- Crowsoniella relicta
- Micromalthus debilis
- Anthocoleus habiensis (incertae sedis)
- Bothynophora elegans (incertae sedis)
- Chalepocarabus elongatus (incertae sedis)
- Clathrocupes anthrilegnotus (incertae sedis)
- Clinomerus laticellus (incertae sedis)
- Cupedites minutissimus (incertae sedis)
- Doggeriopsis stonefieldiana (incertae sedis)
- Ensicupes guyanensis (incertae sedis)
- Euteticoleus radiatus (incertae sedis)
- Gansucupes attenuatus (incertae sedis)
- Hebeicoleus sertulatus (incertae sedis)
- Helopides hildsiensis (incertae sedis)
- Ironicus nothus (incertae sedis)
- Kakoselia anglia (incertae sedis)
- Katapontisus brodiei (incertae sedis)
- Kelidus bolbus (incertae sedis)
- Liassocupes (incertae sedis)
- Mesothoris (incertae sedis)
- Nannocurculionites carlsoni (incertae sedis)
- Parabuprestites rugulosus (incertae sedis)
- Paracurculionites parvulus (incertae sedis)
- Paradoggeria acuminata (incertae sedis)
- Pseudosilphites triassicus (incertae sedis)
- Sinocupes (incertae sedis)
- Stigmenamma heeri (incertae sedis)
- Synodus changnaensis (incertae sedis)
- Tenthyridium (incertae sedis)
- Tripocoleus rambulosus (incertae sedis)
- Zigadenia tuberculata (incertae sedis)
With 42 extant species the Archostemata are the smallest of the currently recognised four major subgroups of Coleoptera. Representatives of Archostemata occur on all continents with the exception of Antarctica. However, the distribution is quite heterogeneous, with the east Asiatic & Australian region harbouring most species (22) and Europe having only one extant species. During the Mesozoic the group was much more diverse, which is documented by several hundred fossil species. Most of them have been found in European and Siberian fossil sites (Ponomarenko 1969). Recently, many new fossils of Cretaceous and Jurassic age have been described from Chinese localities (Tan & Ren 2009).
Even the oldest known fossils of Coleoptera, like the lower Permian Tshekardocoleus magnus Rhodendorf, 1944, are extremely similar to extant Archostemata in all recognisable characters. However, Tshekardocoleus and other Permian beetles are not members of the extant Archostemata but either represent early side-branches of extant Coleoptera or are part of their stem-lineage. Members of the extant monophylum appear in the upper Permian for the first time (Ponomarenko 1969, Beutel 1997, Beutel et al. 2008).
The name Archostemata was introduced by Kolbe (1908) who also established the taxon as a suborder of Coleoptera. At this time the only known family was Cupedidae. Subsequently, Forbes (1922, 1926) summarised the then known genera Cupes, Omma and Micromalthus under the name Archostemata (Atkins 1958). Today the Archostemata comprise the Cupedidae (33 species) and Ommatidae (6 species), and Micromalthus debilis LeConte, 1878, Crowsoniella relicta Pace, 1975 and probably Sikhotealinia zhiltzovae Lafer, 1996 (Atkins 1963; Lawrence & Newton 1995; Hörnschemeyer 1998; Beutel & Haas 2000; Beutel & Hörnschemeyer 2002a, Beutel et al. 2008, Hörnschemeyer 2009). The phylogenetic relationships of the latter three species are still uncertain (Hörnschemeyer 2009, Ge et al. 2010).
The relationships between Archostemata and the other three groups of Coleoptera (Myxophaga, Adephaga, Polyphaga) have long been uncertain and are still under discussion to some degree (Atkins 1958, 1963, Kukalová-Peck & Lawrence 1993, Hörnschemeyer 1998, Beutel & Haas 2000, Caterino et al. 2002, Leschen & Beutel 2004, Hughes et al. 2006). Presently, a position as sistergroup of all other Coleoptera seems to be best supported (Beutel & Haas 2000, Hughes et al. 2006, Friedrich et al. 2009).
Archostemata comprise species from 1.7 mm (Crowsoniella relicta, Micromalthus debilis) to more than 20 mm (Omma stanleyi, Priacma serrata, Rhipsideigma raffrayi) body length. Characteristic features common to all these species can be found in the larval as well as the adult morphology.
According to Friedrich et al. (2009) and Beutel at al. (2008) Larvae are characterized by the combination of a head with an endocarina and posteromedian emargination, mandibles with three blunt apical teeth, narrow, fused mentum and submentum, an enlarged, wedge-shaped ligula and an abdomen with tergal ampullae in the second and later instars. These characters are present in all known archostematan larvae and most likely represent autapomorphies of the group.
In the adults there are some features that, in the extant fauna, are only found in Archostemata, but that probably are plesiomorphic characters that were already present in the last common ancestor of all Coleoptera. In this category fall, e.g. the typical window puncture of the elytra, which is most conspicuous in Cupedidae, as well as the venation of the archostematan hindwings and there type of folding, or more precisely, rolling of the apical part of the hindwing. Also the tuperculous structure of the cuticle and the scales that cover most or all of the cuticle in Cupedidae are characteristic but not apomorphic. The scales are also present in Ommatidae, but smaller or modified and they are missing in Micromalthus debilis and Crowsoniella relicta.
Autapomorphies of adult Archostemata might be the posteriorly consticted and extended head (missing in Micromalthus), the strongly reduced anterior tentorial arms, which are missing entirely in some species, the mouthparts with an enlarged, lid-like prementum, an absent mentum and modified associated muscles as well as mesocoxal cavities that are bordered by the metepisterna and a distinct median ridge on the first ventrite of the abdomen (Friedrich et al. 2009, Beutel et al. 2008, Hörnschemeyer 2005, 2009).
Phylogenetic relationships of extant and fossil genera of Archostemata and early Coleoptera were analysed by Beutel et al. (2008). Previous analyses of phylogenetic relationships of at least some of the archostematan genera include Beutel & Hörnschemeyer (2002a, b), Hörnschemeyer et al. (2002), Hörnschemeyer et al. (2006). Hörnschemeyer (2009) investigated the phylogeny of Archostemata on species level with 37 of 42 extant species taken into account. In all analyses the monophyly of Ommatidae and of Cupedidae is confirmed and the positions of Micromalthus debilis and Crowsoniella relicta are most uncertain. Originally, C. relicta was placed with Ommatidae by Crowson (1975) and M. debilis somtimes is associated with Cupedidae. More often these two small and highly modified species appear together (as sistergroups) in various positions among the other Archostemata, as e.g. in the analyses by Hörnschemeyer (2009). These results suggest that many characters used in the analyses developed convergently in similar ways in these two species. However, characters of the male genitalia might indicate that contrary to previous assumptions, C. relicta is closely related to Cupedidae and M. debilis a member of Ommatidae, probably closely related to Omma (Hörnschemeyer 2009).
An interesting result of the analysis of Hörnschemeyer (2009) is that Tenomerga, as defined by Neboiss (1984) most likely is paraphyletic with the African Tenomerga leucophaea forming a monophyletic grouping together with the North American Cupes capitatus and African/Madagascan Rhipsideigma.
Despite of all these analyses the phylogenetic relationships of archostematan beetles are still not satifactorily understood. Especially the positions of M. debilis, C. relicta and the enigmatic Sikhotealinia zhiltzovae Lsafer, 1996 need further investigation. Perhaps an analysis based on genetic data might bring some clarification.
|The tree represents the relationships of Archostemata as reconstructed by Hörnschemeyer (2009).||The tree represents the relationships of Archostemata and early fossil beetles as reconstructed by Beutel et al. (2008).|
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