1. ARCHOSTEMATA KOLBE, 1908
John F. Lawrence and Hermes E. Escalona
The genera Cupes Latreille and Omma Newman were included in the suborder Adephaga by Ganglbauer (1903), Kolbe (1901, 1903) and Lameere (1903), based primarily on wing venation, while Kolbe (1908) excluded cupedids from Adephaga and placed them in a separate group, Archostemata, within what is now known as Polyphaga. Forbes (1926) considered Archostemata to be one of the three suborders of beetles, containing Cupedidae Laporte (including Ommatidae Sharp & Muir) and Micromalthidae Barber, based on wing folding, while by Böving & Craighead (1931) and Peyerimhoff (1933) supported this based on larval evidence. This classification was used in Crowsonâs (1955) classic work on the order. In a later paper, Crowson (1962) made the first attempt to integrate the work of paleontologists, such as A. Handlirsch, R. J. Tillyard, B. B. Rohdendorf and A. B. Martynov, on cupedid-like fossils with knowledge of the extant archostematan fauna known at that time. Ponomarenko (1964, 1966, 1968, 1969) published the first major work dealing with Archostemata, which was defined in a broad sense to include virtually all known Permian and Mesozoic fossils sufficiently complete to be described (11 families, 74 genera and 186 species) but lacking the distinctive features of either Polyphaga or Adephaga. The Lower Permian Tshekardocoleidae and several Upper Permian groups were placed in Protocoleoptera and Archecoleoptera, respectively, by Crowson (1975), but were considered to form a coleopterid stem group by KukalovĂĄ-Peck (1991). This was followed by Beutel (1997) and Beutel & Friedrich (2008), who added the Triassic Triadocupedidae to this paraphyletic stem group. Within the remaining Archostemata, relationships are still not clearly understood, particularly among the taxa with non-clathrate elytra (without window-punctures) such as Jurodidae, Ademosynidae, Schizophoridae and Catiniidae, and their possible affinities with the other three suborders, Adephaga, Myxophaga and Polyphaga. Numerous publications have appeared in the past 25 years on fossil Archostemata primarily from localities in Kazakhstan, Russia, Mongolia and northern China but also on those in the UK, USA, Spain, Australia and Argentina; included are Ashman et al. (2015); Jarzembowski & Wang (2016); Jarzembowski et al. (2012, 2013a, 2013b, 2015); Kirejtshuk (2005); Kirejtshuk et al. (2010a, 2010b, 2016); Lubkin (2003); Martin (2010); Martins-Neto et al. (2006); Ponomarenko (1994, 1997, 2000, 2004, 2006); Ponomarenko & Martinez-DelclĂČs (2000); Ponomarenko & Ren (2010); Ponomarenko et al. (2014); Ren & Tan (2006); Ren et al. (2006); Soriano & DelclĂČs (2006); Tan & Ren (2006a, 2006b, 2006c, 2007, 2009); Tan et al. (2007a); Tan et al. (2005); Tan et al. (2006a, 2006b, 2007b, 2007c); Tan et al. (2006c, 2013, 2012) and Yan et al. (2014).
In the study of Archostemata by Kirejtshuk et al. (2016), virtually all archostematans with clathrate or reticulate elytra are considered to belong to the single family Cupedidae, extending back to the Triassic, but without an adequate phylogenetic analysis and additional support from other members of the paleontological community, we continue to recognise the families Ommatidae and Cupedidae. Recent discoveries of Cupedidae in the Cretaceous amber of Myanmar may be found in the papers by Jarzembowski et al. (2017a, 2017b, 2017c).
Some important characters defining this suborder may be found in extant members of the Ommatidae and Cupedidae, but not in highly derived or little known groups like Micromalthidae (Beutel & Hörnschemeyer 2002a), Crowsoniellidae (Ge et al. 2011) or the doubtfully included Jurodidae (Yan et al. 2014). These include: (1) lack of cervical sclerites (also in Adephaga and Myxophaga); (2) external propleuron extending well in front of procoxal cavities (also in Adephaga, reduced in Myxophaga); (3) free, external protrochantin; (4) membranous joint between mesothorax and metathorax (also in some Polyphaga); (5) mesocoxal cavities broadly closed laterally by metanepisternum (narrowly so in a few Polyphaga); (6) well developed and exposed metatrochantins; (7) hind wing without a radial cell formed by meeting of RA1+2 and RA3+4 (also in Adephaga and Myxophaga); (8) hind wing with cross-veins between RP and MP1+2 forming an oblongum cell (also in Adephaga and Myxophaga), (9) hind wing with major fold crossing MP1+2 forming a sharp hinge (also in Adephaga and Myxophaga); (10) larva with 6-segmented legs and paired pretarsal claws (also in Adephaga); (11) larva with ligular sclerome.
There are relatively few extant species of Archostemata: the family Ommatidae contains four Australian species of Omma Newman and two species of Tetraphalerus Waterhouse, occurring primarily in Argentina and Bolivia, and the Cupedidae includes nine genera and 31 species occurring in North America, South America, the eastern Palaearctric and Oriental regions, the East Indies, New Guinea, Australia, New Caledonia, East Africa, South Africa and Madagascar (Neboiss 1984; Lawrence 1999; Hörnschemeyer 2009, 2016; Hörnschemeyer & Beutel 2016; Hörnschemeyer & Yavorskaya 2016). The group has become extinct in Europe, but several European species are known from the early Tertiary (Kirejtshuk 2005; Kirejtshuk et al. 2010a). Several morphological papers have been produced on both larvae (Beutel & Hörnschemeyer 2002a, 2002b; Hörnschemeyer et al. 2002; Grebennikov 2004; Yavorskaya et al. 2015) and adults (Baehr 1975; Hörnschemeyer et al. 2006; Beutel et al. 2008; Friedrich et al. 2009; HĂŒnefeld et al. 2011). General summaries of extant Archostemata may be found in Hörnschemeyer (2016), Hörnschemeyer & Beutel (2016) and Hörnschemeyer & Yavorskaya (2016).
References
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Hörnschemeyer T (2016) 5. Archostemata Kolbe, 1908, 5.1. Crowsoniellidae Iablokoff-Khnzorian, 1983. 5.3. Micromalthidae Barber, 1913. 5.5 Jurodidae Kirejtshuk, 1999 (= Sikhotealinidae). In Handbuch der Zoologie/Handbook of Zoology. Band/Volume IV Arthropoda: Insecta. Teilband/Part 38. Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2nd edn. (Eds RG Beutel and RAB Leschen) pp. 41â43, 48â51, 57â62. Walter de Gruyter, Berlin.
Hörnschemeyer T, Beutel RG (2016) 5.4. Ommatidae Sharp & Muir, 1912. In Handbuch der Zoologie/Handbook of Zoology. Band/Volume IV Arthropoda: Insecta. Teilband/P...