Aballayetal 2016

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An Illustrated Key to, and Diagnoses of the Species of Tenebrionidae

(Coleoptera) Associated with Decaying Carcasses in Argentina
Article in Annales Zoologici · December 2016

DOI: 10.3161/00034541ANZ2016.66.4.021
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A N N A L E S Z O O L O G I C I (Warszawa), 2016, 66(4): 703-726
AN ILLUSTRATED KEY TO, AND DIAGNOSES OF

THE SPECIES OF TENEBRIONIDAE (COLEOPTERA)
ASSOCIATED WITH DECAYING CARCASSES IN ARGENTINA
FERNANDO H. ABALLAY1,*, GUSTAVO E. FLORES1, VIOLETA A.

SILVESTRO2, NOELIA I. ZANETTI3,4 and NÉSTOR D. CENTENO4
1Laboratorio de Entomología, Instituto Argentino de Investigaciones de las Zonas

Áridas (IADIZA, Centro Científico Tecnológico CONICET Mendoza), Casilla de
correo 507, 5500 Mendoza, Argentina.
2
Laboratorio de Entomología, Instituto de Biodiversidad y Biología
Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos
Aires (UBA), e Instituto de Biodiversidad y Biología Experimental y Aplicada
(IBBEA), CONICET-UBA, Buenos Aires, Argentina;
3Cátedra de Parasitología Clínica, Departamento de Biología, Bioquímica y
Farmacia, Universidad Nacional del Sur, San Juan 670, Bahía Blanca 8000,
Provincia de Buenos Aires, Argentina.
4Laboratorio de Entomología Aplicada y Forense, Universidad Nacional de
Quilmes, Roque Sáenz peña 180, B1876BXD, Bernal, Buenos Aires, Argentina
*
Corresponding author: e-mail: [email protected]
Abstract.— In this study we identified adults of Tenebrionidae associated mainly with pig
carcasses and human cadavers lying in different arid and semiarid areas of Argentina. This
article provides an illustrated key to 29 Tenebrionidae species of the potential forensic
importance, accompanied by diagnoses, and habitus photographs of these taxa. The
recorded species of darkling beetles belong to four subfamilies: Alleculinae (one species of
Alleculini), Diaperinae (two species of Crypticini), Tenebrioninae (15 species, ten belonging
to the tribe Scotobiini, three to Opatrini, two to Alphitobiini and one species of Tenebrionini),
and Pimeliinae (10 species, three belonging to the tribe Epitragini, two to Evaniosomini, two
to Praociini, one species from each of the tribes Trilobocarini, Edrotini and Elenophorini).
A new species of Eutelocera Solier (Pimeliinae: Praociini) collected from decomposing pig
carcasses is described: E. cadaverina Flores and Zanetti sp. nov. Two species are new
records for Argentina: Conibius (Gondwanodilamus) franzi Kaszab and Alphitobius
laevigatus (Fabricius); three species for Bolivia: Blapstinus punctulatus Solier, Salax
lacordairei Guérin-Méneville, and Vaniosus profana (Kulzer); and one species for
Uruguay: Blapstinus punctulatus Solier. A discussion is presented on the potential
forensic importance of some species collected from human cadavers and pig carcasses.
Key words.— darkling beetles, Alleculinae, Diaperinae, Tenebrioninae, Pimeliinae,

forensic, cadavers, arid lands.
INTRODUCTION carcasses, and its forensic importance has been fre-

quently documented (Smith 1986, Battán and Linhares
The order Coleoptera forms one of the major in- 2011, Prado e Castro et al. 2013, Caballero and León-
sect groups involved in the decomposition process of Cortés 2014, Zanetti et al. 2015a, b). Certain beetle
PL ISSN 0003-4541 © Museum and Institute of Zoology PAS

doi: 10.3161/00034541ANZ2016.66.4.021
704 F. H. ABALLAY, G. E. FLORES, V. A. SILVESTRO, N. I. ZANETTI and N. D. CENTENO
species are associated with advanced stages of decom- earlier works (Mégnin 1894, Luederwaldt 1911, Goff et
position (Smith 1986) and their usefulness in estimat- al. 1986) to most recent ones (Aballay et al. 2012, San-
ing the minimum Post-Mortem Interval (PMI) has been tos et al. 2014, Zanetti et al. 2015a, b). However, these
highlighted by several authors (Kulshrestha and Satpa- taxa have been considered accidental on cadavers
thy 2001, Schroeder et al. 2002, Midgley et al. 2010). (Özdemir and Sert 2009, Corrêa et al. 2014). This is
The main families recorded in decomposition experi- probably caused by specific methodology used in most
ments or on human corpses are Dermestidae Latreille, of the studies, which are concluded a few days after the
Cleridae Latreille, Histeridae Gyllenhaal, Staphyli- body reaches the last stage of decomposition (remains),
nidae Latreille, Nitidulidae Latreille, Scarabaeidae but decomposition continues to progress while tenebri-
Latreille, Silphidae Latreille , and Trogidae MacLeay onids would be present on completely dried corpses
(Smith 1986, Ösdermir and Set 2009, Lefebvre and (Mégnin 1894, Smith 1986, Lefebvre and Gaudry 2009).
Gaudry 2009). Besides those coleopteran families usu- Larval development in some Tenebrionidae species
ally used for making estimates of the post-mortem normally takes about a year (Santos et al. 1988), but it
interval, species of other families such as the Tenebrio- can extend to two, with the larval period lasting
nidae Latreille have frequently been reported in foren- throughout the winter (Smith 1986). Mégnin (1894) re-
sic studies (Voss et al. 2011, Aballay et al 2012, Za- corded Tenebrionidae on human corpses up to three
netti et al. 2015a, b). years or more after death, and Lefebvre and Gaudry
The family Tenebrionidae comprises about 2,300 (2009) associated 100 % of records of tenebrionids on
genera and 20,000 species worldwide (Matthews et al. human corpses with the end of the first generation of
2010), with 478 genera and 4,624 species estimated to colonizing species (Diptera: Calliphoridae). Tenebrion-
occur in the Neotropical region (Almeida and Mise idae are related to the final stage of decomposition in
2009). Members of the subfamilies Pimeliinae Latreille, arid and semi-arid environments, and for this reason
Tenebrioninae Latreille and Diaperinae Latreille are they were probably not collected in the first forensic
ground-dwelling scavengers, usually inhabiting open studies conducted in Argentina, where the experiments
and arid or semiarid lands (Matthews et al. 2010). performed by Oliva (2001) and Centeno et al (2002)
Some of them feed on plant material including decaying involved humid environments.
plants, dead wood, lichens, mosses, fungal fruiting bod- Adult Tenebrionidae have been mentioned in foren-
ies and stored products (Almeida and Mise 2009, sic studies on decomposing pig carcasses (Matuszews-
Matthews et al. 2010). Others feed on dead animal ki et al. 2008, Voss et al. 2011, Battán and Linhares
material, for example: Eschatomoxys pholeter Thom- 2011, Aballay et al. 2012, Caballero and León-Cortés
as and Pape, 2007 which was found feeding on dead 2014, Zanetti et al 2015a, b) and on human corpses
free-tailed bat (Pape et al. 2007); Gyriosomus elon- (Lefebvre and Gaudry 2009, Mariani et al. 2014, Intro-
gatus Waterhouse, 1843 on dead lizards (Pizarro- na et al. 2011, Aballay pers. obs.). Tenebrionidae re-
Araya 2010); Vaniosus profana (Kulzer, 1956) on pig corded on carcasses were identified at family (Battán
corpses; and Achanius antofagastensis Flores and and Linhares 2011, Caballero and León-Cortés 2014),
Aballay, 2015 on pig and llama carcasses (Flores and genus (Arnaldos et al. 2005, Santos et al. 2014), and at
Aballay 2015). Alphitobius diaperinus (Panzer, species levels in recent forensic studies conducted in
1796) and A. laevigatus (Fabricius, 1781) are frequent arid and semiarid lands in Argentina (Aballay et al.
inhabitants of bat caves where they scavenge on guano 2008, 2012, Zanetti et al. 2015 a, b). The correct identi-
as well as on sick or dead bats (Peck 2005, Matthews et fication of insects at species level and knowledge of
al. 2010). In poultry houses, Aphitobius diaperinus their life history as well as the duration of each stage of
feeds on feathers, dead, dying or newly born chicks development lead to accurately establishing the PMI
(Matthews et al. 2010). Under experimental condi- (Turchetto and Vanin 2004). In South America, the use-
tions, larvae of the genera Tenebrio Linnaeus, 1758 fulness of Tenebrionidae as PMI indicators has not
and Eleodes Eschscholtz, 1829 left marks on bones of been established because of the absence of taxonomic
chicken, pig, horse and sheep (Holden et al. 2013). keys allowing for their identification, as well as
Adults and larvae of Tenebrionidae were collected because of the scant documentation of the detailed life
from pig carcasses for use in decomposition experi- cycle of species. A key to the main families of South
ments conducted in Argentina’s arid and semiarid American Coleoptera of forensic importance was pub-
lands (Aballay 2012, Zanetti et al. 2015 a, b). Larvae of lished (Almeida and Mise 2009), which includes a list of
unidentified species were collected from human three identified and four unidentified species of Tene-
corpses in advanced stages of decomposition (Lefebvre brionidae, but which does not include a key for identi-
and Gaudry 2009, Introna et al. 2011, Mariani et al. fying these species.
2014). The objectives of this paper are to provide an illus-
Tenebrionid beetles have been recorded on carcass- trated key to the Tenebrionidae species associated
es by a wide range of forensic entomology studies, from with decaying carcasses in Argentina and to present
AN ILLUSTRATED KEY TO SPECIES OF TENEBRIONIDAE IN ARGENTINA 705
diagnoses, habitus photographs, substratum and dis- bottle trap” made from one 2.5-liter clear plastic soft
tribution for these species. drink bottle 11 cm in diameter, with its top cut off and
turned over, forming a funnel connected to two cham-
bers, an upper collecting chamber and a lower bait
MATERIAL AND METHODS chamber. The lower one is baited with rotting chicken
flesh and works as an olfactory attractant. The upper
Most of the specimens (1421 out of 1509) were col- chamber of the funnel-shaped bottle ends in a 0.5-liter
lected during forensic studies of decomposing pig car- plastic container with 50% propylene glycol as preser-
casses, because this species is the preferred animal vative and captures the insects. This trap was active in
model for forensic entomological studies (Goff 1993). the field for 25 days and protected by rocks to avoid
Decomposition experiments were conducted in the vertebrate scavengers. Collecting sites in Argentina
Argentinean provinces of Mendoza, San Juan, Cata- belong to the biogeographic provinces of Monte (Men-
marca and Buenos Aires. doza and San Juan), Puna (Catamarca) and Espinal
Most of the darkling beetles specimens studied (Buenos Aires) (Cabrera and Willink 1980).
were collected during the entire decomposition process Voucher specimens are housed in the entomological
from carcasses of 28 pigs and one llama. In Mendoza, collection of the Instituto Argentino de Investigaciones
the study was performed on the campus of the Centro de las Zonas Áridas (Mendoza, Argentina) and Univer-
Científico and Tecnológico CCT CONICET-Mendoza sidad Nacional del Sur, Bahía Blanca (Argentina). Dis-
(32°53’ 53.3”S, 68°52’26.2”W, 850 m altitude). Speci- tributional data were obtained from the following
mens were gathered from 12 pig carcasses during the collections: Instituto Argentino de Investigaciones de
four seasons of the year. In San Juan, Tenebrionidae las Zonas Áridas (IADIZA, Mendoza Argentina), Museo
were sampled in summer from two decomposing pig Argentino de Ciencias Naturales “Bernardino Rivada-
carcasses, placed on the campus of Facultad de Cien- via” (Buenos Aires, Argentina), and National Museum
cias Exactas Físicas y Naturales, Universidad Nacional of Natural History, Smithsonian Institution (Washing-
de San Juan (31°32’34.1”S, 68°34’38.2”W, 673 m alti- ton DC, USA). We followed the classification proposed
tude). Whereas in Catamarca (Antofagasta de la Sier- by Matthews et al. (2010) for assigning species and
ra, 26°01’32.3”S, 67°20’36.5”W, 3600 m altitude), they genera to tribes and subfamilies.
were sampled in spring from decomposing carcasses of Tenebrionidae were examined using a NiKon SMZ
two pigs and one llama during spring. In Buenos Aires, 745 microscope. Measurements (given in millimeters)
the experiments were carried out on a field of the Uni- were taken with an ocular micrometer. Overall body
versidad Nacional del Sur, Bahía Blanca (38°41’41”S, length was measured from the apex of the labrum to
62°15’10”W, 70 m altitude), and specimens were the apex of the elytra. The terminology used in the key
gathered from 12 decomposing pig carcasses during and diagnoses follows recent papers dealing with Tene-
the four seasons of the year. For collecting and pre- brionidae (Aalbu 2006, Matthews et al. 2010, Silvestro
serving specimens, we followed the methods by Cen- and Flores 2012, Silvestro et al. 2012, Flores and Abal-
teno et al. (2002) and Aballay et al. (2008, 2012). Tene- lay 2015). Digital photographs of the specimens were
brionids were collected by hand on corpses and from taken with a Canon S50 camera adapted to a NiKon
six pitfall traps placed approximately 40 cm from SMZ 745 stereomicroscope. Final images of the speci-
each carcass during the entire process of carcass mens were produced using the image-stacking free-
decomposition. Other six pitfalls were placed 15 m ware CombineZM (Hadley 2014).
from each carcass as controls, but these control Type specimens of the newly described species are
traps caught local epigean tenebrionids. The tenebri- deposited in the following collections:
onids attracted by the carcasses were collected with FMNH – Field Museum of Natural History, Chicago,
the pitfall traps that had been placed close to the USA;
corpses. IADIZA – Instituto Argentino de Investigaciones de las
Additional Tenebrionidae specimens were obtained Zonas Áridas, Mendoza, Argentina;
using another three types of collection procedures: the IFML – Instituto y Fundación Miguel Lillo, San
first was conducted on human cadavers at various Miguel de Tucumán, Argentina;
places in Mendoza province, authorized by Mendoza’s MACN – Museo Argentino de Ciencias Naturales
Medical Forensic Committee of Mendoza; the second Bernardino Rivadavia, Buenos Aires, Argen-
was carried out during field trips to various Argen- tina;
tinean provinces, assessing carcasses of cow (Bos tau- MLPA – Museo de La Plata, La Plata, Argentina;
rus), fox (Lycalopex griseus), horse (Equus cabal- MNHN – Muséum National d’Histoire Naturelle,
lus), whale (Eubalaena australis), lesser rhea (Pte- Paris, France;
rocnemia pennata), all found outdoors; for the third NHMB – Natural History Museum, Basel, Switzer-
procedure we used a trap defined here as “baited land.
Figure 1. Sites where tenebrionid species of forensic importance were sampled in provinces of Argentina (see Table 1 for a list of species sampled
at each site) Provinces: 1 – Chaco, 2 – Catamarca, 3 – San Juan, 4 – Mendoza 5 – San Luis, 6 – Córdoba, 7 – Buenos Aires, 8 – Chubut.
RESULTS –. Eyes emarginate anteriorly by epistomal canthus

almost completely divided, leaving only one omma-
The list of Tenebrionidae of potential forensic tidium in their narrowest part (Fig. 7); body dull;
importance in Argentina comprises 29 species included pronotum with lateral margins concave, narrowed
in four subfamilies (Alleculinae, Diaperinae, Tenebri- toward posterior angles, punctures on disc sepa-
oninae and Pimeliinae Latreille) and twelve tribes rated by one puncture diameter (Fig. 9) . . . . . . . . .
(Alleculini Laporte, Crypticini Brullé, Alphitobiini Reit- . . . . . . . . . . . . Alphitobius laevigatus (Fabricius)
ter, Opatrini Brullé, Scotobiini Solier, Tenebrionini La- 7. Pronotum widest before midpoint (Figs. 10, 11);
treille, Trilobocarini Lacordaire, Edrotini Lacordaire, anterior margin of clypeus armed with a row of
Evaniosomini Lacordaire, Epitragini Blanchard, Eleno- teeth directed downwards, with one large tooth at
phorini Solier, and Praociini Eschscholtz) distributed middle (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
in eight provinces (Fig. 1). In order to make a more –. Pronotum widest at or behind midpoint, or at base;
accurate use of the key, a diagnosis and habitus photo- anterior margin of clypeus lacking row of teeth . . . 9
graphs are provided for each species. 8. Clypeus short, not covering the labrum, anterior
margin nearly at same level as antennal insertions;
eyes protruding outward, upper margin lacking
Key to species of Tenebrionidae associated carina; length of epicanthus 2.5 x the length of eye
with carcasses in Argentina (Figs. 10, 12) . . . . . . . . Vaniosus profana (Kulzer)
–. Clypeus covering the labrum, with anterior margin
1. Tarsal claws pectinate (Fig. 2) . . . . . . . . . . . . . . . . . produced anteriorly to antennal insertions; eyes
. . . . . . . . . . . . . . . . . . . . Lobopoda breyeri Brèthes not protruding, upper margin with a carina well
– . Tarsal claws simple . . . . . . . . . . . . . . . . . . . . . . . . . 2 marked, starting posteriorly to the epicanthus
2. Eyes completely divided into separate lower and and ending in the posterior margin of eye; length
upper parts by epistomal canthus (Figs. 3–5) . . . 3 of epicanthus equal to length of eye (as in Fig. 11;
–. Eyes entire or emarginate anteriorly by epistomal Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
canthus (Figs. 6, 7), not completely divided . . . . . 5 . . . Achanius antofagastensis Flores and Aballay
3. Pronotum widest at base or behind midpoint, pos- 9. Abdomen without exposed intersegmental mem-
terior angles acute, projected backwards; anten- branes apical to visible sternites three and
nomere 11 as long as 10; dorsal surface of body four . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
clothed with two kinds of setae, one stout, dark –. Abdomen with exposed intersegmental membranes
brown, the other finer, golden, dense (Fig. 3) . . . . . apical to visible sternites three and four . . . . . . 17
. . . . . . . . . . Trichoton roigi Ferrer and Moragues 10. Anterior margin of clypeus trilobate, divided by
–. Pronotum widest at or before midpoint, posterior two undulating incisions; head widest at epican-
angles straight to obtuse, not projected backwards; thus, before eyes (Figs. 14, 15) . . . . . . . . . . . . . . . . .
antennomere 11 longer than 10; dorsal surface . . . . . . . . . . . . Salax lacordairei Guérin-Méneville
of body clothed in sparse, fine, golden setae (Figs. –. Anterior margin of clypeus concave, straight or
4, 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 convex; head widest at or behind eyes . . . . . . . . 11
4. Pronotum with anterior margin concave; elytron 11. Eyes protruding outward; mandibles with a dorsal
with nine rows of punctures and intervals with cusp on basal half, shorter on the left than on the
dense punctures, epipleuron reaching the apex right (as in Fig. 12); labrum partially sclerotized,
(Fig. 4) . . . . . . . . . Blapstinus punctulatus Solier dorsal mandibular cusps slightly overlap the la-
–. Pronotum with anterior margin straight; anterior brum (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
half of elytron with surface densely punctate but . . . . . . . . . Hylithus tentyroides Guérin-Méneville
punctures not aligned in rows, epipleuron abruptly –. Eyes not protruding; mandibles lacking dorsal
abbreviated before apex (Fig. 5) . . . . . . . . . . . . . . . . cusp on basal half; labrum exposed, entirely sclero-
. . . . . . . . . . . . . . . . . . . . . . Conibius franzi Kaszab tized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
5. Eyes emarginate anteriorly by epistomal canthus, 12. Humeri not elevated, elytra fused along the midline
not completely divided (Figs. 6, 7) . . . . . . . . . . . . . 6 and abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
–. Eyes entire: rounded, elliptical, or semicircular, –. Humeri elevated, macropterous metathoracic
narrow, never emarginate . . . . . . . . . . . . . . . . . . . 7 wings present, elytra not fused along the midline
6. Eyes emarginate anteriorly by epistomal canthus and abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
leaving 3 to 4 ommatidia in their narrowest part 13. Large size, usually more than 3.0 cm in length; ely-
(Fig. 6); body shiny; pronotum with lateral mar- tral epipleuron with an undulate ridge bearing
gins subparallel toward posterior angles, punc- transverse, small carinae, modified as strigil;
tures on disc separated by two puncture diameters antennomere 3 longer than 4 + 5 combined; subge-
(Fig. 8) . . . . . . . Alphitobius diaperinus (Panzer) nal process contiguous with mentum, maxillary
articulation of cardo partially covered by subgena parallel in posterior half; prosternal process sub-
and submentum (Figs. 17, 18). . . . . . . . . . . . . . . . . . rectangular, forming a straight angle; elytron dark
. . . . . Megelenophorus americanus (Lacordaire) brown, lacking spots (Fig. 28) . . . . . . . . . . . . . . . . . .
–. Size smaller, less than 1.5 cm in length; elytral epi- . . . . . . Gondwanocrypticus platensis Fairmaire
pleuron lacking ridge or small carinae; anten- –. Pronotum widest at midpoint, lateral margins con-
nomere 3 shorter than 4 + 5 combined; subgenal cave; prosternal process absent; elytron light
process remote from mentum, maxillary articula- brown with two spots and apex black (Fig. 29) . . .
tion of cardo visible (Fig. 19) . . . . . . . . . . . . . . . . 14 . . . . . . . . Poecilocrypticus formicophilus Gebien
14. Medium size, usually more than 1.0 cm in length; 20. Body brown to light brown; head, antennomeres
pronotum widest at base; antennae moniliform, 1–7 and legs clothed in dense woolly, yellow setae;
antennomere 11 shorter and narrower than anten- head elongated behind the eyes, widest at epican-
nomere 10; maxillary palps with last segment axe- thus, before eyes; lateral margins of pronotum with
shaped, widest at apex (Fig. 20) . . . . . . . . . . . . . . . . a constriction at base (Figs. 30, 31) . . . . . . . . . . 21
. . . . . . . . . . . . . . . . . . . . . . Praocis ecostata Kulzer –. Body black or black to reddish; head, anten-
–. Size smaller, less than 0.5 cm in length; pronotum nomeres 1–7 and legs clothed in sparse short, finer
widest behind midpoint; antennae clavate, anten- setae; head not elongated behind the eyes, widest
nomere 11 as long as 10; maxillary palps with last at eyes; lateral margins of pronotum lacking con-
segment subcylindrical, widest at midpoint (Fig. striction at base (Figs. 32–38) . . . . . . . . . . . . . . 22
21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. Head without horn or protuberances; pronotum
Eutelocera cadaverina Flores and Zanetti sp. nov. with two elevated, sharp, central protuberances;
15. Pronotum widest at base, anterior angles rounded elytra with four complete carinae (Fig. 30) . . . . . . .
sloping towards venter, posterior angles acute; ely- . . . . . . . . . . Leptynoderes strangulata Fairmaire
tra with punctures aligned in ten rows (Fig. 22) . . . –. Head with a horn in the clypeus and two protuber-
. . . . . . . . . . . . . . . Hemasodes minutus (Steinheil) ances between the eyes; pronotum with two large,
–. Pronotum widest at or behind midpoint, anterior elongate, central protuberances and four small
angles acute, directed forwards, posterior angles protuberances, two forward and two at the sides of
obtuse (Figs. 23, 24); punctures on elytra dense but the centrals; elytra with nine complete carinae
not aligned in rows (Figs. 25, 26) . . . . . . . . . . . . 16 (Fig. 31) . . . . Leptynoderes nordenskioldi Kulzer
16. Pronotum with lateral margins concave; anten- 22. Anterior margin of profemur with a tooth or a
nomere 11 as long as 10; frons with two tufts of sharp spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
setae inward to eyes (Figs. 23, 25) . . . . . . . . . . . . . . –. Anterior margin of profemur wavy, lacking tooth or
. . . . . . . . . . . Omopheres scabripennis (Steinheil) spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
–. Pronotum with lateral margins subparallel; anten- 23. Anterior margin of profemur with a sharp spine;
nomere 11 longer than 10; frons lacking tufts of pronotum without grooves or foveae, with dense
setae (Figs. 24, 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . punctures regularly distributed, lateral margin not
. . . . . . . . . . . . . . . Omopheres difficilis (Steinheil) curved upwards; elytron with nine longitudinal
17. Antennomeres 8–10 longer than wide; eyes ellipti- rows of conical tubercles, each interval with a row
cal; last tarsomeres lacking longitudinal groove on of sparse tubercles (Fig. 32) . . . . . . . . . . . . . . . . . . .
anterior margin; macropterous metathoracic wings . . . . . . . . . . . . Emmallodera perlifera Burmeister
present, elytra not fused along the midline and –. Anterior margin of profemur with a rounded tooth;
abdomen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 pronotum with a central longitudinal groove, a
–. Antennomeres 8–10 wider than long; eyes narrow, fovea on each side, and a posterior transverse
semicircular; last tarsomeres with longitudinal groove, disc almost smooth, lateral quarters with
groove on anterior margin; elytra fused along the large and dense punctures, lateral margin curved
midline and abdomen . . . . . . . . . . . . . . . . . . . . . . 20 upwards; elytron with nine longitudinal rows of
18. Size large, usually more than 2.5 cm in length; last conical tubercles, each interval with a row of
visible abdominal sternite with groove along outer sparse punctures (Fig. 33) . . . . . . . . . . . . . . . . . . . .
margin; apical ring on antennomeres 7–11 with . . . . . . . . . Scotobius muricatus Guérin-Méneville
simple sensilla (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . 24. Small size: length: 10–11 mm; pronotum with small,
. . . . . . . . . . . . . . . . . Zophobas opacus (Sahlberg) sparse punctures, width 1.8 times the length; elytra
–. Size smaller, less than 0.5 cm in length; last visi- almost subquadrate, 1.20 times longer than wide,
ble abdominal sternite lacking groove along outer lateral margins nearly parallel (Fig. 34) . . . . . . . . .
margin; apical ring on antennomeres 7–11 with . . . . . . . . . . . . . Scotobius ovalis Guérin-Méneville
compound, stellate sensoria, visible as white cir- –. Medium to large size: length: 19–30 mm; pronotum
cles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 with large, dense punctures; width not exceeding
19. Pronotum widest at base, lateral margins arcuate, 1.5 times the length; elytra elongate, 1.36 times
longer than wide, lateral margins oval (Figs. groove, clypeus lower than frons. Maxillary palps axe-
35–38) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 shaped, distal margin four times width than base. Eyes
25. Elytra with rows of punctures and intervals form- emarginate anteriorly by epistomal canthus not com-
ing broad carinae slightly raised, turning back- pletely divided. Antennae filiformis, antennomere 11 as
wards into rows of tubercles and converging pos- long as 10. Pronotum widest at base or at midpoint,
tero-laterally (Fig. 35, 36) . . . . . . . . . . . . . . . . . . . 26 with lateral margins subparallel in posterior half; disc
–. Elytra without rows of punctures, with rows of flat, lateral quarters sloping towards venter, posterior
tubercles all along (Figs. 37–39) . . . . . . . . . . . . . 27 angles straight, posterior margin bisinuate, wider than
26. Pronotum widest behind midpoint, with punctures anterior margin. Elytron with nine longitudinal rows of
uniformly distributed, anterior angles acute and sunken punctures and intervals slightly raised with
produced, posterior angles straight; elytron with small disperse punctures; humeri elevated, macropter-
punctures of same size or smaller than pronotum ous metathoracic wings present, elytra not fused along
punctures (Fig. 35) . . . . . . . . . . . . . . . . . . . . . . . . . . . the midline and abdomen. Abdomen with exposed
. . . . . . . . . . . . . Scotobius punctatus Eschscholtz intersegmental membranes apical to visible sternites
–. Pronotum widest at midpoint, with punctures not three and four. Tarsal claws pectinate (Fig. 2).
uniformly distributed, denser on margins, anterior Distribution. Argentina: provinces Buenos Aires,
angles straight to obtuse, not produced, posterior Catamarca, Chubut, Córdoba, La Rioja, Mendoza, Neu-
angles obtuse; elytron with punctures larger than quén, Rio Negro, San Juan, San Luis, Tucumán (collec-
pronotum punctures (Fig. 36) . . . . . . . . . . . . . . . . . . tion data IADIZA).
. . . . . . . . . . . . . . . . . . Scotobius andrassyi Kaszab
27. Pronotum widest at midpoint, with 2 to 4 irregular,
smooth and slightly raised areas; anterior margin Trichoton roigi Ferrer and Moragues, 2001
as wide as posterior margin; elytra with rows of (Tenebrioninae, Opatrini)
large round tubercles, with only 10–16 tubercles (Fig. 3)
per row (Fig. 37) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . Scotobius pilularius Germar Diagnosis. Medium size (length: 7.2–8.1 mm, width:
–. Pronotum widest behind midpoint, without smooth 3.5–4.1 mm). Body dark brown, oval, dorsal surface
raised areas; anterior margin different width than clothed with two kinds of setae, one stout, dark brown,
posterior margin; elytra with rows of medium size the other finer, golden, dense. Head and pronotum with
tubercles, more than 25 tubercles per row (Figs. 38, punctures separated by less than one puncture diame-
39) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 ter, frons punctures same size of clypeal punctures.
28. Large size: length 26–30 mm; pronotum with ante- Eyes completely divided into separate lower and upper
rior margin wider than posterior margin, anterior parts by epistomal canthus. Antennae moniliform,
angles produced; elytra with rows of conical tuber- antennomere 11 as long as 10. Pronotum widest at base
cles aligned (Fig. 38) . . . . . . . . . . . . . . . . . . . . . . . . . or behind midpoint, disc very raised, lateral quarters
. . . . . . . . . . . . . . . . . Scotobius casicus Lacordaire flat, posterior angles acute, projected backwards, pos-
–. Medium size: length 15–17 mm; pronotum with terior margin bisinuate, wider than anterior margin.
anterior margin narrower than posterior margin, Elytron with nine longitudinal rows of big punctures
anterior angles not produced; elytra with rows of and intervals carina-shape slightly raised, with small
rounded tubercles not aligned (Fig. 39) . . . . . . . . . . punctures; humeri elevated, macropterous metatho-
. . . . . . . . . . . . . . . . . . . . Scotobius granosus Solier racic wings present, elytra not fused along the midline
and abdomen. Abdomen with exposed intersegmental
membranes apical to visible sternites three and four.
DIAGNOSES OF SPECIES Distribution. Argentina: provinces Buenos Aires
(Zanetti et al. 2015 a, b), Córdoba, Formosa, La Rioja,
Lobopoda breyeri Brèthes, 1910 Mendoza, Salta, San Juan, Santa Fe (collection data
(Alleculinae, Alleculini) IADIZA).
(Fig. 2)
Diagnosis. Medium size (length: 11.8–17.6 mm, Blapstinus punctulatus Solier, 1851
width: 3.8–5.9 mm). Body dark brown, elongate, dorsal (Tenebrioninae, Opatrini)
surface clothed in dense, short, fine, brown setae. (Fig. 4)
Pronotum with punctures separated by more than two
puncture diameters, frons punctures same size of Diagnosis. Small size (length: 4.9–6.2 mm, width:
clypeal punctures, separated by less than one punc- 2.1–2.5 mm). Body black to dark brown, oval, dorsal
ture diameter. Clypeal suture as a vertical, bisinuate surface clothed in sparse, short, fine, golden setae.
Head and pronotum with punctures separated by one Alphitobius diaperinus (Panzer, 1796)
puncture diameter, frons punctures same size of (Tenebrioninae, Alphitobiini)
clypeal punctures. Eyes completely divided into sepa- (Figs 4, 6)
rate lower and upper parts by epistomal canthus.
Antennae moniliform, antennomere 11 longer than 10. Diagnosis. Small size (length: 5.9–7.4 mm, width:
Pronotum widest at or before midpoint, disc and later- 2.5–3.3 mm). Body shiny, dark brown, oval, dorsal sur-
al quarters raised, posterior angles straight, not pro- face clothed in sparse, very short, fine setae on punc-
jected backwards, posterior margin almost straight, tures. Frons punctures same size of clypeal punctures.
wider than anterior margin. Elytron with nine longitu- Eyes emarginate anteriorly by epistomal canthus not
dinal rows of sunken punctures and intervals with dis- completely divided, leaving 3 to 4 ommatidia in their
perse punctures; humeri weakly elevated, macropter- narrowest part (Fig. 6), head widest at epicanthus,
ous metathoracic wings present, elytra not fused along before eyes. Antennae capitate, antennomeres 6 to 10
the midline and abdomen. Abdomen with exposed wider than long, asymmetrically expanded apically
intersegmental membranes apical to visible sternites (Fig. 6), antennomere 11 longer than 10. Pronotum
three and four. widest at or before midpoint, with lateral margins sub-
Distribution. Argentina: provinces Buenos Aires parallel toward posterior angles, disc and lateral quar-
(Zanetti et al. 2015 a, b), Catamarca, Chaco, Chubut, ters raised, posterior angles straight, not projected
Córdoba, Entre Ríos, Jujuy, La Pampa, La Rioja, Men- backwards, posterior margin bisinuate, wider than
doza, Neuquén, Salta, San Juan, San Luis, Santa Fe anterior margin; punctures separated by less than two
Santiago del Estero, Río Negro, Tucumán, (collection puncture diameters on lateral quarters and by two
data IADIZA, MACN); Bolivia: Cochabama department puncture diameters on disc. Elytron with eight longitu-
(collection data IADIZA) new record for Bolivia; Chi- dinal rows of sunken punctures and intervals with
le: provinces Bio-Bio, Cachapoal, Chiapa, Cordillera, shallow punctures; humeri weakly elevated, macro-
Elqui, Linares, Maipo, Malleco Ñuble, Quillota, Santia- pterous metathoracic wings present, elytra not fused
go, Talca, (Vidal and Guerrero 2007). Uruguay: depart- along the midline and abdomen. Abdomen with ex-
ments Canelones (Atlántida), Colonia, Montevideo posed intersegmental membranes apical to visible ster-
(Carrasco) (collection data MACN) new record for nites three and four.
Uruguay. Distribution. Argentina: provinces Buenos Aires,
Entre Ríos, La Rioja, Mendoza, Río Negro, San Juan
(collection data IADIZA).
Conibius (Gondwanodilamus) franzi Kaszab, 1969 Remarks. It is currently cosmopolitan, introduced
(Tenebrioninae, Opatrini) in South America and recently recorded in Argentina
(Fig. 5) (Cecco et al. 2005), it has been associated with many
stored products (Spilman 1991) and in poultry farms
Diagnosis. Small size (length: 3.9–4.6 mm, width: (Cecco et al. 2005, Matthews et al. 2010).
1.2–1.8 mm). Body black to dark brown, elongate, dor-
sal surface clothed in sparse, very short, fine setae
on punctures. Head, pronotum and elytron with punc- Alphitobius laevigatus (Fabricius, 1781)
tures separated by one puncture diameter, frons (Tenebrioninae, Alphitobiini)
punctures same size of clypeal punctures. Eyes com- (Figs 5, 7)
pletely divided into separate lower and upper parts by
epistomal canthus. Antennae moniliform, antennomere Diagnosis. Small size (length: 5.6–6.3 mm, width:
11 longer than 10. Pronotum widest before midpoint, 2.3–2.5 mm). Body dull, dark brown, oval, dorsal sur-
disc and lateral quarters raised, posterior angles face clothed in sparse, very short, fine setae on punc-
obtuse, not projected backwards, posterior margin tures. Frons punctures same size of clypeal punctures.
concave, narrower than anterior margin. Elytron Eyes emarginate anteriorly by epistomal canthus
densely punctate, on anterior half punctures not almost completely divided, leaving only one ommatidi-
aligned in rows; epipleuron abruptly abbreviated um in their narrowest part (Fig. 7), head widest at epi-
before apex, humeri not elevated, metathoracic wings canthus, before eyes. Antennae capitate, anten-
present, reduced. Abdomen with exposed interseg- nomeres 6 to 10 wider than long, asymmetrically
mental membranes apical to visible sternites three and expanded apically (Fig. 7), antennomere 11 longer than
four. 10. Pronotum widest at midpoint, with lateral margins
Distribution. Argentina: provinces La Pampa, Men- concave, narrowed toward posterior angles, disc and
doza, Neuquén, Salta, San Juan (collection data lateral quarters raised, posterior angles obtuse, not
IADIZA), new record for Argentina. Chile: provinces projected backwards, posterior margin bisinuate,
Chañaral, El Loa (Vidal and Guerrero 2007). wider than anterior margin; punctures uniformely
distributed, separated by one puncture diameter. Ely- punctures separated by two puncture diameters, frons
tron with eight longitudinal rows of sunken punctures punctures same size of clypeal punctures. Right
and intervals with shallow punctures; humeri weakly mandible with a dorsal cusp on the basal half, left
elevated, macropterous metathoracic wings present, mandible with a dorsal sharp raised edge; anterior
elytra not fused along the midline and abdomen. margin of clypeus armed with a row of teeth directed
Abdomen with exposed intersegmental membranes downwards, with one large tooth at middle. Eyes
apical to visible sternites three and four. round, upper margin with a carina well marked (Fig.
Distribution. Argentina: Mendoza province (collec- 13). Antennae moniliform, antennomeres 6 to 10 trian-
tion data IADIZA). gular, expanded outwards, antennomere 11 longer
Remarks. It is currently cosmopolitan and intro- than 10. Pronotum widest before midpoint, disc and lat-
duced in South America, it has been associated with eral quarters raised, with anterior angles rounded,
many stored products (Spilman 1991), new record for sloping towards venter, posterior angles obtuse, not
Argentina. projected backwards, posterior margin concave, nar-
rower than anterior margin. Elytron with ten longitudi-
nal rows of punctures; humeri elevated, metathoracic
Vaniosus profana (Kulzer, 1956) wings present, reduced, narrow, reaching the ¾ of
(Pimeliinae, Evaniosomini) length of elytra. Abdomen without exposed interseg-
(Figs 10, 12) mental membranes apical to visible sternites three and
four.
Diagnosis. Small size (length: 3.6–4.8 mm, width: Distribution. Argentina: Catamarca province over
1.3–1.8 mm). Body dark brown, elongate, dorsal sur- altitude of 3000 m (Flores and Aballay 2015).
face clothed in sparse, very short, fine setae on punc-
tures. Head and pronotum with punctures separated by
less than one puncture diameter, frons punctures same Salax lacordairei Guérin-Méneville, 1834
size of clypeal punctures. Mandibles with a dorsal cusp (Pimeliinae, Trilobocarini)
on basal half, longer on the right than on the left (Figs 14, 15)
mandible (Fig. 12); anterior margin of clypeus armed
with a row of teeth directed downwards, with one large Diagnosis. Medium size (length: 7.9–9.9 mm, width:
tooth at middle (Fig. 12). Eyes protruding outward. 3.1–4.6 mm). Body black to dark brown, elongate, dor-
Antennae moniliform, antennomere 11 as long as 10. sal surface clothed in dense, short, fine setae. Head,
Pronotum widest before midpoint, disc raised, lateral pronotum and elytron with punctures separated by one
quarters flat, posterior angles obtuse, not projected puncture diameter; frons punctures larger than clypeal
backwards, posterior margin concave, narrower than punctures. Anterior margin of clypeus trilobate, divid-
anterior margin. Elytron with ten longitudinal rows of ed by two undulating incisions. Eyes round, upper mar-
punctures, only the anterior quarter of intervals with gin of eyes with a carina well marked; head widest at
disperse punctures; humeri weakly elevated, metatho- epicanthus, before eyes (Fig. 14). Antennae monili-
racic wings present, reduced, as long as elytra, not form, antennomere 11 shorter than 10; antennomere 10
folded. Abdomen without exposed intersegmental twice the width of antennomere 11 (Fig. 14). Pronotum
membranes apical to visible sternites three and four. widest at or before midpoint, disc and lateral quarters
Distribution. Argentina: provinces Catamarca, raised, posterior angles straight, not projected back-
Córdoba, La Rioja, Mendoza, Salta, San Juan, Santiago wards, posterior margin bisinuate, wider than anterior
del Estero, Tucumán (Flores and Aballay 2015), and margin. Elytron with punctures not aligned in rows,
Formosa; Bolivia: Santa Cruz department (collection with five to seven superficial, longitudinal grooves and
data USNM), new record for Bolivia. intervals carina-shape slightly raised; humeri elevated,
macropterous metathoracic wings present, elytra not
fused along the midline and abdomen. Abdomen with-
Achanius antofagastensis Flores and Aballay, 2015 out exposed intersegmental membranes apical to visi-
(Pimeliinae, Evaniosomini) ble sternites three and four.
(Fig. 11) Distribution. Argentina: provinces Catamarca,
Córdoba, La Pampa, La Rioja, Mendoza, Neuquén,
Diagnosis. Small size (length: 3.5–4.9 mm, width: Río Negro Salta, San Juan, San Luis, Santiago del
1.1–1.5 mm). Body elongate, head and prothorax dark Estero, Tucumán (collection data IADIZA); Bolivia:
brown, elytra light brown, with a dark brown spot on Cochabama department (collection data IADIZA) new
central third extending backwards along the suture; record for Bolivia; Chile: provinces Antofagasta, El
antennae, legs and mouth parts light brown as elytra, Loa (collection data IADIZA; Vidal and Guerrero,
dorsal surface glabrous. Head and pronotum with 2007).
Hylithus tentyroides Guérin-Méneville, 1834 bearing transverse, small carinae, modified as strigil.
(Pimeliinae, Edrotini) Abdomen without exposed intersegmental membranes
(Fig. 16) apical to visible sternites three and four.
Distribution. Argentina: provinces Catamarca,
Diagnosis. Small size (length: 5.4–7.9 mm, width: Formosa, La Rioja, Mendoza, Neuquén, Río Negro,
2.1–3.1 mm). Body black to dark brown, elongate, dor- Salta, San Juan, Santiago del Estero, Tucumán (collec-
sal surface clothed in sparse, very short setae on punc- tion data IADIZA and MACN).
tures. Head and pronotum with punctures separated by
less than two puncture diameters, frons punctures
same size of clypeal punctures. Mandibles with a dor- Praocis ecostata Kulzer, 1958
sal cusp on basal half, shorter on the left than on the (Pimeliinae, Praociini)
right (as in Fig. 12); labrum partially sclerotized, dor- (Figs 19, 20)
sal mandibular cusps slightly overlap the labrum. Eyes
round, protruding outward. Antennae moniliform, Diagnosis. Medium size (length: 9.6–12.1 mm,
antennomere 11 longer than 10. Pronotum widest at width: 6.1–8.1 mm). Body black to dark brown, oval,
midpoint, disc and lateral quarters not raised, posteri- dorsal surface of pronotum and elytron clothed in
or angles obtuse, not projected backwards, posterior sparse, very short setae on punctures; with noticeable
margin as wide as anterior margin. Elytron with nine punctures on pronotum and elytra not uniformly dis-
longitudinal rows of punctures smaller than head and tributed, separated by more than two puncture diame-
pronotum punctures, intervals with punctures same ters, bigger on elytra than on pronotum. Clypeal suture
size aligned in rows, making visible the surface with as horizontal groove not covered by frons, clypeus low-
eighteen rows of punctures; humeri elevated, macro- er than frons. Eyes elliptical. Antennae moniliform,
pterous metathoracic wings present, elytra not fused antennomere 11 shorter and narrower than antenno-
along the midline and abdomen. Abdomen without mere 10. Pronotum widest at base, lateral margins
exposed intersegmental membranes apical to visible arcuate, well marked; disc very raised, lateral quarters
sternites three and four. raised, posterior angles acute, projected backwards;
Distribution. Argentina: provinces Buenos Aires, posterior margin bisinuate, wider than anterior mar-
Catamarca, Chubut, Córdoba, La Rioja, Mendoza, Rio gin. Elytron with two longitudinal carinae slightly
Negro, Salta, San Juan, Tucumán (collection data IADI- raised, equidistant between suture and lateral margin;
ZA and MACN; Zanetti et al. 2015 a, b). humeri not elevated, elytra fused along the midline and
abdomen. Abdomen without exposed intersegmental
Megelenophorus americanus (Lacordaire, 1830) Distribution. Argentina: provinces Catamarca,
(Pimeliinae, Elenophorini) Jujuy and Salta, over altitude of 3000 m (collection data
(Figs 17, 18) IADIZA).
Diagnosis. Large size (length: 29.9–40.7 mm, width:

11.4–15.9 mm). Body black, elongate, dorsal surface Eutelocera cadaverina Flores and Zanetti sp. nov.
clothed in sparse, very short setae arising on small pro- (Pimeliinae, Praociini)
tuberances in frons, pronotum and elytra. Clypeus with (Fig. 21)
sparse punctures on anterior half, frontoclypeal and
genoclypeal sutures well marked and deep; subgenal Diagnosis. Eutelocera cadaverina sp. nov. dif-
process contiguous with mentum, maxillary articula- fers from E. viatica Solier 1840 by having pronotum
tion of cardo partially covered by subgena and submen- with posterior angles obtuse, rounded, not produced
tum (Fig. 18). Eyes semicircular, narrow. Antennae fil- backwards, posterior margin convex, elytron with
iformis, antennomere 3 longer than 4 + 5 combined; small, isolated protuberances and dorsal surface
antennomere 10 and 11 shorter than remaining. Prono- clothed with two kinds of setae, one stout, the other fin-
tum separated from elytra, lateral margins concave, er, while in E. viatica the pronotum has posterior
well marked, carina shape, widest at midpoint, disc angles straight produced backwards, posterior margin
very raised, lateral quarters raised, posterior angles biconcave, elytron with three rows of big protuber-
obtuse, not projected backwards, posterior margin ances on posterior half, lateral margin formed by big
straight, narrower than anterior margin. Elytron plain, protuberances, and dorsal surface clothed in sparse,
with lateral margins carina-shape, humeri not elevat- finer setae.
ed, elytra fused along the midline and abdomen; Description. Small size (length: 3.8–4.9 mm, width:
pseudopleuron with two carinae; elytral epipleuron 1.8–2.5 mm). Body black to dark brown, oval, dorsal
broadened in posterior half, with an undulate ridge surface clothed with two kinds of setae, one stout, the
other dense, finer. Clypeal suture as horizontal groove metathoracic wings present, elytra not fused along the
not covered by frons, clypeus lower than frons. Eyes midline and abdomen. Abdomen without exposed inter-
elliptical. Antennae clavate, antennomere 11 as long as segmental membranes apical to visible sternites three
10. Pronotum with small protuberances, widest behind and four.
midpoint, lateral margins concave, well marked; disc Distribution. Argentina: provinces Buenos Aires,
raised, lateral quarters flat, posterior angles obtuse, La Rioja, Mendoza, San Juan, San Luis, Santiago del
not produced backwards; posterior margin convex, Estero (collection data IADIZA and MACN).
wider than anterior margin. Elytron with deep, sparse
punctures and small, isolated protuberances on distal
third and on pseudopleuron; humeri not elevated, ely- Omopheres scabripennis (Steinheil, 1874)
tra fused along the midline and abdomen. Abdomen (Pimeliinae, Epitragini)
without exposed intersegmental membranes apical to (Figs 23, 25)
visible sternites three and four.
Etymology. Its name is derived from the Latin word Diagnosis. Medium size (length: 10.8–15.5 mm,
cadaverina, prope of carcase, meaning that feed on width: 3.9–7.1 mm). Body brown to dark brown, elon-
dead bodies. gate, dorsal surface of pronotum and elytra clothed in
Type material. Holotype, male: [Argentina, Buenos sparse, golden, very short setae; punctures on prono-
Aires, Dto. Bahia Blanca, Bahia Blanca, Altos del Pali- tum and elytra separated by less than two puncture
hue, on pig carcase, 38°41’41’’S, 62°15’10’’W, 70 m, diameters. Frons with two tufts of setae inward to eyes.
17.II.2011, coll: N. Zanetti] [Eutelocera / cadaverina Eyes round. Antennae filiformis, antennomeres 8 to 10
sp. n. / HOLOTYPUS male / Det. G. Flores and / N. asymmetrically expanded outwards, antennomere 11
Zanetti 2016] (IADIZA). Allotype, female: with the as long as 10. Pronotum widest at or behind midpoint,
same data as holotype except date: 9.IX.2010 (IADIZA). anterior angles acute, directed forwards, lateral mar-
Paratypes with the same data as holotype except date: gins concave, well marked; disc and lateral quarters
one female 28.VIII.2010 (IADIZA); two males and one raised, posterior angles obtuse, not projected back-
female 10.XI.2010 (IADIZA); one male 10.XII.2010 wards; posterior margin bisinuate, wider than anterior
(MLPA); three males 1.III.2011 (2 MACN, 1 MLPA); four margin. Elytron with noticeable, dense punctures not
males 1.IV.2011 (2 FMNH, 2 NHMB); one male aligned in rows; humeri elevated, macropterous
12.IV.2011 (MNHN); one male 17.II.2011 (IADIZA); one metathoracic wings present, elytra not fused along the
male 18.X.2011 (IFML); one paratype not sexed midline and abdomen. Abdomen without exposed inter-
8.X.2010 (IFML). segmental membranes apical to visible sternites three
Distribution. Argentina: Known only from the typi- and four.
cal locality, Altos del Palihue, Bahia Blanca, Buenos Distribution. Argentina: provinces Catamarca,
Aires province. Chaco, Córdoba, La Rioja, Mendoza, Neuquén, Rio
Remarks. Prior to this study Eutelocera Solier, Negro, San Juan, San Luis, Santiago del Estero (collec-
1840 was monotypic. Now the genus is comprised of tion data IADIZA and MACN); Bolivia (Freude 1967).
two species: E. viatica and E. cadaverina sp. nov.
Omopheres difficilis (Steinheil, 1874)

Hemasodes minutus (Steinheil, 1874) (Pimeliinae, Epitragini)
(Pimeliinae, Epitragini) (Figs 24, 26)
(Fig. 22)
Diagnosis. Medium size (length: 10.1–15.1 mm,
Diagnosis. Small size (length: 5.1–8.2 mm, width: width: 4.1–6.6 mm). Body brown to dark brown, elon-
2.0–3.4 mm). Body brown, elongate, dorsal surface of gate, dorsal surface of pronotum and elytron clothed in
pronotum and elytron clothed in sparse, golden, very sparse, golden, very short setae; punctures on prono-
short setae; with punctures on pronotum and elytra tum and elytra separated by two puncture diameters.
separated by less than two puncture diameters. Eyes Eyes round. Antennae filiformis, antennomeres 8 to 10
round. Antennae filiformis, antennomere 11 as long as asymmetrically expanded outwards, antennomere 11
10. Pronotum widest at base, anterior angles rounded longer than 10. Pronotum widest at or behind midpoint,
sloping towards venter, lateral margins arcuate, well anterior angles acute, directed forwards, lateral mar-
marked; disc raised, lateral quarters not raised, poste- gins subparallel, well marked; disc and lateral quarters
rior angles acute, projected backwards; posterior raised, posterior angles obtuse, not projected back-
margin bisinuate, wider than anterior margin. Elytron wards; posterior margin convex, wider than anterior
with punctures aligned in 10 rows, intervals with margin. Elytron with dense punctures not aligned in
sparse punctures; humeri elevated, macropterous rows; humeri elevated, macropterous metathoracic
wings present, elytra not fused along the midline and longitudinal rows of punctures and intervals with
abdomen. Abdomen without exposed intersegmental sparse punctures; humeri weakly elevated, macropter-
membranes apical to visible sternites three and four. ous metathoracic wings present, elytra not fused along
Distribution. Argentina: provinces Catamarca, the midline and abdomen. Abdomen with exposed
Chaco, La Rioja, Mendoza, Neuquén, Rio Negro, Salta, intersegmental membranes apical to visible sternites
San Juan, San Luis, Santiago del Estero (collection three and four.
data IADIZA and MACN); Bolivia and Paraguay (Freu- Distribution. Argentina: provinces Buenos Aires
de 1967). (Zanetti et al. 2015 a, b), Catamarca, Córdoba, For-
mosa, La Pampa, La Rioja, Mendoza, Salta, San Juan,
San Luis, Santiago del Estero, Tucumán (collection
Zophobas opacus (Sahlberg, 1823) data IADIZA and MACN) and Misiones; Uruguay: Mon-
(Tenebrioninae, Tenebrionini) tevideo department (Fairmaire 1883). Introduced in the
(Fig. 27) United States (Steiner and Swearingen 2014).
Diagnosis. Large size (length: 22.9–29.3 mm, width:

7.5–9.4 mm). Body black, elongate, surface of pronotum Poecilocrypticus formicophilus Gebien, 1928
and elytron glabrous. Eyes elliptical. Antennae monili- (Diaperinae, Crypticini)
form, antennomere 11 with apex asymmetrically (Fig.
curved outwards, longer than 10. Pronotum almost
smooth with sparse punctures, widest at or before mid- Diagnosis. Small size (length: 2.8–3.2 mm, width:
point, anterior angles rounded, lateral margins con- 1.1–1.3 mm). Body oval, light brown to reddish, elytron
cave, well marked; disc and lateral quarters raised, light brown with two spots and apex black, dorsal sur-
posterior angles obtuse, not projected backwards; pos- face glabrous. Anterior margin of clypeus straight.
terior margin slightly bisinuate, wider than anterior Eyes elliptical. Antennae moniliform, antennomere 11
margin. Elytron with punctures aligned in nine rows, with apex asymmetrically curved outwards, longer
intervals smooth; humeri elevated, macropterous than antennomere10; apical ring on antennomeres
metathoracic wings present, elytra not fused along the 7–11 with compound, stellate sensoria, visible as white
midline and abdomen. Abdomen with exposed interseg- circles. Pronotum with punctures separated by one
mental membranes apical to visible sternites three and puncture diameter, widest at midpoint, disc and lateral
four; last visible abdominal sternite with groove along quarters raised, lateral margins concave, posterior
outer margin. angles straight, not projected backwards, posterior
Distribution. America, from United States to margin straight, wider than anterior margin. Elytron
Argentina (Ferrer 2011). In Argentina: provinces Cha- with nine longitudinal rows of punctures and intervals
co, Córdoba, Formosa, Mendoza, Salta, San Juan, San- with sparse, smaller punctures; humeri weakly elevat-
ta Fe, Santiago del Estero (collection data IADIZA). ed, macropterous metathoracic wings present, elytra
not fused along the midline and abdomen. Abdomen
with exposed intersegmental membranes apical to vis-
Gondwanocrypticus platensis Fairmaire, 1883 ible sternites three and four.
(Diaperinae, Crypticini) Distribution. Argentina: provinces Buenos Aires,
(Fig. 28) Córdoba, Mendoza (collection data IADIZA and MACN;
Gebien 1928) and Brazil (Rio Grande do Sul; Gebien
Diagnosis. Small size (length: 3.6–4.9 mm, width: 1928). Introduced in the United States (Steiner 1982).
1.8–2.5 mm). Body dark brown to black, oval, dorsal
surface clothed in sparse, very short, fine setae.
Clypeal suture not marked, anterior margin of clypeus Leptynoderes strangulata Fairmaire, 1903
contiguous with genae, forming an arcuate anterior (Tenebrioninae, Scotobiini)
margin of head. Eyes elliptical. Antennae filiformis, (Fig. 30)
antennomere 11 with apex asymmetrically curved out-
wards, longer than 10; apical ring on antennomeres Diagnosis. Medium size (length: 9.4–14.4 mm,
7–11 with compound, stellate sensoria, visible as white width: 3.9–6.5 mm). Body brown to light brown, usually
circles. Pronotum with punctures separated by one covered with grains of sand. Head elongated behind the
puncture diameter, widest at base, disc and lateral eyes, without horn or protuberances on clypeus. Head,
quarters raised, lateral margins arcuate, parallel in antennomeres 1–7 and legs covered with dense woolly,
posterior half, posterior angles straight, not projected yellow setae, setae not covering completely the cuticle.
backwards, posterior margin straight, twice the length Eyes narrow, semicircular. Antennae moniliform,
of anterior margin. Elytron with nine imperceptible antennomere 11 slightly longer than 10. Pronotum with
two acute central protuberances; lateral margin frons punctures two times larger than clypeal punc-
expanded upwards with a constriction at base; posteri- tures. Eyes narrow, semicircular. Antennae monili-
or angles straight, produced backwards; posterior mar- form, antennomere 11 slightly longer than 10. Prono-
gin straight, wider than anterior margin. Elytron with tum widest at midpoint, with dense punctures of same
seven carinae: first adjoining elytral suture, 2 and 4 as size of clypeal punctures regularly distributed; lateral
rows of acute tubercles only visible in posterior third; margin not curved upwards; anterior angles acute; pos-
carina 3 entire, raised, acute and smooth; carina 5 terior margin straight, wider than anterior margin.
raised with tubercles; 6 and 7 like rows of acute tuber- Elytron with nine longitudinal rows of conical tuber-
cles inconspicuous for the grains of sand; humeri cles, the first three rows inconspicuous anteriorly,
weakly elevated, elytra fused along the midline and each interval with a row of sparse tubercles, more
abdomen. Abdomen with exposed intersegmental mem- noticeable on the sides; humeri not elevated, elytra
branes apical to visible sternites three and four. fused along the midline and abdomen. Legs with punc-
Distribution. Argentina: Catamarca, Tucumán tures; anterior margin of profemur with a sharp spine.
(Kulzer 1955); Córdoba, La Rioja, Northern Mendoza, Abdomen with exposed intersegmental membranes
San Juan (collection data IADIZA). apical to visible sternites three and four.
Distribution. Argentina provinces Catamarca, La
Rioja, Mendoza, Neuquén, Salta, San Juan, Santiago
Leptynoderes nordenskioldi Kulzer, 1955 del Estero (Kulzer 1955), Córdoba (collection data
(Tenebrioninae, Scotobiini) IADIZA).
(Fig. 31)
Diagnosis. Medium size (length: 11.0–13.0 mm, Scotobius muricatus Guérin-Méneville, 1834
width: 4.9–5.8 mm). Body brown to light brown, dull, (Tenebrioninae, Scotobiini)
sometimes covered with grains of sand. Head elongated (Fig. 33)
behind the eyes with a horn on the clypeus and two
acute and elongate protuberances between the eyes. Diagnosis. Large size (length: 18.2–29.9 mm, width:
Head, antennomeres 1–7 and legs covered with dense 8.2–12.2 mm). Body black, dull, elongated. Head with
woolly, yellow setae, covering completely the cuticle. punctures separated less than two puncture diameters;
Eyes narrow, semicircular. Antennae moniliform, anten- frons punctures three to four times larger than clypeal
nomere 11 slightly longer than 10. Pronotum with two punctures. Eyes narrow, semicircular. Antennae moni-
large, elongate central protuberances and four small liform, antennomere 11 slightly longer than 10. Prono-
protuberances, two forward and two at the sides of the tum widest at midpoint, with a central longitudinal
centrals; lateral margin expanded upwards with a con- groove, a fovea on each side, and a posterior transver-
striction at base; posterior angles acute, produced back- se groove; disc almost smooth, lateral quarters of pro-
wards; posterior margin straight, wider than anterior notum with large and dense punctures of same size of
margin. Elytron with nine carinae: first adjoining elytral clypeal punctures; lateral margin curved upwards; an-
suture, 2 as row of small tubercles slightly raised, more terior angles rounded and downwards; posterior margin
pronounced in posterior third; 3 and 4 more raised than straight, as wide as anterior margin. Elytron with nine
the remaining, acute and smooth, backwards become in longitudinal rows of conical tubercles well defined even
elongate tubercles; the remaining carinae as rows of backwards and sideways; humeri not elevated, elytra
acute tubercles sometimes elongated backwards; fused along the midline and abdomen. Legs with incon-
humeri weakly elevated, elytra fused along the midline spicuous punctures; anterior margin of profemur with
and abdomen. Abdomen with exposed intersegmental a rounded tooth. Abdomen with exposed intersegmental
membranes apical to visible sternites three and four. membranes apical to visible sternites three and four.
Distribution. Argentina: South Buenos Aires Distribution. Argentina provinces Córdoba, La
(Zanetti et al. 2015 a, b), Chubut, Southern Mendoza, Pampa, San Luis (collection data IADIZA) Buenos Aires
Neuquén, Rio Negro (collection data IADIZA). (Zanetti et al. 2015a, b); Uruguay; Paraguay (Kulzer
1955).
Emmallodera perlifera Burmeister, 1875

(Tenebrioninae, Scotobiini) Scotobius ovalis Guérin-Méneville, 1834
(Fig. 32) (Tenebrioninae, Scotobiini)
(Fig. 34)
Diagnosis. Large size (length: 20.9–28.8 mm,
width: 9.1–13.4 mm). Body black, dull, oval. Head with Diagnosis. Medium size (length: 10.3–11.5 mm,
punctures separated less than two puncture diameters; width: 4.6–6.0 mm). Body black, dull, oval and glabrous.
Head punctures separated by more than three puncture glabrous. Head and pronotum with punctures of same
diameters; frons punctures of same size than clypeal size. Eyes narrow, semicircular. Antennae moniliform,
punctures, distributed uniformly. Eyes narrow, semicir- antennomere 11 slightly longer than 10. Pronotum
cular. Antennae moniliform, antennomere 11 slightly widest at midpoint, with punctures separated by more
longer than 10. Pronotum widest just behind midpoint, than two puncture diameters on the disc and separat-
with a central longitudinal groove and punctures small- ed by less than one puncture diameter at lateral quar-
er than frons punctures, separated by one to three punc- ters; lateral margin flattened, slightly curved upwards;
ture diameters; lateral margin marked, not elevated; anterior angles straight to obtuse, not produced; poste-
anterior angles acute, rounded, slightly produced; posterior angles obtuse; posterior margin straight, narrower
rior angles obtuse, rounded; posterior margin straight, than anterior margin. Elytron with nine longitudinal
wider than anterior margin. Elytron with nine carinae rows of punctures larger than pronotum punctures;
slightly raised and intervals with longitudinal rows of intervals carina-shape slightly raised withouth or with
large punctures slightly marked; on the carinae and small disperse punctures, turning backwards into rows
intervals with small disperse punctures uniformly dis- of tubercles and converging postero-laterally; humeri
tributed; humeri elevated, elytra fused along the midline not elevated, elytra fused along the midline and
and abdomen. Legs with punctures same size than abdomen. Legs with punctures same size than prono-
pronotum punctures; anterior margin of profemur wavy, tum punctures, anterior margin of profemur wavy,
lacking tooth. Abdomen with exposed intersegmental lacking tooth. Abdomen with exposed intersegmental
membranes apical to visible sternites three and four. membranes apical to visible sternites three and four.
Distribution. Argentina: Buenos Aires province; Distribution. Argentina: Northwest Mendoza and
Uruguay: Montevideo department (Kulzer 1955). Center-Southwest San Juan provinces; Chile: Los
Andes, Santiago, Cordillera, and Cachapoal provinces
(Silvestro et al. 2012).
Scotobius punctatus Eschscholtz, 1831
(Tenebrioninae, Scotobiini)
(Fig. 35)
Scotobius pilularius Germar, 1823
(Fig. 37)
width: 6.5–8.0 mm). Body black to reddish, dull, gla-
brous. Head and pronotum with punctures of same size,
separated by less than two puncture diameters uniform-
width: 7.6–9.9 mm). Body black, dull, glabrous. Head
ly distributed. Eyes narrow, semicircular. Antennae
punctate, on frons not uniform, on clypeus uniformly
moniliform, antennomere 11 slightly longer than 10. Pro-
distributed, separated by less than two puncture diam-
notum widest behind midpoint; lateral margin flattened,
eters, both of same size. Eyes narrow, semicircular.
slightly curved upwards; anterior angles acute, pro-
Antennae moniliform, antennomere 11 slightly longer
duced; posterior angles straight; posterior margin
than 10. Pronotum widest at midpoint, with punctures
straight, wider than anterior margin. Elytron with nine
same size than head punctures, separated by less than
longitudinal rows of punctures of same size or smaller
one puncture diameter, with two to four irregular,
than pronotum punctures; intervals carina-shape slight-
smooth and slightly raised areas; lateral margin flat,
ly raised with small disperse punctures, turning back-
slightly curved upwards; anterior angles rounded, not
wards into rows of tubercles and converging postero-lat-
produced; posterior margin straight, as wide as anteri-
erally; humeri not elevated, elytra fused along the mid-
or margin. Elytron with nine longitudinal rows of round
line and abdomen. Legs with punctures same size than
large tubercles, with only 10–16 tubercles per row;
pronotum punctures, anterior margin of profemur wavy,
humeri weakly elevated, elytra fused along the midline
lacking tooth. Abdomen with exposed intersegmental
and abdomen. Legs punctures lesser than pronotum
punctures; anterior margin of profemur wavy, lacking
Distribution. Argentina: South and West Mendoza
tooth. Abdomen with exposed intersegmental mem-
and West Neuquén provinces; Chile: Malleco province
branes apical to visible sternites three and four.
(Silvestro et al. 2012).
Distribution. Argentina: provinces Buenos Aires,
Chubut, Córdoba, Corrientes, Entre Ríos, Formosa, La
Scotobius andrassyi Kaszab, 1969 Pampa, La Rioja, Mendoza, Neuquén, Río Negro, San
(Tenebrioninae, Scotobiini) Luis, Tucumán; Uruguay: departments Canelones,
(Fig. 36) Lavalleja, Montevideo; Brazil: Rio Grande do Sul; Chile:
Cautín province (Kulzer, 1955, Zanetti et al. 2015a, b
Diagnosis. Medium size (length: 21.3–13.8 mm, cited as S. miliaris Billberg 1815, Silvestro and Flores
width: 8.9–5.7 mm). Body black to reddish, dull, 2016, this volume).
2 3 4
12
5 8 9
13
14
10 11 15
Figures 2–15. (2–5) Habitus: 2 – Lobopoda breyeri, upper left showing tarsal claws pectinate; 3 – Trichoton roigi; 4 – Blapstinus punctulatus;
5 – Conibius franzi; (6–7) Eyes emarginate anteriorly by epistomal canthus: 6 – Alphitobius diaperinus, 7 – Alphitobius laevigatus, (8–11)
Habitus: 8 – Alphitobius diaperinus, 9 – Alphitobius laevigatus, 10 – Vaniosus profana, 11 – Achanius antofagastensis (Figs 10 and 11
images from Flores and Aballay 2015: Figs. 2 and 14, reproduced with permission); (12–14) Head, frontal view showing mouthparts: 12 – Vaniosus
profana; 13 – Achanius (Ambigatus) bicolor Kulzer (images from Flores and Aballay 2015: Figs. 6 and 19, reproduced with permission);
14 – Salax lacordairei, (15) Habitus: 15 – Salax lacordairei. Scale bars 2 mm.
18
19
16 17 20
23
24
21 22 25
26 27 28 29
Figures 16–29. 16 – Hylithus tentyroides, 17 – Megelenophorus americanus; (18, 19) Head, ventral view showing mouthparts:
18 – Megelenophorus americanus, 19 – Praocis ecostata. White arrow indicates subgenal process; (20–22) Habitus: 20 – Praocis ecostata,
21 – Eutelocera cadaverina sp. nov. (paratypus), 22 – Hemasodes minutus, (23–24) Pronotum: 23 – Omopheres scabripennis,
24 – Omopheres difficilis, (25–39) Habitus: 25 – Omopheres scabripennis, 26 – Omopheres difficilis, 27 – Zophobas opacus,
28 – Gondwanocrypticus platensis, 29 – Poecilocrypticus formicophilus. Scale bars 2 mm.
30 31 32
33 34 35
36 37 38
Figures 30–38. (30) Leptynoderes strangulata, (31) Leptynoderes nordenskioldi, (32) Emmallodera perlifera, (33) Scotobius muricatus,
(34) Scotobius ovalis, (35) Scotobius punctatus (image from Silvestro et al. 2012: Fig. 1, reproduced with permission), (36) Scotobius
andrassyi (image from Silvestro et al. 2012: Fig. 3, reproduced with permission), (37) Scotobius pilularius, (38) Scotobius casicus. Scale
bars 2 mm.
Remarks. This species was known in publications midline and abdomen. Legs with punctures same size
as Scotobius miliaris since 173 years ago due to than pronotum punctures; anterior margin of profemur
incorrect synonymy (Silvestro and Flores 2016, this wavy, lacking tooth. Abdomen with exposed interseg-
volume). The taxonomic history can be found in Silve- mental membranes apical to visible sternites three and
stro and Flores (2016, this volume). four.
Distribution. Argentina: provinces Córdoba, San
Luis, Santiago del Estero, Tucumán (collection data
Scotobius casicus Lacordaire, 1830 IADIZA).
(Fig. 38)
Diagnosis. Large size (length: 21.9–30.0 mm, DISCUSSION

width: 9.2–12.0 mm). Body black, dull, elongated. Head
with punctures separated by less than two puncture The identification key and diagnoses of particular
diameters, frons punctures same size of clypeal punc- species presented in this paper are the first attempt to
tures. Eyes narrow, semicircular. Antennae monili- gather information concerning the species of Tenebri-
form, antennomere 11 slightly longer than 10. Prono- onidae associated with carcasses. The 29 species stud-
tum widest behind midpoint, with punctures same size ied here (Table 1) represent four of the nine currently
than frons punctures, separated by less than three recognized subfamilies of Tenebrionidae (Matthews et
puncture diameters, denser towards the margins and al. 2010): Alleculinae (one species of Alleculini), Dia-
in the central longitudinal groove slightly marked; lat- perinae (two species of Crypticini), Tenebrioninae (15
eral margin flattened, slightly curved upwards; anteri- species, ten belonging to the tribe Scotobiini, three to
or angles produced; posterior margin convex, narrow- Opatrini, two to Alphitobiini and one species of Tene-
er than anterior margin. Elytron with nine rows of con- brionini), Pimeliinae (10 species, three belonging to the
ical tubercles well defined even backwards and side- tribe Epitragini, two to Evaniosomini, two to Praociini,
ways; humeri not elevated, elytra fused along the mid- one species from each of the tribes Trilobocarini, Edro-
line and abdomen. Legs with punctures same size than tini and Elenophorini). The tribe Scotobiini is the most
numerous in this study, comprising species of the three
pronotum punctures; anterior margin of profemur
genera that occur in Argentina, all of which have been
wavy, lacking tooth. Abdomen with exposed interseg-
recorded feeding on corpses and rotting chicken flesh,
mental membranes apical to visible sternites three
which suggests that the members of this tribe have
and four.
a preference for feeding on cadaver tissue. It should be
Distribution. Argentina: provinces Buenos Aires
(Zanetti et al. 2015 a), Mendoza, Neuquén, San Luis
(Kulzer 1955), Santiago del Estero (collection data
IADIZA).
Scotobius granosus Solier, 1838

(Fig. 39)

width: 6.6–8.9 mm). Body black, dull, glabrous. Head
with punctures separated by less than two puncture
diameters; frons punctures larger than clypeal punc-
tures, uniformly distributed. Eyes narrow, semicircu-
lar. Antennae moniliform, antennomere 11 slightly
longer than 10. Pronotum widest behind midpoint,
punctures same size than frons, separated by less than
two puncture diameters, denser towards the margins
and in the central longitudinal groove; lateral margin
slightly marked, flat; anterior angles rounded, not pro-
duced; posterior margin straight, wider than anterior
margin. Elytron with nine longitudinal rows of round
tubercles, with more than 30 tubercles per row, not
aligned; humeri not elevated, elytra fused along the Figure 39. Scotobius granosus. Scale bar 2 mm.
noted that all species of the tribes Scotobiini, Eleno- americanus, Blapstinus punctulatus and Scotobius
phorini and Praociini are flightless, while the remain- pilularius are also omnivorous beetles, the first has
ing species of the other tribes are able to fly actively, been reported to eat dry fruits of Bulnesia retama
and therefore exhibiting different capabilities to find (Flores and Debandi 2004) and to feed on cow and horse
a carcass. carcasses (Aballay et al. 2008); the second has been
Twelve species of Tenebrionidae constitute new recorded feeding on seedlings of cotton, sunflower, cap-
records for the cadaver fauna in Argentina: Omo- sicum, potato, grapevine, corn and seeds of cotton and
pheres difficilis, O. scabripenis, Hemasodes min- corn (Flores and Debandi 2004), and in the present
utus, Zophobas opacus, Alphitobius laevigatus, study it was collected eating on lesser rhea, cow and pig
Conibius franzi, Eutelocera cadaverina sp. nov., carcasses (Table 1); and the third was reared on fruit
Scotobius granosus, S. ovalis, S. punctatus, S. an- (banana, apple), vegetables and meat in order to obtain
drassyi and Emmallodera perlifera. With exception eggs and larvae (Silvestro and Michat 2016, this vol-
of the last three species (collected only in chicken-bait- ume), and was also collected eating on pig carcasses
ed trap) all of them were collected mostly from carcass- (Zanetti et al. 2015 a, b as Scotobius miliaris).
es (Table 1). The remaining species associated with Blapstinus punctulatus and Scotobius pilular-
carcasses listed in this key were previously recorded ius were collected over a wide area, whereas Eutelo-
for the country in Buenos Aires (Zanetti et al. 2015 a, cera cadaverina sp. nov appears to have a restricted
b), Mendoza (Aballay 2012, Flores and Aballay 2015), distribution. Moreover, Praocis ecostata and Acha-
Catamarca (Flores and Aballay 2015), and San Juan nius antofagastensis exclusively inhabit high alti-
provinces (Aballay et al. 2008, 2012). Herein we pres- tude environments. Differences among species in geo-
ent here two new records for Argentina: Conibius graphic distribution and behavior may be attributed to
(Gondwanodilamus) franzi and Alphitobius laevi- the conditions characterizing the environment, such as
gatus; three new for Bolivia: Salax lacordairei, Bla- climate, vegetation and substrate (Cepeda-Pizarro
pstinus punctulatus, Vaniosus profana; and one 1989, Cartagena and Galante 2001). These conditions
new for Uruguay: Blapstinus punctulatus. determine adaptations of tenebrionids to arid and
Hylithus tentyroides has been listed in a catalog semiarid ecosystems since many species of the family
as being phytophagous, eating corn seed and cotton show a predisposition in to occupy dry habitats
seedlings (Flores and Debandi 2004), and in recent (Matthews et al. 2010). These beetles play an impor-
decomposition experiments it was recorded on pig car- tant role in these environments by facilitating nutrient
casses (Aballay et al. 2008, 2012, Zanetti et al. 2015 a, recycling (Santos et al. 1988, Cepeda-Pizarro 1989).
b). During the present study, this species was collected This variation in the distribution of species of Tenebri-
from horse, whale, pig, chicken carcasses, and human onidae can help as a forensic indicator in case of
cadavers, showing a wide distribution across Argenti- corpse transport.
na (from Salta to Chubut) and being recorded from sea Further studies are needed to state the specific time
level to 2200 m (Table 1). Alphitobius diaperinus has period in the cadaver succession that a species is in,
been collected from pig carcasses (Matuszewski et al. which would make the species cited in the present arti-
2008); this species can feed on dead chickens and bats cle more predictable, and which could be used to esti-
(Matthews et al. 2010) and has been considered a pred- mate PMI indicators based on succession patterns. In
ator on larvae and pupae of Musca domestica Lin- addition, immature stages of Tenebrionidae can be
naeus, 1758 (Despins et al. 1988); in the present re- useful in forensic entomology because they are reared
search, it was collected from pig carcasses and human in the corpses and collected in advanced stages of
corpses. Alphitobius diaperinus was recorded on decomposition (Lefebvre and Gaudry 2009, Introna et
human cadavers at the end of the advanced decay al. 2011, Aballay 2012, Mariani et al. 2014, Zanetti et
stage with presence of larvae III and pupae of Calli- al. 2015a, b) but the duration of larval development is
phoridae (Diptera), it can possibly feed on larvae and variable and depends on the species. Because informa-
pupae of Calliphoridae or on cadaver tissues. Salax tion about larvae of Tenebrionidae species is limited
lacordairei was also listed as phytophagous, eating (Doyen 1994), research studies should be conducted on
cotton seed and seedlings (Flores and Debandi 2004) their life cycle, and essential to this end is a correct
and has previously been collected from pig carcasses identification of adult Tenebrionidae. In fact, of the 29
(Aballay et al. 2008, 2012); during this study, it was col- species collected in this study, only the larvae of three
lected from pig, cow and human carcasses. This spe- of them are known: Alphitobius laevigatus (Klaus-
cies and Hylithus tentyroides likely feed on dry ca- nitzer 1996), Alphitobius diaperinus (Hayashi 1966)
daver tissue. Salax lacordairei, Hylithus tentyro- and Scotobius pilularius (Silvestro and Michat 2016,
ides and Alphitobius diaperinus constitute new re- this volume). We expect that this key will contribute to
cords on human corpses, but their potential as forensic future studies addressing the potential role of carrion-
indicators needs to be investigated. Megelenophorus frequenting Tenebrionidae as PMI indicators.
Table 1. List of Tenebrionidae species collected on vertebrate carcasses and from baited traps in Argentina and their geographic distribution by provinces.
722
Sampling method: m = hand on the carcass, pt = pitfall trap, bt = baited bottle trap
Species Substratum/ Sampling Province Geographic coordinates Altitude Collector/reference

N°
carcasses method (m)
Achanius antofagastensis 13 Pig m, pt Catamarca 26°01’32”S, 67°20’36”W 3578 Flores and Aballay (2015)
Alphitobius diaperinus 3 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 843 Aballay, F. (2012)
Alphitobius laevigatus 2 Pig m Mendoza 32°53’49”S, 68°52’24”W 843 Aballay, F. (2012)
Alphitobius diaperinus 28 Human m Mendoza 32°58’36”S, 68°37’24”W 653 Aballay, F. (forensic cases)
Blapstinus punctulatus 1 Cow m Mendoza 34°02’46”S, 67°54’37”W 565 Flores, G.
Blapstinus punctulatus 3 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 845 Aballay, F. (2012)
Blapstinus punctulatus 43 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Blapstinus punctulatus 1 Lesser rhea m Chubut 42°20’21”S, 63°42’05”W 25 Flores, G. Carrara, R. Cheli, G.
Blapstinus punctulatus 1 Pig m, pt San Juan 31°32’34”S, 68°34’38”W 676 Aballay et al. (2008)
Conibius franzi 5 Pig pt Mendoza 32°53’49”S, 68°52’24”W 845 Aballay, F. (2012)
Emmallodera perlifera 1 Chicken bt Mendoza 33°36’09”S, 69°18’45”W 1400 Jofré, F., Aballay, F.
Eutelocera cadaverina sp. nov. 46 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Gondwanocrypticus platensis 2 Pig pt Mendoza 32°53’49”S, 68°52’24”W 847 Aballay, F. (2012)
Gondwanocrypticus platensis 103 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Hemasodes minutus 19 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 853 Aballay, F. (2012)
Hylithus tentyroides 161 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Hylithus tentyroides 54 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 842 Aballay, F. (2012)
Hylithus tentyroides 1 Human m Mendoza 32°31’23”S, 68°56’4”W 1247 Aballay, F. (forensic cases)
Hylithus tentyroides 7 Chicken bt Mendoza 34°15’00”S, 69°21’45”W 2200 J Jofré, F., Aballay, F.
Hylithus tentyroides 1 Chicken bt Mendoza 33°36’58”S, 69°30’21”W 2600 Jofré, F., Aballay, F.
Hylithus tentyroides 2 Horse m Mendoza 35°49’01”S, 68°39’57”W 1877 Roig, S., Flores, G.
Hylithus tentyroides 1 Whale m Chubut 42°39’48”S, 64°59’22”W 2 Rojas Lanús, D.
Hylithus tentyroides 3 Pig m, pt San Juan 31°32’34”S, 68°34’38”W 675 Aballay et al. (2008, 2012)
F. H. ABALLAY, G. E. FLORES, V. A. SILVESTRO, N. I. ZANETTI and N. D. CENTENO
Leptynoderes nordenskioldi 17 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Leptynoderes strangulata 2 Pig m, pt Catamarca 26°01’33”S, 67°20’42”W 839 Aballay, F. (2012)
Lobopoda breyeri 1 Pig m Mendoza 32°53’49”S, 68°52’24”W 846 Aballay, F. (2012)
Megelenophorus americanus 5 Cow m San Juan 32°13’00”S, 67°47’38”W 520 Aballay et al. (2008)
Megelenophorus americanus 1 Horse m San Juan 31°18’36”S, 68°26’31”W 1392 Aballay et al. (2008)
Table 1. Continued.
Species Substratum/ Sampling Province Geographic coordinates Altitude Collector/reference

N°
carcasses method (m)
Omopheres difficilis 6 Pig m Mendoza 32°53’49”S, 68°52’24”W 851 Aballay, F. (2012)
Omopheres scabripennis 11 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 852 Aballay, F. (2012)
Omopheres scabripennis 3 Chicken bt Mendoza 34°06’29”S, 69°06’42”W 1400 Jofré, F., Aballay, F.
Poecilocrypticus formicophilus 3 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 844 Aballay, F. (2012)
Praocis ecostata 1 Llama m Catamarca 26°01’32”S, 67°20’36”W 3578 Aballay, F. (2012)
Salax lacordairei 205 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 841 Aballay, F. (2012)
Salax lacordairei 1 Human m Mendoza 32°30’17”S, 68°09’08”W 572 Aballay, F. (forensic cases)
Salax lacordairei 2 Cow m San Juan 31°32’54”S, 67°25’27”W 560 Aballay, F.
Salax lacordairei 15 Pig m, pt San Juan 31°32’34”S, 68°34’38”W 673 Aballay et al (2008, 2012)
Salax lacordairei 6 Cow m San Luis 32°22’08”S, 67°09’37”W 551 Aballay, F.
Scotobius casicus 25 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Scotobius granosus 1 Cow m Cordoba 33°06’35”S, 64°17’54”W 432 Gómez, R.
Scotobius pilularius 176 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Scotobius muricatus 42 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al (2015a)
Scotobius ovalis 166 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
Scotobius punctatus 2 Chicken bt Mendoza 34°14’20”S, 69°24’23”W 2509 Aballay, F.
Scotobius punctatus 8 Chicken bt Mendoza 34°15’00”S, 69°21’45”W 2200 Jofré, F., Aballay, F.
Scotobius punctatus 2 Chicken bt Mendoza 34°15’28”S, 69°17’32”W 1800 Jofré, F., Aballay, F.
Scotobius andrassyi 6 Chicken bt Mendoza 32°30’13”S, 69°03’11”W 2600 Jofré, F., Aballay, F.
Scotobius andrassyi 3 Chicken bt Mendoza 32°29’08”S, 69°06’55”W 3000 Jofré, F., Aballay, F.
Trichoton roigi 8 Pig m, pt San Juan 31°32’34”S, 68°34’38”W 674 Aballay et al. (2008, 2012)
Trichoton roigi 189 Pig pt Mendoza 32°53’49”S, 68°52’24”W 840 Aballay, F. (2012)
Trichoton roigi 6 Pig m, pt Buenos Aires 38°41’41”S, 62°15’10”W 70 Zanetti et al. (2015a)
AN ILLUSTRATED KEY TO SPECIES OF TENEBRIONIDAE IN ARGENTINA
Vaniosus profana 91 Pig m, pt Mendoza 32°53’49”S, 68°52’24”W 849 Flores and Aballay (2015)
Zophobas opacus 1 Fox m Chaco 26°3005”S, 61°11’39”W 124 Gómez, R.
723
ACKNOWLEDGMENTS Caballero, U. and J. L. León-Cortés. 2014. Beetle succession

and diversity between clothed sun-exposed and shaded
pig carrion in a tropical dry forest landscape in Southern
We acknowledge directors of Medical Forensic
Mexico. Forensic Science International, 245: 143–150.
Committee of Mendoza and medical Hugo San Martino, Cabrera, A. L. and A. Willink. 1980. Biogeografía de América
Eduardo Berte and Javier Salinas for the collecting Latina, 2nd ed. Monografía N°13, Serie Biología, OEA. Se-
permission on human carcasses; Nelly Horak for cor- cretaría General de la OEA, Washington, DC, 122 pp.
rection of the English language; Mariana Chani Posse Cartagena, M. C. and E. Galante. 2001. Ecología de Tentyria
for suggestions improving the manuscript; Morales laevis Solier, 1835 y T. peiroleri Solier, 1835 (Coleopte-
family (Antofagasta de la Sierra, Catamarca) for their ra, Tenebrionidae, Tentyriini). Sessio Conjunta d’Entomo-
willingness to carry out the field study; Ana María Scol- logia ICHN SCL, 11: 25–34.
lo for mounting specimens; Daniel Rojas Lanús and Cecco, L., González, H., Deluchi, P., Barrios, H. and M. De
Rodrigo Gómez for his collecting efforts; Rodolfo Car- Franceschi. 2005. Determinación de los estados de des-
arrollo de Alphitobius diaperinus en granjas avícolas.
rara for GIS assistance; Patrice Bouchard and two
Revista Argentina de Producción Animal, 25: 93–99.
anonymous reviewers for suggestions improving this
Centeno, N. D., Maldonado, M. and A. Oliva. 2002. Seasonal
paper. This study was supported by the Consejo
patterns of arthropods occurring on sheltered and unshel-
Nacional de Investigaciones Científicas y Técnicas tered pig carcasses in Buenos Aires Province (Argentina).
(CONICET, Argentina) and in different periods of its Forensic Science International, 126: 63–70.
carrying out by grants from the Agencia Nacional de Cepeda-Pizarro, J. G. 1989. Actividad temporal de tenebrióni-
Promoción Científica y Tecnológica (ANPCYT, Argenti- dos epígeos (Coleoptera) y su relación con la vegetación
na): PICT 38127; PICT 26000; PICT 2013–514: Universi- arbustiva en un ecosistema árido de Chile. Revista Chile-
dad Nacional de Quilmes; PIP 112-201101-00987 (CON- na de Historia Natural, 62: 115–125.
ICET, Argentina), and Servicios Tecnológicos de Alto Correa, R. C., Almeida, L. M. and M. O. Moura. 2014.
Nivel (STAN CONICET N°826). Coleoptera associated with buried carrion: potential
forensic importance and seasonal composition. Journal of
Medical Entomology, 51(5): 1057–1066.
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An Illustrated Key to, and Diagnoses of the Species of Tenebrionidae

Article in Annales Zoologici · December 2016

Fernando Hernán Aballay Violeta A. Silvestro

Noelia Zanetti Nestor Centeno

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AN ILLUSTRATED KEY TO, AND DIAGNOSES OF

FERNANDO H. ABALLAY1,*, GUSTAVO E. FLORES1, VIOLETA A.

1Laboratorio de Entomología, Instituto Argentino de Investigaciones de las Zonas

Key words.— darkling beetles, Alleculinae, Diaperinae, Tenebrioninae, Pimeliinae,

INTRODUCTION carcasses, and its forensic importance has been fre-

PL ISSN 0003-4541 © Museum and Institute of Zoology PAS

RESULTS –. Eyes emarginate anteriorly by epistomal canthus

Diagnosis. Large size (length: 29.9–40.7 mm, width:

Omopheres difficilis (Steinheil, 1874)

Diagnosis. Large size (length: 22.9–29.3 mm, width:

Emmallodera perlifera Burmeister, 1875

Diagnosis. Large size (length: 21.9–30.0 mm, DISCUSSION

Scotobius granosus Solier, 1838

Diagnosis. Medium size (length: 15.2–19.9 mm,

Species Substratum/ Sampling Province Geographic coordinates Altitude Collector/reference

Species Substratum/ Sampling Province Geographic coordinates Altitude Collector/reference

ACKNOWLEDGMENTS Caballero, U. and J. L. León-Cortés. 2014. Beetle succession

Received: June 14, 2016

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