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Taxonomy of the Reticulate Beetles of the Subfamily Cupedinae (Coleoptera: Archostemata), with a Review of the Historical Development A.G

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Taxonomy of the Reticulate Beetles of the Subfamily Cupedinae (Coleoptera: Archostemata), with a Review of the Historical Development A.G Invertebrate Zoology, 2016, 13(2): 61–190 © INVERTEBRATE ZOOLOGY, 2016 Taxonomy of the reticulate beetles of the subfamily Cupedinae (Coleoptera: Archostemata), with a review of the historical development A.G. Kirejtshuk1,2, A. Nel2, P.A. Kirejtshuk3 1 Zoological Institute of the Russian Academy of Sciences, Universitetskaya emb. 1, St. Petersburg, 199034, Russia. E-mail: [email protected], [email protected] 2 Institut de Systématique, Évolution, Biodiversité, ISYEB - UMR 7205 – CNRS, MNHN, UPMC, EPHE, Muséum national d’Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, CP 50, Entomologie, F-75005, Paris, France. E-mail: [email protected] 3 6th Linia, 49-34, St. Petersburg, 199004, Russia. E-mail: [email protected] ABSTRACT:The paper is mainly devoted to the studies of the fossil Cupedidae from the Paleocene of Menat (Puy-de-Dôme, France) and to a preliminary generic analysis of the subfamily Cupedinae. All modern and fossil taxa of genus rank which could belong to Cupedinae are revised. The previous division of this subfamily into tribes (including Priacmini and Mesocupidini) is regarded as not advisable. An annotated list of the fossil species of the subfamily Cupedinae is compiled and revised. New keys to subfamilies of Cupedidae and to genera and subgenera of Cupedinae, and also keys for species of Cainomerga subgen.n. and Cupes found in deposits from Menat are given. As a result, several new genera and subgenera are proposed, i.e. the fossil Apriacma gen.n., from the early Cretaceous (the type species: Priacma tuberculosa Tan, Ren et Shin, 2006), Cre- tomerga gen.n., from the early Cretaceous (the type species: Priacmopsis subtilis Tan et Ren, 2006), Cupopsis gen.n. from the late Cretaceous (the type species: Paracupes svetkoi Lubkin, 2003), Cainomerga subgen.n., from the Paleocene (the type species: Mesocupes (Cainomerga) brevicornis sp.n.), Menatops gen.n., from the Paleocene (the type species: Cupes orbiculatus Kirejtshuk, Nel et Collomb, 2010), Taxopsis gen.n., from the late Eocene (the type species: Cupes motschulskyi Kirejtshuk, 2005), Cainocups gen.n., from the latest Oligocene (the type species: Cainocups aixensis sp.n.), as well as the Recent Paracupoides subgen.n., from central Ecuador, South America (the type species: Paracupes ascius Neboiss, 1989). The following species are described from the Paleocene deposits of Menat: Mesocupes (Cainomerga) brevicornis sp.n., M. (C.) fraternus sp.n., M. (C.) palaeocenicus sp.n., M. (C.) ponti sp.n., Menatops bartenevi sp.n., Cupes distinctissimus sp.n., C. simillimus sp.n., or commented: Menatops orbiculatus (Kirejtshuk, Nel et Collomb, 2010), comb.n. transferred from Cupes], and Mesocupes (Cainomerga) immaculatus (Piton, 1940), comb.n. transferred from Zonabris (Meloidae). Besides, some cupedine fossils from Menat remain without both description and generic attribution. Kirghizocupes Ponomaren- ko, 1966 is transferred from the Triadocupedinae to Cupedinae and synonymized with Mesocupoides Ponomarenko, 1969, syn.n., Pulchicupes Ren, 1995 is transfered from the Taldycupedidae to Cupedinae and provisionally synonymized with Latocupes Tan et Ren, 2006, in which also three species described in “Mesocupes” are transfered: L. angustilabi- alis (Tan, Huang et Ren, 2007), comb.n., L. collaris (Tan, Huang et Ren, 2007), comb.n. and L. latilabialis (Tan, Huang et Ren, 2007), comb.n. The genus Mesocupes is divided into the subgenera Mesocupes s.str., Anaglyphites Ponomarenko, 1964, stat.n., and Cainomer- ga subgen.n., with Priacmopsis minimus Ponomarenko, 1997 transferred to the first subgenus [Mesocupes (Mesocupes) minimus, comb.n.]. The genus Apriacma gen.n. is proposed for four fossil species previously described in “Priacma”: A. clavata (Tan, Ren et Shih, 2006), comb.n., (?) A. latidentata (Tan, Ren et Shih, 2006), comb.n., P. renaria 62 A.G. Kirejtshuk et al. (Tan, Ren et Shih, 2006), comb.n., and A. tuberculosa (Tan, Ren et Shih, 2006), comb.n., while the remaining fossils species described in “Priacma” are left without generic allocation: “Priacma” corrupta Ponomarenko, 1986, “P.” longicapitis Ponomarenko, 1997, “P.” oculata Ponomarenko, 1997, “P.” striata Ponomarenko, 2000, and “P.” sanzii Soriano et Delclós 2006. The genera Asimma Ponomarenko, 1966, and Pterocupes Ponomarenko, 1966 are transferred from the subfamily Triadocupedinae to Cupedinae, albeit excluding Pterocupes leptocerus Ponomarenko, 1966, which needs revision to clarify its generic attribution. The genus Ensicupes Hong, 1976 is transferred from the family Taldicupedidae to Cupedidae and provisionally assigned to the subfamily Cupedi- nae. The genera Helopides Roemer, 1876, Longaevicupes Ren, 1995, Lupicupes Ren, 1995, Mesothoris Tillyard, 1916, Nannocurculionites Handlirsch, 1906, Parabuprestites Handlirsch, 1906, Paracurculionites Handlirsch, 1906, and Pseudosilphites Zeuner, 1930 are also regarded as members of Cupedidae without subfamily allocations. The positions of Anthocoleus Hong, 1983, Argentinocupes Martins-Neto et Gallego in Martins-Neto et al., 2006a, Bothynophora Heer, 1865, Celocoleus Hong, 1983, Chengdecupes Hong 1983, Cupedites Ponomarenko, 1985, Doggeriopsis Handlirsch, 1906, Euteticoleus Hong, 1983, Gansucupes Hong, 1982, Hebeicoleus Hong, 1992, Hebeicupes Zhang, 1986, Ironicus Handlirsch, 1906, Katapontisus Handlirsch, 1906, Kelidus Handlirsch, 1906, Liassocupes Zeuner, 1962, Mesotricupes Hong, 1982, Metacupes Gardiner, 1961, Paradoggeria Handlirsch, 1906, Petalocupes Hong, 1982, Stigmenamma Handlirsch, 1906, Synodus Hong, 1982, Tentyridium Westwood, 1854, and Tetrocupes Hong 1983 are also reconsid- ered. Both genera Clathrocupes Hong, 1980, originally described in the Taldycupedidae, and Eumecoleus Haupt, 1950, described in Oedemeridae, are regarded as archostematan genera without family assignment, although the latter seems to belong to cupedids. The genus Lecanopteryx Haupt, 1950 is regarded as a beetle without subordinal placement. The species taxon “punctomarginum Dunstan, 1923” is transferred from the genus Elaterium Westwood, 1854 to Mesothoris [Mesothoris punctomarginum (Dunstan, 1923), comb.n.]. The taxa Adinolepis Neboiss, 1984 and Ascioplaga Neboiss, 1984 are considered as subgenera of the one genus (Adinolepis). The position of the genus Priacma is discussed in special detail. It is characterized by a peculiar (Priacma-type) aedeagus which is unique among the modern Archostemata, but also shared some similarity with the Middle Jurassic genus Gracilicupes Tan, Ren et Shin, 2006. Some of its structures (“volsellar plate”, “digitus volsellaris” and “cuspis volsellaris”, after Edwards, 1953) are known in the structure of the male genitalia of numerous hymenopterans, but not in other holometab- olans, including almost all modern representatives of the order Coleoptera. However, many archostematans and some polyphagans of different groups show probable derivatives of these structures which are traceable and can be used for phylogenetic reconstructions. The apomorphies of the order Coleoptera in general, suborder Archostemata, family Cupedidae and its subfamilies are reviewed. The problem of origin of the order is discussed, overviews of the historical development of the suborder Archostemata and subfamily Cupedinae are given. The following scheme of divergences of the order Coleoptera is proposed: Tshek- ardocoleidea + {Schizophorimorpha + [Cupedinae + (Ommatinae + other families of Cupedomorpha)]}. How to cite this article: Kirejtshuk A.G., Nel A., Kirejtshuk P.A. 2016. Taxonomy of the reticulate beetles of the subfamily Cupedinae (Coleoptera, Archostemata), with a review of the historical development // Invert. Zool. Vol.13. No.2. P.61–190. doi: 10.15298/ invertzool.13.2.01 KEY WORDS: fossil and recent Archostemata, Cupedidae, Cupedinae, classification, phylogeny, new genera and species, synonymies, fossil record, historical development. Taxonomy of the reticulate beetles of the subfamily Cupedinae 63 Таксономия лакомок подсемейства Cupedinae (Coleoptera: Archostemata), с обзором исторического развития А.Г.Кирейчук1, 2, А. Нель2, П.А. Кирейчук3 1 Зоологический институт РАН, Университетская наб. 1, Санкт-Петербург, 199034, Рос- сия. E-mail: [email protected], [email protected] 2 Институт систематики, эволюции и биоразнообразия, ISYEB - UMR 7205 – CNRS, MNHN, UPMC, EPHE, Национальный музей естественной истории, Университеты Сорбоны, ул. Кювье, 57, CP 50, Энтомология, F-75005, Париж, Франция. E-mail: [email protected] 3 6-я линия, 49-34, Санкт-Петербург, 199004, Россия. E-mail: [email protected] РЕЗЮМЕ: Настоящая работа посвящена преимущественно изучению ископаемых купедид (Cupedidae) из палеоценового захоронения Мена (Пюи-де-Ром, Франция) и предварительному анализу родов подсемейства Cupedinae. Все современные и вы- мершие таксоны, которые могли относиться к Cupedinae, ревизованы. Предшеству- ющее деление подсемейства на трибы (включая Priacmini и Mesocupidini) рассматри- вается как нецелесообразное. Составлен и ревизован аннотированный список иско- паемых видов подсемйства Cupedinae. Приводится новые определительные таблицы для подсемейств Cupedidae, а также для родов и подродов Cupedinae, а также определительные таблицы для видов Cainomerga subgen.n. и Cupes, найденныех в захоронении Мена. В результате предложено несколько новых родов и подродов, а именно: вымершие Apriacma gen.n. из раннего мела (типовой вид: Priacma tuberculosa Tan, Ren et Shin, 2006), Cretomerga gen.n. из раннего мела (типовой вид: Priacmopsis subtilis Tan et Ren, 2006), Cupopsis gen.n. из позднего мела (типовой вид: Paracupes
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