Trogidae

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Scarabaeiformia
Superfamilia: Scarabaeoidea

Familia: Trogidae
Subfamiliae: Troginae - †Avitortorinae - †Prototroginae

Overview of genera

Afromorgus – Glyptotrox – Madagatrox – Omorgus – Polynoncus – Trox – †Avitortor – †Cretomorgus – †Prototrox

[source: Catalogue of Life: 2011 Annual Checklist, plus (3) Madagatrox – †Avitortor – †Cretomorgus
]
Name

Trogidae MacLeay, 1819
References

Smith, A.B.T. 2006. A review of the family-group names for the superfamily Scarabaeoidea (Coleoptera) with corrections to nomenclature and a current classification. Coleopterists Society monograph, 5: 144–204. DOI: 10.1649/0010-065X(2006)60[144:AROTFN]2.0.CO;2 PDF.

Additional references

Bezděk, A. & Hájek, J. 2009. Catalogue of type specimens of beetles (Coleoptera) deposited in the National Museum, Prague, Czech Republic - Scarabaeoidea: Bolboceratidae, Geotrupidae, Glaphyridae, Hybosoridae, Ochodaeidae and Trogidae. Acta Entomologica Musei Nationalis Pragae 49(1): 297–332. PDF Reference page.
Diéguez, V.M. 2008. Present knowledge of the Trogidae of Chile (Coleoptera: Scarabaeoidea). Revista chilena de entomología, 34: 11–28. PDF.
Nikolajev, G.V. 2016. Taxonomic composition of the family Trogidae (Coleoptera: Scarabaeoidea) of the Russian fauna. Caucasian Entomological Bulletin 12(1): 81–91. PDF. (in Russian, with English abstract.) Reference page.
Pittino, R. 2010: A new genus and species of Trogidae MacLeay, 1819 from Madagascar, and a new species of Omorgus Erichson, 1847 from Argentina (Coleoptera, Trogidae). Kogane, (11): 75-83.
Catalogue of Palearctic Coleoptera. Vol. 3 ed. I. Lobl, & A. Smetana, Apollo Books, Stenstrup, Denmark, 2006. ISBN 87-88757-59-5, p. 79
Scholtz, C.H. 1982. Catalogue of world Trogidae (Coleoptera: Scarabaeoidea). Entomology memoir Department of Agriculture and Fisheries Republic of South Africa 54: 1–27.
Scholtz, C.H. 1986. Phylogeny and systematics of the Trogidae (Coleoptera: Scarabaeoidea). Systematic entomology 11: 355–363. DOI: 10.1111/j.1365-3113.1986.tb00186.x
Scholtz, C.H. 1990. Revision of the Trogidae of South America (Coleoptera: Scarabaeoidea). Journal of natural history 24(6): 1391–1456. DOI: 10.1080/00222939000770841
Strümpher, W.P. et al. 2014: Phylogeny of the family Trogidae (Coleoptera: Scarabaeoidea) inferred from mitochondrial and nuclear ribosomal DNA sequence data. Systematic entomology 39(3): 548-562. DOI: 10.1111/syen.12074 Reference page.
Zídek, J. 2013: Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea). Insecta mundi, (0314) Reference page.

Links

Australian Faunal Directory
Trogidae in SYNOPSIS OF THE DESCRIBED COLEOPTERA OF THE WORLD
Tree of Life Web Project. 2007. Trogidae. Hide beetles. Version 02 May 2007 (temporary) in The Tree of Life Web Project
Family Trogidae (atlas of beetles of Russia)

Vernacular names
Deutsch: Erdkäfer
English: hide beetles
한국어: 송장풍뎅이과
lietuvių: Smiltvabaliai
norsk: Knokkelbiller
русский: Падальники

Trogidae, sometimes called hide beetles, is a family of beetles with a distinctive warty or bumpy appearance. Found worldwide, the family includes about 300 species contained in four or five genera.[1]

Trogids range in length from 2.5 to 20.0 mm. Their shape is oblong to oval, with a generally flat abdomen. Their color ranges from brown to gray or black, and they often encrust their bodies with soil. They resemble scarab beetles with heavy limbs and spurs.

They are scavengers and are among the last species to visit and feed on carrion. They are most often found on the dry remains of dead animals. Both adults and larvae eat feathers, fur, and skin. Some species are found in bird and mammal nests. Details of the life histories of many species are poorly known, since many are specialized to particular types of nests. They are often overlooked by predators and collectors due to their behaviors of covering their bodies with soil and becoming motionless when disturbed.

This group may also be considered Troginae, a subfamily of the Scarabaeidae. The common name "skin beetle" is sometimes used in reference to these beetles, but more often refers to species of the Dermestidae.

Origins

Hide beetles are found worldwide. More species of Trogidae are found in dry environments instead of moist environments, typically temperate plains areas. Each genus is more diverse in different regions of the world. Trox is found in the Holarctic/Ethiopian area, Omorgus within the southern continents, and Polynoncus is found in South America.[2]

Controversy exists over whether Trogidae is a family of its own or a subfamily of Scarabaeidae.[2] One major reason for the dispute between classifications is the possible evolution of the ommatidium in the eyes . Different environmental pressures and predators may have led to the adaptation of ommatidium structures within this family. For example, the more advanced and numerous the ommatidium, the more present the larger the ability of the insect to escape and elude predators. The Trogidae may have evolved in Australia.[3]

The oldest known fossils are from the Aptian Zaza Formation of Russia and the Shar-Tolgoy and Dzun-Bain Formations of Mongolia, assignable to the extant genus Trox as well as the extinct genera Cretomorgus and Paratrox. Other known fossils include Kresnikus beynoni found in Cenomanian aged Burmese amber.[4]
Anatomy

Trogidae are characterized by their distinct soil-encrusted, warty, or bumpy appearance. They are usually brown, gray, or black in color and are covered with short, dense setae. Their body shape is oblong to oval with a flat abdomen and their length varies from 2 to 20 mm. The antenna of hide beetles are usually short and clubbed.[5] The hardened elytra of Trogidae, which are generally covered with small knobs giving the beetle their rough appearance, meet along the midline of the body and cover the entire abdomen and well-developed wings. Their heads are bent down and covered by the pronotum.[6] They also have heavy limbs and spurs resembling those of scarab beetles. Trogidae larvae are a creamy yellow/white in color, except at their caudal end which darkens as it accumulates feces. Their heads are dark and heavily sclerotized. The abdominal segments of hide beetles have at least one or more transverse rows of setae.[7]
Diet and habitat

Predators rarely attack species of Trogidae. They avoid detection and predation due to their soil covering and motionless behavior. Birds prey upon hide beetles that have invaded the bird nests.[7]

Species of Trogidae often feed on decomposing carcasses.[8] In one lab experiment done in 1998 by the Department of Zoology at the University of Melbourne, the hide beetles ate all tissues on a sheep carcass and left the bones. Along with carrion, hide beetles are found within the pellet of many animal species, on other decaying dry matter, and around birds’ and mammals’ nests and feathers, as well as aging bones.
Mating habits and life cycle

Little is known about the specifics of the Trogid life cycle. Female Trogids of several species produce chirping noises in order to attract males to their burrows for mating.[9] After impregnation of the female by the male, the female lays the eggs and the larvae hatch after an unknown amount of time. During decomposition of a carcass, the beetles leave their nests to feed on the carrion. As the last succession of insects to appear on the carcass, both larvae and adults can be found feeding on the dry remains. At the site of the carcass, an impregnated female digs small, vertical columns underneath the carcass to lay her eggs, allowing the larvae to locate food after hatching. Trogidae usually have three to five instars.[7]
Forensic importance

The use of Trogidae in forensic entomology is unknown at this time. Though they typically arrive last in the order of succession, they can be the first in succession on burned and charred bodies. After the burned skin is eaten away by the trogids, the corpse (with now-exposed, "fresher" surfaces) allows for viable colonization by other forensically important insects that help determine accurate post mortem interval estimates.[10]

Various species of Trogidae have been used by museums to clean up skeletons by eating any remaining dried material left on the skeletons, leaving them clean for display. This method of bone-stripping has been used by some museums for many years, as it is the most effective method.[11]
Research

The Chinese Academy of Sciences funded a study on the classification of this family of beetles.[12] The forensic importance of African Trogidae and other carrion-associated beetles is being studied at the University of Pretoria. This project is investigating how the presence of beetles on carrion affects the infestation of other arthropods in carrion in Africa.[13]
Taxonomy

The family Trogidae contains the following subfamily and genera:[14]

†Avitortorinae Nikolajev, 2007
†Avitortor Ponomarenko, 1977
†Kresnikinae Tihelka, Huang & Cai, 2020
†Kresnikus Tihelka, Huang & Cai, 2020
Omorginae Nikolajev, 2005
†Cretomorgus Nikolajev, 2007
Omorgus Erichson, 1847
Polynoncus Burmeister, 1876
†Prototroginae Nikolajev, 2000
†Prototrox Nikolajev, 2000
Troginae MacLeay, 1819
Glyptotrox Nikolajev, 2016
†Paratrox Nikolajev, 2009
Phoberus MacLeay, 1819
Trox Fabricius, 1775

References

Jameson, Mary Liz (2002): "Trogidae", in Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2002), vol. 2
Deloya, C. (2005): Omorgus rodriguezae especie nueva de México y clave para separar las especies del género para centro y norteamérica (Coleoptera: Trogidae) Archived 28 September 2007 at the Wayback Machine. Folia Entomol. Mex 44: 121-129. PDF
LAWRENCE, J. F. and E. B. BRITTON. 1991. Coleoptera. The Insects of Australia, 2nd edition, Volume 1, pp. 543-683. Melbourne University Press, Carlton.
Tihelka, Erik; Huang, Diying; Cai, Chenyang (15 July 2019). "A new subfamily of hide beetles from the Cretaceous of northern Myanmar (Coleoptera: Scarabaeoidea: Trogidae)". Historical Biology: 1–8. doi:10.1080/08912963.2019.1641705. ISSN 0891-2963.
Watson, L. and Dallwitz, M. J.. 2003 onwards. British Insects: the Families of Coleoptera: Trogidae Version: 9 April 2007. http://delta-intkey.com
Carcass Beetles CSIRO Entomology
Jameson, Mary Liz. Guide to New World Scarab Beetles - Trogidae UNL State Museum - Division of Entomology
Murawski, Darlyne (2013). Ultimate Bug-opedia. Washington, DC: National Geographic Society. p. 153. ISBN 9781426313776.
Alexander, Richard D., Thomas E. Moore, and Robert E. Woodruff. "The evolutionary differentiation of stridulatory signals in beetles (Insecta: Coleoptera)." (1963).
Trogidae[permanent dead link] Australian Biological Resources Study, Department of the Environment and Heritage
Reid, Craig. Bones Stripped Bare. Australian Geographic; Jan-March2005 Issue 77, p15, 1/4p
GuoDong, Ren and Hou Lin. Advance in taxonomic research of the Trogidae. Entomological Knowledge, 2003 (Vol. 40) (No. 6) 505-508
Williams, K.A. and M.H. Villet. A history of South African research relevant to Forensic Entomology. South African Journal of Science 102, January/February 2006. p4.
"Hide Beetles MacLeay, 1819". BioLib.