Family TENEBRIONIDAE Latreille, 1802
Compiler and date details
31 December 2004 - E.G. Matthews, South Australian Museum, North Terrace, Adelaide 5000 [matthews.eric@saugov.sa.gov.au] and P. Bouchard, Agriculture and Agri-Food Canada, Entomology, K. W. Neatby Building, 960 Carling Avenue, Ottawa, ON, K1A 0C6 [bouchardpb@agr.gc.ca]
- Tenebrionites Latreille, 1802.
Type genus:
Tenebrio Linnaeus, 1758.
Introduction
The following checklist includes 1,601 described species of Australian Tenebrionidae (plus 453 synonyms) arranged in 209 genera (200 synonyms) and 11 subgenera, 41 tribes, 13 subtribes, and 8 subfamilies, making this the fifth largest family of beetles on the continent after Curculionidae, Scarabaeidae, Chrysomelidae, and Carabidae. The last published checklist of Australian tenebrionid species is that of Matthews & Bouchard (2008) which lists 1,595 species, 216 genera and subgenera, and 8 subfamilies.
Tenebrionidae is the largest family in the superfamily Tenebrionoidea which comprises 30 world families in the classification of Lawrence & Newton (1995). Analyses by Lawrence (1994) and Slipinski & Lawrence (1999) suggest that the closest families to Tenebrionidae are Chalcodryidae, a small group of three genera now restricted to New Zealand, and Perimylopidae with seven genera occurring in Tasmania and the southern cool temperate parts of South America. These authors did not consider Trachelostenidae, with one Chilean genus, which Watt (1987) suggested may have some relationship to Tenebrionidae. The fact that all its probable relatives are south-temperate groups is an indication that Tenebrionidae itself may be of Gondwanan origin.
Typical tenebrionids may be recognised by their antennal insertions covered by shelf-like extensions of the genae, the prosternal process not strongly expanded behind the procoxae, a 5-5-4 tarsal formula, the abdomen with the first three visible ventrites immovable and the last two hinged by exposed membranes. These membranes are indicative of the presence of paired defensive glands that are unique within the Tenebrionoidea. Problems arise when there are exceptions to the tarsal formula, which may be 4-4-4 in very small tenebrionids, and when there are no visible abdominal membranes. The latter condition may occur in the Toxicini, even though defensive glands are present, and in the Pimeliinae, Belopini and Cossyphini (all of which are rare in Australia) where they are absent. It is probable that absence of the defensive glands is primitive in the family and that such glands arose between the 7th and 8th abdominal segments at least twice independently (Tschinkel & Doyen 1980) and between the 8th and 9th segments at least once (in the tribe Adeliini). The chemistry of the defensive secretions was analysed by Brown et al. (1992).
The classification of the family adopted here has evolved from the morphological and phylogenetic studies of Watt (1974), Doyen & Lawrence (1979), Doyen & Tschinkel (1982), Doyen et al. (1990), and Matthews & Bouchard (2008) as well as other works of more limited scope, and is probably close to the general consensus at the present time, being practically the same as that adopted by Aalbu et al. (2002) for the North American fauna. There are three major lineages, termed branches here and divisions in Aalbu et al.: the pimelioid, lagrioid and tenebrionoid. No doubt changes will occur in the future as knowledge increases. For instance, the ostensibly pimelioid Zolodininae, which occur only in Tasmania and New Zealand, will probably be found to be a separate lineage, perhaps the most basal in the family, and the lagrioid branch is evidently a paraphyletic cluster of lines associated only by shared primitive characters.
Larvae are typified by an elongate subcylindrical 'mealworm' form with few hairs except in Lagriini, and distinguished from the similar 'wireworms' (Elateridae) by having a distinct clypeus and labrum. This larval form evolved to pursue a fungivorous existence in decaying wood, which in common with that of other tenebrionoids, is the ancestral habitat in which many still live (e.g. Zolodininae, Stenochiinae, Alleculinae, Phrenapatinae, Gnathidiini, Ulomini, Amarygmini, Heleini-Cyphaleina). From rotten wood it became possible to move into other substrates such as soil (Opatrinae, Heleini-Heleina, Isopteron of the Adeliini), dry sand (Pimeliinae, Hyociini, Ectychini), leaf litter (most Adeliini), fungal fruiting bodies (Toxicini, Bolitophagini, Diaperini), storage chambers of vertebrates (Alphitobiini, Tenebrionini, Triboliini, some Diaperini and Palorini) and of social insects (Belopini, Cnemeplatiini, some Palorini, Amarygmini, Hyociini, Ectychini and Heleini)) , and surface grazing on trees (Titaenini, Leiochrinini, a few Amarygmini). In the arid zone, species of Platydema (Diaperini) are abundant in the nests of zebra finches. In all these habitats the actual food substance is nearly always fungi or dead plant matter. Animal matter as food, mainly in the form of carrion and faeces, is rare but it is known in Alphitobius, the lesser mealworm common in aviaries, and in desert foraging groups such as adult Heleina. Corticeus of the Hypophlaeini lives in association with scolytids under bark and has been thought to prey on the latter, but it appears now that this is unlikely (Doyen & Lawrence 1979). Some soil-inhabiting larvae have become agricultural pests as live-root feeders (Celibe, Pterohelaeus, Gonocephalum, Isopteron)—(Allsopp 1979, 1986) and as surface feeders on seedlings and strawberries (Adelium, Isopteron). The problem has become exacerbated of late by the widespread adoption of zero-tillage or stubble-retention practices, which leave a lot of plant cover on the soil surface in order to prevent erosion and encourage worms.
Tenebrionid pupae are rarely encountered in nature because of the comparatively short duration and the sedentary nature of the pupal stage. For this reason, only a very small number of pupae have been formally described or illustrated. Steiner (1995) was the first to summarise what is known about the morphology of pupae within the entire family. Of the 125 genera included in Steiner's study, 25 occur in Australia (including the introduced Alphitobius, Blaps, Phaleria, Alphitophagus, and Gnatocerus). Bouchard & Steiner (2004) recently described Australian pupae belonging to the subfamily Coelometopinae.
Tenebrionid pupae are unique in having lateral extensions of the abdominal segments that serve as a defence mechanism in this otherwise immobile life stage. Pairs of coapted, sclerotised 'jaws' on the abdomen of the pupae comprise what is known as the 'gin-trap' defence system. When small arthropods, such as mites, crawl on the body of the pupa and touch the sensilla in cavities between opposing structures of the gin-trap, a reflex is initiated that causes the eventual crushing of the crawling arthropod. This reflex system is thought to protect the pupae from potential predators and parasites during their otherwise defenceless life stage.
While most major tenebrionid groups are represented in Australia, the proportional composition of the fauna here is unique in the world. Most notable is the small number of Pimeliinae, a group that predominates elsewhere, particularly in arid regions. In the world as a whole, Pimeliinae make up half the diversity of the entire family, whereas in Australia native pimeliines are restricted to just three genera with a handful of species in the arid zone: Thorictosoma, Lixionica and Exangeltus. Another genus added recently, Stenosida, is probably based on mislabeled specimens, and two other pimeliines, Ocnera and Salax, are known introductions. Thorictosoma belongs to the Cnemeplatiini, a group which Watt (1992) and Doyen (1993) consider to be the most basal (primitive) of the pimeliines, that is to say, one which probably evolved earliest. Matthews (2000) suggested that Cnemeplatiini, Belopini and some other xerophilic tenebrionids lived in coastal sand dunes along the shores of the Tethys Sea before there was an inland arid zone in Australia, and that they now survive here as the oldest of the arid-zone tenebrionid elements primarily by living in ants' nests. All indications are that Pimeliinae is a very ancient group, but that it was barely able to gain a foothold in Australia before the break-up of Gondwana because of the early absence of deserts here.
Another arid-adapted tenebrionid group, the Opatrini, is very diverse world-wide especially in Africa but is represented by only a few genera in Australia which appear to have invaded from the north late in the Tertiary.
The niches associated with Pimeliinae and Opatrini elsewhere in the world are occupied in Australia by the Tenebrioninae-Heleini and to a lesser extent the genus Isopteron of the Lagriinae-Adeliini. Heleini is a dominant group here in sclerophyll forest and the arid zone, which formed only after desertification began in the mid-Tertiary, and it is this preponderance of Tenebrioninae (evidently a Gondwanan group), coupled with the scarcity of Pimeliinae, which helps to give the Australian tenebrionid fauna its unique character. The quintessential Australian tenebrionid is the pie-dish beetle genus Helea, with many surface-foraging species, which evolved from an ancestor already flattened in order to move under the bark of Eucalyptus trees (Matthews 1993). The wide lateral flanges of Helea serve to shield the head and legs from attack during foraging. Another major characteristic Australian element is Adeliini, richly represented in forests, particularly rainforests. Adeliini also occurs in New Zealand, New Caledonia and Chile, and is clearly one of the groups that flourished in Gondwana. Lastly among the uniquely Australian taxa, we should mention the xerophilic Hyociini and Ectychini of the Diaperinae, both endemic to Australia or nearly so. They are related to the littoral Phaleriini (present in Australia only as one introduced species) and appear to have evolved from ancient coastal sand dune inhabitants.
Also well represented in Australia are groups which are not unique or endemic— too many to list here but we might single out the Tenebrioninae-Amarygmini, Alleculinae and Stenochiinae, all of which are very diverse, mainly in forested habitats. Alleculinae appear among Jurassic fossils in Russia and therefore must have been present in Gondwanan Australia, whereas Amarygmini and Stenochiinae are clearly post-Gondwanan invaders from the north. The latter have even managed to penetrate arid environments in the form of the genus Hypaulax.
Early workers on Australian Tenebrionidae were based originally in France in the early 19th century (A. Latreille and J.B. de Lamarck, who described the specimens brought back by F. Péron on the Baudin expedition, including the first known species of Helea, J.B.A. Boisduval and E. Blanchard, who described the specimens collected on the two Astrolabe expeditions, and T. Lacordaire who established the first comprehensive classification of the family), then in England in the mid 19th to early 20th centuries (W. Kirby, F.P. Pascoe, C.O. Waterhouse, F. Bates, G.C. Champion and K.G. Blair, who described many taxa). During this period also the first Australian workers were W.J. Macleay, T. Blackburn and A.M. Lea, who described tenebrionids in the course of their work on Coleoptera in general, whilst the first worker to specialise on Australian Tenebrionidae was H.J. Carter who worked from 1905 to 1940, naming over 600 Australian species. The most important cataloguer was H. Gebien who compiled a world catalogue of Tenebrionidae (Gebien 1937-1944). After the second world war, descriptive work continued mainly by H. Kulzer in the 1960s, Z. Kaszab until the 1980s and H.J. Bremer and J. Ferrer at the present time. In recent years the emphasis has shifted to studies on higher classification and phylogeny in an attempt to establish a natural system for the family on a world basis. The most significant pioneering work in this field was that of Watt (1974), who integrated larval and adult data for the first time. This was followed by the seminal works of Doyen & Lawrence (1979), Tschinkel & Doyen (1980) and Doyen & Tschinkel (1982) who discerned major evolutionary lineages based largely on the defensive gland and female reproductive systems, as well as larval characters. The latter work, in particular, established the criteria on which the modern classification of the family is based. For Australia, Doyen et al. (1990) applied these criteria to propose a classification of the family and a list of genera. Matthews (1992, 1993 and 1998) then revised the tribes Heleini and Adeliini at generic level on Hennigian principles, Bouchard & Yeates (2001) published a phylogenetic analysis of the genera of Stenochiinae, and Bouchard (2002) did a phylogenetic revision of the rainforest-inhabiting genus Apterotheca of the Cnodalonini.
Persons wishing to identify Australian tenebrionids down to generic level can refer to Matthews & Bouchard (2008). Among keys to species we can mention Bremer (2006 and other works) (Amarygmini), Grimm 2013 (Cryphaeus), Kaszab 1961a and 1961b (Leiochrinini), 1973 (Scotoderus), 1977a and 1978 (Phrenapatini and Gnathidiini), 1977b (Strongyliini), 1982a (Lorelus), 1982b (Uloma), 1987 (Daerosphaerus), and 1988 (Promethis), Halstead 1967 (Palorini), Lawrence & Medvedev 1982 and Medvedev & Lawrence 1983, 1984, 1986 and 1987 (Hyociini), Matthews 1986 (Brises), Merkl 1986 and 1987 (Lagriini), Matthews & Doyen 1989 (Bassianus and Kaszaba), Merkl & Kompantzeva 1996 (Rhipidandrus), Scupola 2000 (Cossyphus) and Bouchard 2002 (Apterotheca).
ACKNOWLEDGEMENTS
We wish to thank the following colleagues who helped with problems of nomenclature, classification and literature search: V.R. Bejsak (University of Sydney), H. J. Bremer (Wellingholzhausen, Germany), J. Ferrer (Haninge, Sweden), R. Grimm (Tübingen, Germany), H. Labrique (Muséum d'Histoire naturelle, Lyon, France), J.F. Lawrence (Australian National Insect Collection), O. Merkl (Hungarian Natural History Museum), C.A.M. Reid (the Australian Museum), W. Schawaller (Staatliches Museum für Naturkunde, Stuttgart, Germany), and W. Steiner, Jr (National Museum of Natural History, Washington DC, USA).
J. Evans, librarian at the South Australian Museum, was very helpful in the hunt for literature, and the Museum's excellent library holdings, particlarly those from the 19th century, made the task of compiling this checklist much easier.
A. Wells, Australian Biological Resources Study, is sincerely thanked for struggling with the difficult task of rendering the original Microsoft Word checklist in internet-compatible format using the program Platypus.
Financial assistance for preparing this list was provided by the Australian Heritage Division of the Department of Environment and Heritage, and the work on Australian Tenebrionidae was supported by a grant from the Australian Biological Resources Study.
Notes
TAXA ERRONEOUSLY RECORDED FROM AUSTRALIA
The following species have been recorded as occurring in Australia based on mislabeled specimens. They have been, or should be, removed from Australian lists and are not included in the following checklist.
Arcothymus coenosus Pascoe, 1866: 476 (Lagriinae-Adeliini). Australia [erroneous]. Holotype BMNH. Syn. with Arcothymus tristis (Montrouzier, 1860: 302) (New Caledonia) by Bates 1872: 99. Arcothymus Pascoe is endemic to New Caledonia (Kaszab 1982a: 228).
Bradymerus antennatus Kulzer, 1951: 544 (Tenebrioninae-Toxicini). N. Australia [erroneous]. Holotype MGF. Syn. with Calymmus berardi (Montrouzier, 1860: 289) (New Caledonia) by Kaszab 1982: 52. Calymmus Montrouzier is endemic to New Caledonia (Kaszab 1982: 50).
Cymbeba dissimilis Pascoe, 1866: 484 (Lagriinae-Adeliini). Australia [erroneous]. Syntypes BMNH. Cymbeba Pascoe is endemic to New Caledonia (Kaszab 1982: 240).
Derispia coccinelloides (Westwood, 1883: 75) (Leiochrodes) (Diaperinae-Leiochrinini). Ceylon. Holotype BMNH. Listed in Carter 1926: 135 but not Australian (Kaszab 1961: 169).
Ecripsis pubescens Pascoe, 1866: 456 (Tenebrioninae-Opatrini). Tas [erroneous]. Syntypes BMNH. Syn. with Ammidium ciliatum Erichson, 1843: 251 (Angola) by Carter 1914: 406. Ammidium Erichson is endemic to Angola (Koch 1956: 49).
Emeax sculpturatus Pascoe, 1866: 450 (Pimeliinae-Nyctoporini). NSW [erroneous]. Syntypes BMNH. Syn. with Nyctoporis cristata Eschscholtz, 1831: 11 (California) by Carter 1914: 406. Nyctoporis Eschscholtz is endemic to the SW United States (Gebien 1937: 698).
Luprops atronitens Fairmaire, 1883: 27 Lagriinae-Lupropini). New Guinea. Syntypes not located. Erroneous Australian locality in Carter 1933: 170. Luprops Hope does not occur in Australia (Doyen et al. 1990: 239).
Menearchus impressosulcatus Carter, 1920: 230 (Opatrinae-Platynotini). NSW [erroneous]. Holotype not located. Syn. with Tenebrio dispar Herbst, 1797: 248 (India) by Carter 1922: 72. Menearchus Carter is endemic to India and Sri Lanka (Iwan 2002: 277).
Mimopeus amaroides Pascoe, 1866: 477 (Tenebrioninae-Heleini). Australia [erroneous]. Syntypes not located. Syn. with Mimopeus elongatus (Brême, 1842: 38) (New Guinea, erroneous) by Bates 1873: 474. Mimopeus Pascoe is endemic to New Zealand (Matthews 1993: 1054).
Thesilea cuprina Fairmaire, 1849: 451 (Stenochiinae-Cnodalonini). Wallis Is. Syntypes not located. Specimens from north Queensland given this name by Carter (1913: 102) are misidentified Titaena Erichson Bates (Doyen et al. 1990: 244).
General References
Aalbu, R.L., Triplehorn, C.A., Campbell, J.M., Brown, K.W., Somerby, R.E. & Thomas, D.B. 2002. Tenebrionidae Latreille, 1802. pp. 463-509 in Arnett, R.H., Thomas, M.C., Skelley, P.E. & Frank, J.H. (eds). American Beetles. Polyphaga: Scarabaeoidea through Curculionoidea. Boca Raton, Florida : CRC Press Vol. 2 xiv 861 pp.
Allsopp, P.G. 1979. Identification of false wireworms (Coleoptera: Tenebrionidae) from southern Queensland and northern New South Wales. Journal of the Australian Entomological Society 18: 277-286
Allsopp, P.G. 1986. Identification of an additional false wireworm Cestrinus trivialis Erichson (Coleoptera: Tenebrionidae) from south Queensland. Journal of the Australian Entomological Society 25: 137-140
Bates, F. 1872. Notes on Heteromera, and descriptions of new genera and species No. 1, No. 3. Entomologist's Monthly Magazine 9: 97-99, 149-152
Bates, F. 1874. Descriptions of new genera and species of Heteromera, chiefly from New Zealand and New Caledonia, together with a revision of the genus Hypaulax and description of an allied new genus from Colombia. Annals and Magazine of Natural History 4 13: 16-23; 102-114
Bouchard, P. 2002. Phylogenetic revision of the flightless Australian genus Apterotheca Gebien (Coleoptera: Tenebrionidae: Coelometopinae). Invertebrate Systematics 16: 449-554
Bouchard, P. & Steiner, W.E. 2004. First descriptions of Coelometopini pupae (Coleoptera: Tenebrionidae) from Australia, Southeast Asia and the Pacific Region with comments on phylogenetic relationships and anti-predator adaptations. Systematic Entomology 29: 101-114
Bouchard, P. & Yeates, D.K. 2001. Phylogenetic relationships of the Australasian Coelometopini (Coleoptera: Tenebrionidae); a quantitative cladistic analysis with a review of biology. Organisms, Diversity and Evolution 1: 17-43
Bremer, H. J 2006. Revision der Gattung Amarygmus Dalman, 1823 sowie verwandter Gattungen. XXXVII. Nachbeschreibungen und Abbildungen australischer Amarygmus-Arten, die von Blackburn beschriebenen wurden. Spixiana (Munich) 1 29: 31-50
Brown, W.V., Doyen, J.T., Moore, B.P. & Lawrence, J.F. 1992. Chemical composition and taxonomic significance of defensive secretions of some Australian Tenebrionidae (Coleoptera). Journal of the Australian Entomological Society 31: 79-89
Carter, H.J. 1913. Revision of Australian species of the subfamilies Cyphaleinae and Cnodaloninae. (Fam. Tenebrionidae). Proceedings of the Linnean Society of New South Wales 38: 61–105, pls VI,VII
Carter, H.J. 1914. Notes on Australian Tenebrionidae, with descriptions of new species. Transactions of the Royal Society of South Australia 38: 369-406
Carter, H.J. 1920. Notes on some Australian Tenebrionidae, with descriptions of new species; — also of a new genus and species of Buprestidae. Proceedings of the Linnean Society of New South Wales 45: 222-249
Carter, H.J. 1922. Australian Coleoptera: Notes and new species. No. ii. Proceedings of the Linnean Society of New South Wales 47(2): 65-82
Carter, H.J. 1930. Check list of the Australian Cistelidae. Order Coleoptera. The Australian Zoologist 6(3): 269–276, pls xviii, xix [20 Aug 1930]
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Doyen, J.T. & Tschinkel, W.R. 1982. Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera). Systematic Entomology 7: 127-183
Erichson, W.F. 1843. Beitrag zur Insecten-Fauna von Angola, in besondere Beziehung zur geographischen Verbreitung der Insecten in Afrika. Archiv für Naturgeschichte 9(1): 199–267
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Gebien, H. 1937. Katalog der Tenebrioniden. Teil I. Pubblicazioni del Museo entomologico “Pietro Rossi” 2: 505-883
Grimm, R. 2013. The genus Cryphaeus Klug, 1833 in Australia, with description of a new species (Coleoptera: Tenebrionidae: Toxicini). Australian Entomologist 40(4): 243-247
Halstead, D.G.H. 1967. A revision of the genus Palorus (sens. lat.) (Coleoptera: Tenebrionidae). Bulletin of the British Museum (Natural History) Entomology 19: 61-148
Iwan, D. 2002. Catalogue of the world Platynotini (Coleoptera: Tenebrionidae). Genus (Wroclaw) 13: 219-323
Kaszab, Z, 1987. Die papuanisch-australischen Arten der Gattung Daerosphaerus Thomson, 1858 (Coleoptera: Tenebrionidae). Acta Zoologica Hungarica 33: 41-85
Kaszab, Z. 1961. Beitrag zur Kenntnis der Tenebrioniden-Tribus Leiochrini (Coleoptera). Annales Historico-Naturales Musei Nationalis Hungarici (Zoologica) 53: 357-380
Kaszab, Z. 1961. Revision der tenebrioniden-Gattung Derispia Lewis (Coleoptera). Acta Zoologica Academiae Scientiarum Hungariae 7: 139-184
Kaszab, Z. 1973. Revision der Arten der Tenebrioniden-Gattung Scotoderus Perroud & Montrouzier (Coleoptera). Folia Entomologica Hungarica 26(2): 257–285
Kaszab, Z. 1977. Die Phrenapatinen des papuanisch-pazifischen Gebietes (Coleoptera: Tenebrionidae). Acta Zoologica Academiae Scientiarum Hungariae 23: 299-339
Kaszab, Z. 1977. Die Tenebrioniden des papuanischen Gebietes. I. Strongyliini (Coleoptera: Tenebrionidae). Pacific Insects Monographs 33: 1-219
Kaszab, Z. 1978. Australische und südpazifische Tenebrioniden der Tribus Phrenapatini und Gnathidiini (Coleoptera) sowie synonymische Bemerkungen. Annales Historico-Naturales Musei Nationalis Hungarici (Zoologica) 70: 163-177
Kaszab, Z. 1982. Die papuanisch-pazifischen Arten der Gattung Lorelus Sharp, 1876 (Coleoptera, Tenebrionidae). Annales Historico-Naturales Musei Nationalis Hungarici (Zoologica) 74: 151-191
Kaszab, Z. 1982. Die Tenebrioniden Neukaledoniens und der Loyauté-Inseln (Coleoptera). Folia Entomologica Hungarica 43: 1-294
Kaszab, Z. 1982. Revision der Australischen Uloma-arten (Coleoptera: Tenebrionidae). Acta Zoologica Academiae Scientiarum Hungariae 28(3-4): 233–291
Kaszab, Z. 1988. Katalog und Bestimmungstabelle der Gattung Promethis Pascoe, 1869 (Coleoptera, Tenebrionidae). Acta Zoologica Hungarica 34(2-3): 67–170
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Matthews, E.G. 1992. Classification, relationships and distribution of the genera of Cyphaleini (Coleoptera: Tenebrionidae). Invertebrate Taxonomy 6: 437-522
Matthews, E.G. 1993. Classification, relationships and distribution of the genera of Heleini (Coleoptera: Tenebrionidae). Invertebrate Taxonomy 7: 1025-1095
Matthews, E.G. 1998. Classification, phylogeny and biogeography of the genera of Adeliini (Coleoptera: Tenebrionidae). Invertebrate Taxonomy 12: 685-824
Matthews, E.G. 2000. Origins of Australian arid-zone tenebrionid beetles. Invertebrate Taxonomy 14: 941-951
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Medvedev, G.S. & Lawrence, J.F. 1987. Tenebrionid beetles of the tribe Hyocini (Coleoptera, Tenebrionidae) of Australia. IV. A key to genera and subgenera. [In Russian]. Entomologicheskoe Obozrenie (English translation as Entomological Reviews) 66: 803-816
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History of changes
| Published | As part of group | Action Date | Action Type | Compiler(s) |
|---|---|---|---|---|
| 01-Jul-2020 | TENEBRIONOIDEA Latreille, 1802 | 28-Nov-2018 | MODIFIED | |
| 12-May-2014 | MODIFIED |