccessful.
A small and a large type of fossil dung beetle brood balls
were found at Laetoli, similar to the abundant brood balls Coprinisphaera laetoliensis Krell, ichnosp. n.
described as Coprinisphaera Sauer, 1955, from South Derivatio nominis. Adjective meaning ‘from Laetoli’, the
America. Genise et al. (2000) have already assigned the type locality.
“structure resembling a dung ball of Heliocopris” mentioned
by Sands (1987: 423) to this ichnogenus and counted Laetoli
in the localities with Coprinisphaera ichnofacies. All for-
mally described scarab ichnofossils are from the Americas
(Krell 2007). The fossil dung beetle brood balls were recently
revised by Genise (2004) and Laza (2006) and classified into
nine ichnospecies. To facilitate the integration of the Laetoli
ichnofossils into the current ichnological classification, they
will be named although ichnospecies-specific characters are
often not clearly visible due to the replacement of original
material and infilling with calcium carbonate.
These balls represent the prepared portions of resources
that dung beetles form from available feces as food provision
for their larvae. The brood balls of large tunnelling dung Fig. 19.10 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood
ball, Laetoli, holotype, EP 224/04. Cut; (a) cut half; (b) counterpart
beetles are generally covered by a layer of soil to prevent
desiccation (Halffter and Edmonds 1982), whereas the sur-
face of brood balls of large rollers such as Scarabaeus L.,
1758 or Kheper Janssens, 1940 is either only smoothened
(Lengerken 1954) or covered by the mother beetle’s feces
(Sato and Imamori 1987).
Whereas the American fossil brood balls are mostly hol-
low spheres with the original soil cover preserved, the Laetoli
fossils are steinkerns of calcium carbonate rendering it
impossible to identify whether the outer layer was originally
formed by soil or dung. I interpret the faint line between the
outer layer and the infilling (Figs. 19.10–19.12) as the border Fig. 19.11 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood
between the original dung portion and larval chamber. The ball, Laetoli, EP 1719/03. Cut; (a) half cut; (b) counterpart

Fig. 19.12 Lazaichnus amplus isp.n., traces of kleptoparasites in fossil dung beetle brood ball, Laetoli. Holotype of L. amplus in paratype of
Coprinisphaera laetoliensis, EP 1719/03; (a) half cut; (b) counterpart; (c) outer opening of L. amplus
19 Beetles 543

opening (55.5 mm × 38.8 mm) on one side and a deep small

hole (10 mm diameter) on the other. Height: 48 mm; equato-
rial diameter: 60 mm. Filled with a lighter (replaced dung)
and a darker substance in the area of the small hole and prob-
ably indicating the development space of a kleptoparasite
(paratype of Lazaichnus amplus ichnosp. n., described
below). Due to the large, irregular opening the ichnospecies-
specific collar is missing. Thus, this specimen is not desig-
nated as a paratype of C. laetoliensis.
Biology. The large upper opening (see Fig. 19.14) might
have been caused by a vertebrate predator. Modern exam-
Fig. 19.13 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood ples of brood balls opened by bat-eared foxes (Otocyon
ball, Laetoli, holotype, EP 224/04. Before cut; (a) from top; (b) lateral.
megalotis (Desmarest, 1822)) show similarly shaped holes
Scale in mm
(see Fig. 19.15). These foxes excavate dung beetle brood
balls on a regular basis to prey on the larvae (Nel and Maas
Holotype. EP 224/04 (Figs. 19.10, 19.13), Laetoli Locality 2004).
15: Upper Ndolanya Beds; deposited in the National Museum EP 1719/03, Specimen A (Figs. 19.11, 19.16), Laetoli
of Tanzania, Dar es Salaam. Locality 15: Upper Ndolanya Beds, cut medianly in half;
The specimen is the fossilized empty, abandoned brood flat pear-shaped; height: 48.5 mm, equatorial diameter:
ball, replaced and filled by calcium carbonate. Cut medianly 58.6 mm, maximum width of neck: 27 mm; bottom flat with
in half. Maximum height: 53.4 mm; equatorial diameter: central round area of ca. 14 mm diameter (contact zone of
55.5 mm; diameter of upper opening: 22.5 mm. Regular brood ball to ground); lateral thickness of the wall 6–8.5 mm,
sphere with wide upper opening surrounded by collar in form 4–10 mm at the bottom; no signs of kleptoparasitic or sec-
of a flat bulge (11–12 mm wide). Bottom with a small, oval ondary intrusion. Surface with six dorsoventral ribs which
depression of about 13 mm diameter. upper lateral thickness were possibly caused by filled cracks of the surrounding
of the wall: ca. 4.5 mm, about 10 mm at the bottom. soil. Because of the different shape (rather pear-shaped than
Biology. Thin wall, broad opening and undamaged outer spherical), which is similar of the shape of Vondrichnus
surface indicate that the beetle hatched successfully; no signs planoglobus Duringer et al., 2007 (Fig. 19.10a, d; termite
of kleptoparasitic or secondary intrusion. nest), it is not designated as a paratype. Because a thick
Paratype. EP 1719/03, Specimen B (see Fig. 19.12) (of 3 outer wall can be distinguished from the inner chamber, and
specimens), Laetoli Locality 15: Upper Ndolanya Beds; because it forms part of a series of three fossil brood balls
deposited in the National Museum of Tanzania, Dar es likely found together (the other two being typical
Salaam. Cut medianly in half, some outer cracks fixed by Coprinisphaera), it is identified as Coprinisphaera. Since
glue; spherical with upper opening; height: 47.5 mm, equa- the three balls were not found in close proximity, I do not
torial diameter: 54 mm; opening: 22 mm; thickness of the classify them as Quirogaichnus Laza, 2006 which was intro-
wall: 10.5–15.8 mm, bottom with two deep holes of 9–10 mm duced for clusters of brood balls in a distinct chamber or
diameter (ventral hole: Fig. 19.12a, b; ventrolateral hole: cavity.
Fig. 19.12c). Biology. The thin walls and undamaged outer surface
Biology. The size of the holes in the outer wall indicate indicate a regular development of the larva. However, an
infestation by larger kleptoparasites of genera such as upper opening is not clearly defined. The irregular inter-
Onthophagus Latreille, Hyalonthophagus Palestrini or nal part of the neck could be the opening (diameter
Pedaria Laporte, being the holotype of Lazaichnus amplus 13 mm).
ichnosp.n. (described below). Kleptoparasitic species invade EP 1719/03 Specimen C, Laetoli Locality 15: Upper
and lay their eggs in a dung portion collected by another spe- Ndolanya Beds: flat pear-shaped with flat bottom; height:
cies, generally reducing the reproductive success of the host 47.9 mm; equatorial diameter: 60.6 mm; surface with large,
significantly (Rougon and Rougon 1980; Gonzáles-Megías deep cracks; no indication of kleptoparasitic or secondary
and Sánchez-Piñero 2003). The thick walls show that the intrusion. Because of this suboptimal infilling or incipient
dung material had not been exhausted, probably due to disintegration this specimen was not cut. Due to poor preser-
unsuccessful or abbreviated development of the host larva. vation, I do not designate it as a paratype.
The upper opening might indicate that the host larva might Description. Spherical, solid structures with sometimes
have hatched. flattened underside resulting in a slight pear-like shape;
Additional material. EP 1077/01 (see Fig. 19.14), Laetoli height: 47.5–53.4 mm; equatorial diameter: 54.0–55.5 mm
Locality 15: Upper Ndolanya Beds. Flat sphere with large (all specimens: 53.4–60.6 mm). Dorsal opening on top with
544 F.-T. Krell and W. Schawaller

Fig. 19.14 Lazaichnus amplus isp.n., traces of kleptoparasites in fossil view in large opening showing the dark internal trace of L. amplus.
dung beetle brood ball, Laetoli. Paratype of L. amplus in C. laetoliensis, Scale (for Fig. 19.14a–c) in mm
EP 1077/01; (a) from top; (b) bottom; (c) lateral; (d) outer opening; (e)

Fig. 19.15 Recent dung beetle brood balls excavated and preyed on by bat-eared foxes (Otocyon megalotis) in the Laetoli area, showing an irregu-
lar large opening. Scale: 30 mm. Specimens deposited in National Museum of Tanzania, Dar es Salaam. Photo: T. Harrison

surrounding bulge-like collar. Outer wall of sphere between Diagnosis. Spherical to slightly pear-shaped structure
4.5 and more than 15 mm thick. Internal (filled) chamber with single chamber and upper opening with neck; differs
opens directly to the upper aperture. from Coprinisphaera kraglievichi (Roselli, 1939) from
19 Beetles 545

Fig. 19.16 Coprinisphaera laetoliensis isp.n., fossil dung beetle brood ball, Laetoli, EP 1719/03. Before cut; (a) from top; (b) bottom; (c) lateral.
Scale in mm

Uruguay and Argentina by the larger size (more than 53 mm

equatorial diameter versus 32–42.4 mm).
Discussion. The size of dung beetle brood balls varies
intraspecifically, but generally stays within 20–30%
(Lengerken 1954; Klemperer and Boulton 1976; Klemperer
1983; Sato and Imamori 1987). A brood ball of 60.6 mm
diameter is unlikely to belong to the same species as one of
32 mm diameter. Although, according to a recently published
convention (Bertling et al. 2006), the possible tracemaker
should not be considered as defining character for ichnospe-
cies, I consider it inappropriate to typologically combine
specimens under one ichnospecific name that are most likely
produced by different tracemaker species.
Trace maker. Sands’s (1987) interpretation of a
Coprinisphaera from Laetoli as resembling a brood ball of
Heliocopris Hope might be right. The large size and the
shape (spherical to slightly pear shaped) of the brood balls
are typical for extant Heliocopris (cf. Klemperer and Boulton
1976), but could also be produced by large Catharsius Hope
species or even by large rollers such as Kheper Janssens
(Sato and Imamori 1987) in which the mother beetle scrapes
the brood balls into a spherical shape during the development
of the larva.
Nomenclatural note. Coprinisphaera Sauer, 1955 is a junior
subjective synonym of Fontanai Roselli, 1939 (Laza 2006), but
is in prevailing usage (Krell 2007). Genise et al. (2006) and
Krell (2008) proposed conservation of Coprinisphaera with
the International Commission on Zoological Nomenclature.
The ICZN (International Commission on Zoological
Nomenclature 2008) has ruled that Coprinisphaera is to be
maintained as the valid name.
Coprinisphaera ndolanyanus Krell, ichnosp. n.
Fig. 19.17 Coprinisphaera ndolanyanus isp. n., fossil dung beetle brood
Derivatio nominis. Adjective meaning “belonging to ball, Laetoli. Holotype, EP 824/01. Before cut; (a) lateral; (b) from top.
Ndolanya”, the stratigraphic unit where it was found. Scale in mm
Holotype. EP 824/01 (Fig. 19.17), Laetoli Locality 18:
Upper Ndolanya Beds; deposited in the National Museum of cut medianly in half. Pear-shaped (sphere with neck on top),
Tanzania, Dar es Salaam; maximum height: 33.7 mm; tip of neck slightly impressed. Homogenous steinkern; no
equatorial diameter: 27.1–27.6 mm, width neck: 10.5–12 mm; internal structures visible.
546 F.-T. Krell and W. Schawaller

holotype of C. kheprii only. The equatorial diameter of the

smallest C. kheprii would fall into the specific size range of
C. ndolanyanus, but the minimum height of C. kheprii is
50 mm, much larger than in the latter.
Trace maker. Pear-shaped brood balls are typical for dung
rollers (Scarabaeini, Gymnopleurini) (Lengerken 1954) that
are abundant in recent Afrotropical coprocenoses.

Lazaichnus amplus Krell, ichnosp. n.

Derivatio nominis. Amplus (Latin) = vast, spacious;
adjective.
Holotype. EP 1719/03, Specimen B (Fig. 19.12) (para-
type of Coprinisphaera laetoliensis ichnosp.n.), Laetoli
Locality 15: Upper Ndolanya Beds; deposited in the National
Museum of Tanzania, Dar es Salaam. Coprinisphaera is cut
Fig. 19.18 Coprinisphaera ndolanyanus isp. n., fossil dung beetle medianly in half, hitting one hole of Lazaichnus amplus
brood ball, Laetoli. EP 3335/00. Scale in mm
(11.4 mm diameter). The other hole has an outer diameter of
8–11 mm.
Paratype. EP 1077/01 (Fig. 19.14) (Coprinisphaera lae-
Additional Material. EP 3335/00 (Fig. 19.18), Laetoli
toliensis), Laetoli Locality 15: Upper Ndolanya Beds; depos-
Locality 18: Upper Ndolanya Beds; maximum height:
ited in the National Museum of Tanzania, Dar es Salaam.
30.0 mm; equatorial diameter: 27.5–28.4 mm; cut medianly
The chamber behind the opening of Lazaichnus is visible in
in half. Sphere, one half with smooth surface, damaged on
the large upper opening in Coprinisphaera. Hole diameter:
the other side (possible neck missing), next to damage with
14–15.3 mm with lateral lighter infilling, diameter of proper
oval depression; no internal structures visible.
hole: 8.5 mm.
Description. Pear-shaped solid structure with upper impres-
Description. Holes in fossil dung beetle brood balls of
sion in long neck, probably indicating a former opening;
8–15.3 mm diameter, with a simple gallery or an expanding
height: 30.0–33.7 mm; equatorial diameter: 27.1–28.4 mm;
chamber extending into the inner core of the brood ball.
neck diameter: 10.5–12.0 mm. No internal structure visible.
Diagnosis. Holes leading to a gallery in fossil dung beetle
Diagnosis. Pear shaped solid structure; differs from
brood balls (Coprinisphaera), differ from the only other spe-
Coprinisphaera kheprii Laza, 2006 (height of holotype
cies of the ichnogenus, Lazaichnus fistulosus Mikulaš and
56.1 mm) from Argentina by the smaller size and the longer,
Genise, 2003, by the larger diameter of the opening and the
more slender upper protuberance. The upper protuberance of
simple structure of the gallery, which forms a simple tube or
C. tonnii Laza, 2006 is thicker and more bulge-like than in
a chamber in L. amplus, but often a more complex system of
C. ndolanyanus, containing an egg chamber. Coprinisphaera
tubes in L. fistulosus.
ndolanyanus differs from the globular C. laetoliensis from
Trace maker. The large diameter of the holes indicates
the same locality by the smaller size and the shape.
kleptoparasites much larger than Cleptocaccobius Cambefort
Discussion. A distinct outer wall of the brood balls is the
or kleptoparasitic Aphodius Illiger (the most abundant repre-
diagnostic character that distinguishes Coprinisphaera Sauer
sentatives of this guild in Africa). Current Afrotropical klep-
(dung beetle brood balls) from Pallichnus Retallack, 1984
toparasites of a matching size belong to Onthophagus,
(supposedly pupal chambers). Neither specimen of C. ndola-
Hyalonthophagus and Pedaria (Scarabaeinae).
nyanus shows this character due to the type of preservation
(original substance replaced by calcium carbonate). However,
the pear-shaped form of the holotype is an unequivocal indi-
cation of scarabaeine brood balls (Halffter and Edmonds
1982) which originally might not have had a different outer Biology
wall structure than C. laetoliensis. Coprinisphaera ndolan-
yanus is much smaller than the holotype of the pear-shaped Schizonychini, Dynastinae, dung beetles and their kleptopar-
C. kheprii, but other known specimens assigned to C. kheprii asites are distributed in all vegetation zones, from tropical
have an equatorial diameter from 28.5 to 60.5 mm. This size rain forest to arid regions. The same is true for the paleoen-
range is much larger than the 20–30% size range of brood viroments of the Coprinisphaera ichnofacies, but most are
balls of a single dung beetle species (see above). Thus, I associated with open grasslands (Genise et al. 2000).
compared the size of C. ndolanyanus with the size of the Schizonychini feed on leaves as adults and on roots as larvae.
19 Beetles 547

Dynastinae feed on various living and rotting plant material. Doyen, J. T., & Poinar, G. O. (1994). Tenebrionidae from Dominican
Thus, the scarab fossils and ichnofossils do not contradict amber. Entomologica Scandinavica, 25, 27–51.
Duncan, I. J., Briggs, D. E., & Archer, M. (1998). Three-dimensionally
the current hypothesis of a savanna-forest ecotone paleoenvi- mineralized insects and millipedes from the Tertiary of Riversleigh,
ronment at Laetoli (Kingston and Harrison 2007; Su and Queensland, Australia. Palaeontology, 41, 835–851.
Harrison 2007). Duringer, P., Brunet, M., Cambefort, Y., Likius, A., Mackaye, H. T.,
Schuster, M., & Vignaud, P. (2000). First discovery of fossil dung
beetle brood balls and nests in the Chadian Pliocene australopithe-
cine levels. Lethaia, 33, 277–284.
Conclusions Duringer, P., Schuster, M., Genise, J. F., Mackaye, H. T., Vignaud, P., &
Brunet, M. (2007). New termite trace fossils: Galleries, nests and fungus
combs from the Chad basin of Africa (Upper Miocene–Lower Pliocene).
Laetoli is one of the few lagerstätten where insect fossils are Palaeogeography, Palaeoclimatology, Palaeoecology, 251, 323–353.
preserved three-dimensionally. Whereas reconstructions are Endrődi, S. (1985). The Dynastinae of the world. Budapest: Akadémiai
Kiadó.
the only way to visualize the body shape of common fossil Ferreira, M. C. (1966). Contribuição para o estudo dos dinastíneos afri-
imprints, the Laetoli fossils are almost undistorted replicas of canos V. Os dinastíneos da região etiópica. Revista de Entomologia
the original bodies which made formal description and naming de Moçambique, 8, 3–348.
of an exceptionally character-rich rhinoceros beetle fossil Gebien, H. (1937). Katalog der Tenebrioniden (Col. Heteromera) Teil I.
Pubblicazioni del Museo Entomologico “Pietro Rossi” Duino, 2,
possible. External morphology of beetles is often a reliable 505–883.
indicator of habits, habitats or soil types (e.g., Medvedev 1965). Genise, J. F. (2004). Ichnotaxonomy and ichnostratigraphy of cham-
Adaptive traits are most clearly expressed in the legs, the body bered trace fossils in palaeosols attributed to coleopterans, ants and
parts interacting most extensively with the physical environ- termites. In D. McIlroy (Ed.), The application of ichnology to palaeo-
environmental and stratigraphic analysis (pp. 419–453). London:
ment. However, with legs missing or only partly preserved the Geological Society.
beetle fossils of Laetoli do not indicate a particular habitat. Genise, J. F., Mángano, M. G., Buatois, L. A., Laza, J. H., & Verde, M.
Extant dung beetles producing brood balls of the size found (2000). Insect trace fossil associations in paleosols: The
at Laetoli are distributed in subtropical and tropical regions. In Coprinisphaera ichnofacies. Palaios, 15, 49–64.
Genise, J. F., Laza, J. H., & Rindsberg, A. K. (2006). The ichnogenus
the Afrotropics, the highest abundance of dung beetles is in Coprinisphaera Sauer 1955 (Ichnotaxa, Insecta, Coleoptera,
savannas (Cambefort and Walter 1991), where we also find the Coprinisphaeridae): Proposed conservation. Bulletin of Zoological
highest density of kleptoparasitic dung beetles (Krell et al. Nomenclature, 63, 243–246.
2003). The existence of large fossil dung beetle brood balls in Gonzáles-Megías, A., & Sánchez-Piñero, F. (2003). Effects of brood
parasitism on host reproductive success: Evidence from larval inter-
Laetoli, some with traces of kleptoparasites, indicates a higher actions among dung beetles. Oecologia, 134, 195–202.
probability for grassland rather than an arboreal area. Grimaldi, D., & Engel, M. S. (2005). Evolution of the insects. New
York: Cambridge University Press.
Acknowledgements We are grateful to Terry Harrison, New York Halffter, G., & Edmonds, W. D. (1982). The nesting behavior of dung
University, for the patient loan of the specimens, to Harry Taylor, Photo beetles (Scarabaeinae). An ecological and evolutive approach.
Unit of The Natural History Museum, London, for most of the photo- México: Instituto de Ecología.
graphs, to Brett Ratcliffe, University of Nebraska State Museum, Harrison, T., & Kweka, A. (2011). Paleontological localities on the
Lincoln, and Roger-Paul Dechambre, Muséum national d’Histoire Eyasi Plateau, including Laetoli. In T. Harrison (Ed.), Paleontology
naturelle, Paris, for helpful comments about the fossil dynastine, and to and geology of Laetoli: Human evolution in context (Geology,
Jorge Genise, Museo Paleontológico Egidio Feruglio, Trelew, Argentina, geochronology, paleoecology and paleoenvironment, Vol. 1,
for careful criticism on a former version of the manuscript. Ken pp. 17–45). Dordrecht: Springer.
Carpenter and Thomas Garner, DMNS Earth Sciences Department, Hörnschemeyer, T. (1994). Ein fossiler Tenebrionidae Ceropria? messel-
helped with cutting the fossil brood balls. ense n. sp. (Coleoptera: Tenebrionidae: Diaperinae) aus dem Mitteleozän
der Grube Messel bei Darmstadt. Courier Forschungsinstitut
Senckenberg, 170, 75–83.
International Commission on Zoological Nomenclature (2008). Opinion
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