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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: agk@mail.ru, Kirejtshuk@gmail.com
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: anel@mnhn.fr
3 6th Linia, 49-34, St. Petersburg, 199004, Russia. E-mail: polly.k@mail.ru
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: agk@mail.ru, Kirejtshuk@gmail.com
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: anel@mnhn.fr
3 6-я линия, 49-34, Санкт-Петербург, 199004, Россия. E-mail: polly.k@mail.ru
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: Priacmatuberculosa
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
svetkoi Lubkin, 2003), Cainomerga subgen.n. из палеоцена [типовой вид: Mesocupes
(Cainomerga) brevicornis sp.n.], Menatops gen.n. из палеоцена (типовой вид: Cupes
orbiculatus Kirejtshuk, Nel et Collomb, 2010), Taxopsis gen.n. из верхнего эоцена
(типовой вид: Cupes motschulskyi Kirejtshuk, 2005), Cainocups gen.n. из позднего
олигоцена (типовой вид: Cainocups aixensis sp.n.), а также современный Paracupoides
subgen.n. из Центрального Эквадора Южной Америки (типовой вид: Paracupes
ascius Neboiss, 1989). Из палеоценового захоронения Мена описаны следующие
виды: 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., а также рассмотрены: Menatops orbiculatus (Kirejtshuk, Nel et
Collomb, 2010), comb.n. перенесенный из рода Cupes, и Mesocupes (Cainomerga)
immaculatus (Piton, 1940), comb.n., перенесенный из Zonabris (Meloidae). Вместе с
тем некоторые остатки купедин остались неописанными и без определения родовой
принадлежности. Род Kirghizocupes Ponomarenko, 1966 перенесен из Triadocupedinae
в Cupedinae и синонимизирован с Mesocupoides Ponomarenko, 1969, syn.n., Pulchicupes
Ren, 1995 перенесен из Taldycupedidae в Cupedinae и предварительно синонимизиро-
ван с Latocupes Tan et Ren, 2006, в который перенесены также виды, первоначально
описанные в составе “Mesocupes”: L. angustilabialis (Tan, Huang et Ren, 2007),
comb.n., L. collaris (Tan, Huang et Ren, 2007), comb.n. и L. latilabialis (Tan, Huang et
Ren, 2007), comb.n. Род Mesocupes разделен на три подрода: Mesocupes s.str.,
Anaglyphites Ponomarenko, 1964, stat.n. и Cainomerga subgen.n., с перенесением и
64
A.G. Kirejtshuk et al.
Priacmopsis minimus Ponomarenko, 1997 в первый подрод (Mesocupes (Mesocupes)
minimus, comb.n.). Род Apriacma gen.n. предложен для четырех видов, прежде опи-
санных в составе “Priacma”: A. clavata (Tan, Ren et Shih, 2006), comb.n., (?) A.
latidentata (Tan, Ren et Shih, 2006), comb.n., A. renaria (Tan, Ren et Shih, 2006), comb.n.,
and P. tuberculosa (Tan, Ren et Shih, 2006), comb.n., в то время как остальные
ископаемые виды, описаные в составе рода “Priacma”, остаются без родовой принад-
лежности: “Priacma” corrupta Ponomarenko, 1986, “P.” longicapitis Ponomarenko,
1997, “P.” oculata Ponomarenko, 1997, “P.” striata Ponomarenko, 2000, and “P.” sanzii
Soriano et Delclуs 2006. Роды Asimma Ponomarenko, 1966 и Pterocupes Ponomarenko,
1966 переведены из подсемейства Triadocupedinae в Cupedinae, однако исключая
Pterocupes leptocerus Ponomarenko, 1966, который нуждается в переизучении для
выяснения родовой принадлежности. Род Ensicupes Hong, 1976 переведен из семей-
ства Taldicupedidae в Cupedidae и предварительно отнесен к подсемейству Cupedinae.
Роды Helopides Roemer, 1876, Longaevicupes Ren, 1995, Lupicupes Ren, 1995, Mesothoris
Tillyard, 1916, Nannocurculionites Handlirsch, 1906, Parabuprestites Handlirsch, 1906,
Paracurculionites Handlirsch, 1906 и Pseudosilphites Zeuner, 1930 также рассматрива-
ются в составе Cupedidae, но без отнесения к какому-либо подсемейству. Положение
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 и Tetrocupes Hong, 1983 также пересмотре-
но. Оба рода Clathrocupes Hong, 1980, первоначально описанный в Taldycupedidae,
и Eumecoleus Haupt, 1950, описанный в Oedemeridae, рассматриваются как архосте-
маты без семейственной принадлежности, хотя последний скорее всего относится к
купедидам. Род Lecanopteryx Haupt, 1950 расстатривается как жук, но без отнесения
к определенному подотряду. Видовой таксон “punctomarginum Dunstan, 1923” пере-
несен из рода Elaterium Westwood, 1854 в Mesothoris [Mesothoris punctomarginum
(Dunstan, 1923), comb.n.]. Таксоны Adinolepis Neboiss, 1984 и Ascioplaga Neboiss,
1984 рассматриваются как подроды одного рода (Adinolepis). Подробно обсуждается
положение рода Priacma. Он характеризуется особым строением эдеагуса, которое
уникально среди современных архостемат, но сходный эдеагус отмечен у среднеюр-
ского рода Gracilicupes Tan, Ren et Shin, 2006. Некоторые структуры [“volsellar plate”,
“digitus volsellaris” и “cuspis volsellaris” согласно Эдварсу (Edwards, 1953)] известны
в строении генителий самцов многих перепончатокрылых, но не у других голомета-
бол, включая почти всех остальных современных представителей отряда Coleoptera.
Однако многие архостематы и некоторые полифаги из различных групп обнаружи-
вают предположительные производные этих структур, которые можно проследить и
использовать для филогенетических реконструкций. Пересмотрены апоморфии от-
ряда Coleoptera в целом, подотряда Archostemata, семейства Cupedidae и его подсе-
мейств. Обсуждается проблема происхождения отряда. Приводятся обзоры истори-
ческого развития подотряда Archostemata и подсемейства Cupedinae. Предложена
следующая схема дивергенции в отряде Coleoptera: Tshekardocoleidea + {Schizo-
phorimorpha + [Cupedinae + (Ommatinae + other families of Cupedomorpha)]}.
Как цитировать эту статью: Kirejtshuk A.G., Nel A., Kirejtshuk P.A. 2016. Taxonomy of
the reticulate beetles of the subfamily Cupedinae (Coleoptera, Archostemata), with a review
Taxonomy of the reticulate beetles of the subfamily Cupedinae
65
of the historical development // Invert. Zool. Vol.13. No.2. P.61–190. doi: 10.15298/
invertzool.13.2.01
КЛЮЧЕВЫЕ СЛОВА: современные и вымершие Archostemata, Cupedidae, Cupedinae,
классификация, филогения, новые роды и виды, синонимия, палеонтологическая
летопись, историческое развитие.
Contents
Introduction........................................................ 65
Repositories ........................................................ 66
Materials and methods ..................................... 66
Abbreviations for elytral veins ......................... 67
Systematic part .................................................. 67
1. Family Cupedidae Laporte, 1836 ................... 67
2. Key to the subfamilies of Cupedidae ............. 71
3. Subfamily Cupedinae Laporte, 1836 ............. 72
4. Key to fossil and recent genera and
subgenera of Cupedinae ............................... 74
5. Subfamily Cupedinae from Menat ................. 78
5.1. Geological setting and locality
information ................................................. 78
5.2. Genus Mesocupes Martynov, 1926 .......... 78
5.3. Key to species of Cainomerga
subgen.n. from Menat ................................ 98
5.4. (?) Mesocupes (Cainomerga) spp. ........... 99
5.5. Menatops gen.n. ...................................... 100
5.6. Genus Cupes Fabricius, 1801 ................. 103
5.7. Key to species of the genus Cupes from
Menat ........................................................ 114
5.8. Genus incertus sp.1 ................................. 114
5.9. Genus incertus sp.2 ................................. 114
5.10. Genus incertus sp.3 ............................... 114
5.11. Genus incertus sp.4 ............................... 115
6. New genera and species of the subfamily
Cupedinae .................................................... 115
6.1. Apriacma gen.n. ...................................... 115
6.2. Cretomerga gen.n.................................... 119
6.3. Cupopsis gen.n. ....................................... 122
6.4. Genus Paracupes. Paracupoides
subgen.n. .............................................. 123
6.5. Taxopsis gen.n. ........................................ 128
7. Subfamily incertae sedis ............................... 129
7.1. Cainocups gen.n. ..................................... 129
8. Lists of fossil taxa re-examined.................... 130
8.1. List of fossil genera and subgenera
of Cupedinae............................................. 130
8.2. List of fossil cupedine genera incertae
sedis
(with indefinite position) ......................... 139
8.3. List of fossil species of Cupedinae
without generic attribution....................... 143
8.4. List of fossil cupedid genera incertae sedis
(uncertain or corrected subfamily), not
Cupedinae or doubtful members of
Archostemata ............................................ 144
8.5. List of fossil species without generic
attribution from Cupedidae ..................... 151
9. Position of the genus Priacma LeConte,
1861 ........................................................... 152
10. Taxonomic rank of Adinolepis Neboiss,
1984 and Ascioplaga Neboiss, 1984 ......... 165
Discussion ......................................................... 165
Historical overview of studies on
Archostemata ............................................ 165
Phylogeny and historical development of the
suborder Archostemata ............................ 169
Phylogeny and historical development of the
subfamily Cupedinae ............................... 173
Conclusions ...................................................... 179
Acknowledgements .......................................... 180
References......................................................... 181
Appendix 1. Materials studied ...................... 187
Introduction
The suborder Archostemata, the oldest group
of living Coleoptera, might have appeared be-
fore the Permian (Kirejtshuk et al., 2013). It is
currently considered as the sole subgroup of this
order which occurred in the Palaeozoic. Retic-
ulate beetles were quite numerous during the
first half of the Mesozoic (Ponomarenko, 2003,
2013). Since the Palaeozoic to Mesozoic bound-
ary, members of the other suborders, Polyphaga
and Adephaga, appeared (Volkov, 2012) and
became even more diverse. Their proportions in
the late Mesozoic and Cenozoic deposits in-
creased progressively while archostematans
gradually grew less numerous and less diverse
(Kirejtshuk, Ponomarenko, 2016). In the Ceno-
zoic outcrops, archostematans are represented
by few specimens and species while many Palae-
ozoic groups of this suborder disappeared com-
pletely. However, this ancient group is still
comparatively well represented in some Ceno-
zoic palaeobiotas. This is the case for the Eocene
Baltic amber, as well as for some outcrops with
compression deposits (Menat, Middle Pale-
66
A.G. Kirejtshuk et al.
ocene; Messel, Middle Eocene; Eckfeld, Mid-
dle Eocene; Bembridge Marls, Eocene/Oli-
gocene; and Pochlovice, Middle Miocene). Few
Archostemata are also recorded from Aix-en-
Provence (latest Oligocene), Oise amber (earli-
est Eocene), Dominican amber (early Miocene),
and Chiapas amber (Oligocene/Miocene)
(Iablokoff-Khnzorian, 1960; Ponomarenko,
1973; Gersdorf, 1976; Tröster, 1993; Kirejt-
shuk, 2005, 2016; Perkovsky, 2007; Kirejtshuk
et al., 2010; in press; etc.). All Cenozoic archo-
stematan fossils belong to the cupedid subfam-
ilies Cupedinae Laporte, 1836 and Ommatinae
Sharp et Muir, 1912, as well as to the family
Micromalthidae Barber, 1913. In the Recent
fauna, this suborder is represented by few spe-
cies in four families: Cupedidae, Jurodidae Pono-
marenko, 1985, Micromalthidae, and Crowso-
niellidae Iablokoff-Khnzorian, 1983, sporadi-
cally scattered all over the areas with nemoral
and subtropical climates.
The family Cupedidae has always been re-
garded as keystone for understanding the evolu-
tion of the order Coleoptera in general. A very
important step forward was the monograph on
the historical development of the suborder Ar-
chostemata, by Ponomarenko (1969). Many new
important data on the evolution of the family
have since been obtained, but most of the
phylogenetic hypotheses published during the
last few decades are mainly based on informa-
tion derived from modern members of Cupe-
didae (e.g. Vulcano, Pereira, 1975; Neboiss,
1984; Hörnschemeyer, 2009; etc.). The close
Recent relatives of Cupes Fabricius, 1801 are
only seldom included in the family Cupedidae,
but rather considered separately in Ommatidae
(the modern genera Omma Newman, 1839, and
Tetraphalerus Waterhouse, 1901), without tak-
ing into account that the Mesozoic relatives of
both Omma and Cupes showed rather mosaic
distributions of the structural features which are
now present only in one or another group (see
discussion below). Studying the cupedine mate-
rial from the Paleocene of Menat, the authors of
this paper came to the conclusion that some of
the examined specimens are more similar to
Mesozoic relatives while others are quite char-
acteristic of Recent archostematan genera and
also present in the early Mesozoic. To trace the
relations between these insects, the authors felt
obliged to undertake a preliminary revision of
the subfamily which, in turn, provided an oppor-
tunity to propose an essay on the historical
development of the subfamily.
After the paper had been submitted for pub-
lication, some additional important materials
were revealed and some useful recommenda-
tions were obtained. This forced the authors to
introduce certain corrections to the composition
of the manuscript and also to add some new
parts. As a result, most illustrations remain
numbered in the initial order, some more plates
being added to the end.
Repositories
BMNH — Natural History Museum in London
(former British Museum of Natural History)
CNU — Capital Normal University, Bejing,
China
CUPC — Cornell University Paleobotanical
Collection, Ithaca, USA
DEIM — Deutsches Entomologisches Institut,
Müncheberg, Germany
GPIM — Geologisch-Palaeontologisches Insti-
tut und Museum, Universität Hamburg, Germany
GZUG — Geowissenschaftlishes Zentrum, Uni-
versität Göttingen, Germany
LNNR — Naturhistorisches Museum, Landes-
sammlung für NatuLandessammlung für Naturkunde
Rheinland-Pfalz, Mainz, Germany
MNHN — Muséum National d’Histoire Na-
turelle, Paris, France
MTM — Menat Town Museum, Puy-de-Dôme,
France
PIN — Palaeontological Institute of Russian
Academy of Sciences, Moscow, Russia
SFNFM — Senckenberg Forschungsinstitut und
Naturmuseum Forschungsstation Grube Messel,
Germany
ZIN — Zoological Institute of Russian Academy
of Sciences, St. Petersburg, Russia
Material and methods
Besides the material from Menat, deposited
in MNHN and in the Menat Town Museum
(Village of Menat, Puy-de-Dôme), the authors
Taxonomy of the reticulate beetles of the subfamily Cupedinae
67
studied many other archostematans from the
Cenozoic stored in various collections, includ-
ing fossil cupedids from Eckfeld (Middle
Eocene), Messel (Middle Eocene), Baltic am-
ber (late Eocene), Oise amber (earliest Eocene),
Aix-en-Provence (latest Oligocene), Bembridge
(Eocene to Oligocene boundary), as well as
Mesozoic archostematans from BMNH, CNU,
MNHN, PIN, and Staatliches Museum für
Naturkunde in Stuttgart. Comparisons with Re-
cent species were based on specimens from
BMNH, MNHN, and ZIN. The species exam-
ined by the authors during the preparation of this
paper are referred to in the descriptions and in
the species list below (see Appendix 1). The
holotypes of all new species here described are
deposited in the collection of MNHN and the
holotypes described in Kirejtshuk et al. (2010) —
in the collection of MTM.
The specimens were studied using an Olym-
pus SCX9 stereomicroscope with an Olympus
camera and a Nikon SMZ1500 with Micro-
scope Eyepiece Camera 9.0MP Aptina Color
CMOS MU900, at MNHN; a Leica MZ 12.0
stereomicroscope with a DFC290 camera, at
ZIN; a Leica MZ 9.0 stereomicroscope with a
DFC290 camera, at PIN; a Leica MZ 12.5
stereomicroscope with a Nikon Digital Camera
DXM1200C, at CNU. Some specimens were
also examined with a Tescan Vega LSU scan-
ning electron microscope, at MNHN (Low Vac-
uum Secondary Electron TESCAN Detector,
LVSTD), to test the characters not readily visi-
ble with optic equipment (punctation, margins
of legs, shape of large sclerites, scales on elytra).
The pictures of specimens from SFNFM and
LNNR were taken by Sonja Wedmann (SFN-
FM) and Herbert Lutz (LNNR); those from
CUPC were taken from Lubkin (2003).
Abbreviations for elytral veins
A1, A2, A3, A4 — anal veins;
CuA — cubitus anterior;
CuP — cubitus posterior;
M — median vein;
R — radial vein;
RA — anterior branch of radial vein;
RP — posterior branch of radial vein;
Sc — subcostal vein.
Systematic part
Order Coleoptera Linnaeus, 1758
Suborder Archostemata Kolbe, 1908
1. Family Cupedidae Laporte, 1836
Cupesidae Laporte, 1836; type genus: Cupes
Fabricius, 1801;
= Brochocoleidae Hong, 1982; type genus: Brochoc-
oleus Hong, 1982; currently considered as a tribe in the
subfamily Ommatinae;
= Ommadidae Sharp et Muir, 1912; type genus: Omma
Newman, 1839; currently considered as a subfamily (Om-
matinae) in the family Cupedidae.
This family is well recognizable both in the
modern fauna and as fossils due to the character-
istic elytral structure consisting of two rows of
cells between not more than five long primary
veins. Their configuration is peculiar, including
frequent fusions of long anal vein (A1) and CuA
at the elytral apices, a squamose vestiture of the
dorsum in most of the Recent and many fossil
species, a more or less tuberculose dorsal sur-
face of the head and pronotum, oval procoxal
cavities broadly open externally, triangular
metacoxae projecting medially, rather narrow
and long legs. Having appeared in the fossil
record in the Middle/late Triassic, the family
Cupedidae is found to have occurred through
the Mesozoic and Cenozoic, although Triado-
cupedinae Ponomarenko, 1966, spanned only
during the second half of the Triassic (Kirej-
tshuk, Ponomarenko, 2016). There are more
than 30 known species in the extant fauna dis-
tributed across many areas, but outside Europe
(and the Western Palaearctics in general) most-
ly in South East Asia. Larvae of the extant
species are known to develop in dead wood of
angiosperm trees only. Adults occur on the
surface of tree trunks and branches, frequently
visiting flowers. A comparatively great number
of Cenozoic fossils in this family indicates that
this group must have strongly declined even
since the Paleogene. Fossil Cupedinae s.str.
have been recorded from the Miocene (Pono-
marenko, 1973) and even reached the Pliocene
in Europe (Gersdorf, 1976). Two ommatine
genera have been found both in modern fauna
and Mesozoic deposits. Tetraphalerites oligo-
cenicus Crowson, 1962, also from the Eocene-
68
A.G. Kirejtshuk et al.
Oligocene boundary (Isle of Wight, UK), is now
transferred from the tribe Ommatini to Bro-
chocoleini, previously regarded as an exclu-
sively Mesozoic group (only up to the Cenom-
anian) (Ponomarenko, Kirejtshuk, in press). The
authors follow the cupedid classification elabo-
rated by Crowson (1962), Atkins (1963) and
Ponomarenko (1969), further developed by
Ponomarenko (1997, 2006; etc.), Kirejtshuk et
al. (2010a, b) and Jarzembowski et al. (2013a,
b). Some changes to the family’s system were
proposed by Tan and Ren (2009), as well as by
Tan et al. (2012). These works were partly
criticized from the nomenclatural and taxonom-
ic viewpoints by Dubois et al. (2013). The
systematic and phylogenetic proposals of those
two publications (Tan, Ren, 2009; Tan et al.,
2012) are here regarded as ill-founded because
of many defects in interpreting some structures
and in the consistency of applying taxonomic
and evolutionary data. Some of the mistakes
have been corrected in Kirejtshuk et al., 2010b,
being further elaborated in the present paper.
Notes on structural peculiarities
The diagnostic characters used to separate
the cupedids rather frequently show a mosaic
distribution among members of different sub-
families, probably as a result of parallelisms in
structural transformations (Kirejtshuk, 2005;
Kirejtshuk et al., 2010a). The characters related
to the structure of the prothorax with different
positions of the procoxae make it possible to
separate comparatively large groups (subfami-
lies). However, other characters used to dis-
criminate genera, being scattered across all oth-
er subfamilies (sculptural peculiarities of the
elytra, head, pronotum, procoxae with/without
fold (plate), antennae, and the articulation of
abdominal ventrites), can scarcely be used as
traits of a significant evolutionary (phylogenet-
ic) meaning because of their mosaic distribution
among members of the family. Atkins (1958a)
discussed a mosaic distribution of characters in
different modern genera of the family, but the
mosaicism in character patterns is even more
expressed among the fossil Mesozoic groups.
The male genitalia of Recent and some fossil
Cupedinae look rather different from those of
Recent Ommatinae. Sharp & Muir (1912), after
comparative studies of this organ, came to the
conclusion that these groups should be regarded
as separate families. Some neontologists follow
this opinion even now (Lawrence, 1999; Bou-
chard et al., 2011; Lawrence et al., 2011; etc.).
In addition to the difference in structure of the
procoxae, they (e.g. Lawrence, 1999) used as
diagnostic the structure of the abdominal ven-
trites which can be flat or overlapping (the
articulation of the abdominal ventrites abutting
in Ommatinae versus tegular in Cupedinae, al-
though the abdominal ventrites of Priacma Le-
Conte, 1861 have abutting type — see below).
This difference appears rather stable among
Recent taxa, but most fossil ommatines are
characterized by an overlapping articulation
between the ventrites (both Notocupedini Pono-
marenko, 1966, and Lithocupedini Ponomaren-
ko, 1969), while many fossil cupedines (all
Mesocupedini Ponomarenko, 1969) show a flat
(simple) articulation between the ventrites, in-
cluding some species of Cupes from Baltic
amber (Kirejtshuk, 2005) (see also below the
notes on the subfamily Cupedinae). As Moravo-
coleus permianus (Tshekardocoleidae Rohden-
dorf, 1944) demonstrates clearly overlapping
abdominal ventrites, it seems appropriate to
preliminarily suggest that such a type of articu-
lation could be an apomorphy for the Archo-
stemata and Coleoptera in general (see the be-
low Discussion), but certainly plesiomorphic
for cupedids. The abutting type in this case is to
be treated as apomorphic in spite of its simple
structure which is usually interpretated as initial
for the beetles in general (Crowson, 1981). The
flat (abutting) or overlapping (tegular) articula-
tion of abdominal ventrites seem to have no
clear regular relation with membrane between
ventrites. In most Recent species of Cupes the
tegular articulation is combined with more or
less connate edges of ventrites which, however,
makes it possible to bend the abdomen in the
places of articulation while the abutting articu-
lation of ventrites in the Recent Priacmaserrata
LeConte, 1861 is combined with hard fusion
Taxonomy of the reticulate beetles of the subfamily Cupedinae
69
making it impossible to move the abdomen in
the places of articulation more than in other
abdominal portions. The Recent ommatines with
the abutting articulation of abdominal ventrites
frequently demonstrate the connate places of
articulation strengthening by narrow thickened
stripes along the posterior and anterior edges of
each ventrite. If the type of articulation in some
fossils can be defined, the problem of membra-
nous connection or level of fusion of boundaries
of ventrites almost in all fossils, remains unsolv-
able and inapplicable in diagnostics. Taking
into account a similar variability of other struc-
tures of both groups (“Cupedidae” and “Omma-
tidae”), only the structure of the prosternum and
procoxae can indeed be supposed as showing
rather reliable characters for the discrimination
of these groups, viz. Cupedinae have oval and
slightly convex procoxae separated by a pros-
ternal process and possessing a fold (plate) in
the posterior part of the coxae oriented anterior-
ly and restricting the movement of the protro-
chanter and, frequently, profemur (Fig. 85). In
contrast, Ommatinae usually show subspherical
or conical procoxae without a fold in the poste-
rior part and a free articulation between the
procoxa and protrochanter; however, some spe-
cies of Omma (i.e. Omma stanleyi Newman,
1839) have a clear plate on the procoxae.
The authors prefer to follow the traditional
interpretation that these taxa fall together with
Triadocupedinae (a group that demonstrates
many structural parallelisms with both cupedines
and ommatidines) into a single family, Cupe-
didae (see the Discussion below). Besides this,
not infrequently many of the genera proposed in
Cupedidae s.lato have been described following
a study of individual elytra, without reliable
characters to put them in any subfamily of those
recognized in the present publication. Accord-
ingly, these genera remain without subfamily
assignment (see below in Chapter 8 and Kirejt-
shuk, Ponomarenko, 2016).
The distance between the procoxae, com-
bined with the characters of elytral structure,
seem to be sufficiently stable to propose a key to
separate the subfamilies. Most species previ-
ously included in the subfamily Triadocupedi-
nae were described in a paper in which one of
the main diagnostic characters was the “procox-
ae separated by more than half of their diame-
ter” (Ponomarenko, 1966a: 48). All of them
originated from the same outcrop (Kyrgyzstan,
Madygen). Later, the same author, having re-
viewed all archostematans, wrote that the mem-
bers of Triadocupedinae had their procoxae
separated by one diameter, combined with the
absence of a prosternal process (Ponomarenko,
1969: 72). Some representatives included in
this subfamily showed rather weakly expressed
veins and some of them a very wide body with
widely explanate elytral sides, whereas some
others had a slender body and narrow elytral
sides. The latter share many similarities with
slender Cupedinae, except the somewhat elon-
gate head and wider pronotal sides. At the same
time, two taxa (viz. the Mesozoic Cretomerga
subtilis comb.n. and the Cenozoic Taxopsis
motschulskyi comb.n.) make the character of
procoxae separation somewhat ambiguous be-
cause of their comparatively widely separated
procoxae, combined with similarities in other
characters to those in the typical Cupedinae.
The character “presence of a prosternal process
between the procoxae” used by Ponomarenko
(1969) always means a more or less isolated
structure when the procoxae are separated and,
therefore, the use of of this kind of characters is
very doubtful. Nevertheless, at this level of our
knowledge it seems advisable to provisionally
maintain a certain taxonomic isolation in the
subfamily Triadocupedinae for the species with
widely separated procoxae and widely explan-
ate pronotal and elytral sides because they also
have some peculiarities in the pattern of their
venation and a rather strongly transverse head.
However, it seems more reasonable to transfer
the slender forms described in Triadocupedinae
to the subfamily Cupedinae: Asimma Ponomar-
enko, 1966, Kirghizocupes Ponomarenko, 1966,
Pterocupes Ponomarenko, 1966, and probably
some species (considered above in Chapter 8)
with a slender body described as members of the
genus Platycupes Ponomarenko, 1966.
The “subvertical” laterosternites seem to be
one of the probable synapomorphies of Omma-
70
A.G. Kirejtshuk et al.
tinae and Cupedinae. This structural feature can
be traced also in other cupedid fossils. For
instance, Mesocupes (Cainomerga) brevicor-
nis sp.n. and M. (C.) palaeocenicus sp.n. show
such a condition (Figs 18–31). It is highly prob-
able that subvertical laterosternites are congru-
ent with steeply sloping elytral sides. These
structures provide the elytra and the distal part
of the body a morphofunctional integrity. Per-
haps something similar occurs in different elat-
eriformians with (sub)vertical laterosternites and
steeply sloping elytral sides (Kasap, Crowson,
1975). However, Cupedinae and Ommatinae, in
contrast to the other archostematans, including
Tshekardocoleidae, have laterosternites held
almost perpendicular to the plane of the ven-
trites (Kirejtshuk et al., 2013).
Quite significant structural peculiarities are
present in the arrangement of the long longitu-
dinal rows of cells more or less reaching the
elytral apices, and in the pattern of fusions of
primary veins on the elytra, which seem to be
rather stable and thus allow grouping some of
the relatives among members of different sub-
families. The most usual number of these rows
can be nine or 10. However, the elytra of some
fossil representatives show 11 long longitudinal
rows (the genera Triadocupes Ponomarenko,
1966, and Platycupes from Triadocupedinae;
Kirghizocupes, probably also Priacmopsis
Ponomarenko, 1966, and Mesocupes from Cu-
pedinae; Notocupes Ponomarenko, 1964, from
Ommatinae etc.). Not infrequently, in addition
to the above-mentioned long longitudinal rows,
there are short rows of cells which are some-
times restricted to the scutellum by a short anal
vein (A2, see below), as well as a short row or
confused rows along the lateral edge in the
anterior part of the elytra (for example, extant
species of the genera Cupes and Prolixocupes;
also fossil species from different subfamilies).
Cainocups aixensis sp.n. (a taxon of an obscure
position in the family Cupedidae) demonstrates
eight clear long longitudinal rows of larger cells
in addition to four vague rows of smaller cells on
the sloping sides of each elytron, along its later-
al edge (see Fig. 76). Finally, a trend in forming
an irregular “cellulation” on the elytra is re-
vealed in many fossil Ommatinae, up to a com-
pletely diffuse and very fine “cellulation” in
some species of Cionocoleus Ren, 1995 (see
Jarzembowski et al., 2013a).
A generalized pattern (ground-plan) of pri-
mary veins at the elytral apices in Cupedidae
seems to have included a long anal vein running
along the suture and reaching the lateral edge of
the elytra (as in many Cupedinae and Ommati-
nae), and fusion of this vein with CuA (as in
many genera of all subfamilies and also Per-
mocupes Martynov, 1933 and some other Per-
mocupedidae Martynov, 1933). Ponomarenko
(1969) termed this long anal vein in Cupedidae
as A2, regarding that it corresponds to the anal
vein which is located lateral to the prescutellar
anal vein in the other archostematans, whereas
A1 is missing in Coleoptera, according to that
viewpoint. Thus, the prescutellar anal vein was
termed as A3 while the long adsutural anal vein
as A4. Such an interpretation has since become
traditional (Kirejtshuk, 2005; Shcherbakov et
al., 2009; etc.). Kukalová (1969) simultaneous-
ly proposed to consider this long anal line as A1,
the short prescutellar anal veins as A2 (Crow-
son, 1981; Merkl, 1988; etc.: “scutellary strio-
la”) and the long adsutural anal vein as A3,
although later she changed her initial scheme
into a less adequate one without proper explana-
tions (Kukalová-Peck, 1991). Fedorenko (2009)
suggested four hypothetical anal veins for a
ground-plan of the elytra in Coleoptera, yet
without affirming the presence of a long adsu-
tural anal vein. The earlier interpretations of
elytral venation beginning from Comstock
(1918), briefly reviewed by Crowson (1962),
include the anal vein along the suture which is
accompanied by a trachea and, according to
Forbes (1922) and Crowson (1962), this vein is
to be termed as A2. This opinion is rooted in
tracheation represented by two anal tracheas.
However, the short “scutellary striola” could
also be interpreted as a short vein with a reduced
trachea. After comparing the ancient represen-
tatives of different Palaeozoic groups of the
order, Kirejtshuk et al. (2013) came to the
conclusion that it appears impossible to trace
back the anal vein considered by Ponomarenko
Taxonomy of the reticulate beetles of the subfamily Cupedinae
71
as A1. Therefore, Kirejtshuk et al. (2013) termed
the long anal vein as A1, the short vein (“scutel-
lary striola”) as A2 and the adsutural line as A3,
while Fedorenko’s hypothesis fails to find a
factual support. At the same time, there are
many transformations in A1 shape which some-
times could be interpreted as phyletic traces
while in other cases transformations could cor-
respond to a sequence of homoplastic changes.
Very frequently, vein A1 does not reach the
lateral edge of the elytra, but ends in the distal
part of the elytral suture (probably in almost all
Triadocupedinae, numerous Ommatinae, only
the genus Prolixocupes among Cupedinae;
somewhat similar transformations occur also in
other archostematan families). Another usual
mode of transformation is a trend to have an
independent ending of the most primary veins
along the lateral edge or, more correctly, fusion
of them with Sc, which is not infrequently scarce-
ly expressed. Such trends can be traced among
cupedines and ommatines, although it is more
characteristic of the latter group (including the
Recent members of Omma and Tetraphalerus),
and only rarely occurs among cupedines (the
genera Mesocupes and Miocupes). The primary
elytral veins of Kirghizocupes, the oldest known
cupedine group, are rather weak and show no
trace of fusion, but gradually become more
narrowly separated by progressively smaller
cells at the apices. In some cases, fusions of the
primary veins follow a different pattern. For
example, Cainocups aixensis sp.n. has vein A1
subrectilinearly reaching the elytral apices and
apically fusing with a vein resulting from the
fusion of CuA, M, and R (see Fig. 76). Finally,
if the Triadocupedinae are generally character-
ized by weakly elevated veins, the elytra of
some fossil groups show only long longitudinal
rows of very large cells sometimes not very
regular in shape (in the cupedine genus Meso-
cupes and some genera in Ommatinae, Litho-
cupes Ponomarenko, 1966, Brochocoleus Hong,
1982, etc.) to completely diffuse and fine cells
without a trace both of rows and veins (Ciono-
coleus). In some cases, all veins become sub-
equally expressed and look like thick rollers
without fusion at the apex (f.i. Anaglyphites
stat.n.: Figs 95–100). Such a condition of elytral
structure looks like intermediate between the
one typical of the members of Cupedinae and
that characteristic of the members of Taldycu-
pedidae Rohdendorf, 1961 and some other Palae-
ozoic groups.
Other quite important characters demon-
strating parallel transformations can be observed
in structural peculiarities of the head (convexi-
ties and tuberculations on its upper surface and
the position of the antennal insertions) and ab-
domen (type of articulations of segments), which
are frequent at least among Cupedinae and
Ommatinae, probably also in Triadocupedinae.
These characters are considered in more detail
for the subgroups of the subfamily Cupedinae
(see below). A similar situation seems to be
characteristic of the other subfamilies.
2. Key to subfamilies of Cupedidae
(based on the key proposed by Ponomarenko,
1969)
1. Elytra with long longitudinal rows of very large
cells and extremely narrow interspaces between
them, veins not elevated; procoxae widely sepa-
rated (distance between them subequal to or
greater than their diameter); elytra comparative-
ly wider than in other subfamilies, up to very
wide, usually flatly or gently sloping and widely
explanate on sides, with apparently more or less
wide epipleura and veins very weakly raised .
............ Triadocupedinae Ponomarenko, 1966
– Elytra with long longitudinal rows of moderately
large cells and interspaces between them making
it possible to observe veins; procoxae usually
narrowly separated (distance between them less
than their diameter) to/or conjoining; elytra fre-
quently with more or less clear veins, their sides
more or less steeply sloping (up to subvertical)
and narrowly explanate; if procoxae widely sep-
arated, elytra narrow, with subvertically slop-
ping sides and rather narrow epipleura (Cre-
tomerga gen.n. and Taxopsis gen.n.) ............. 2
2. Procoxae more or less separated, with a flat fold
oriented anteriorly and covering protrochanter
and base of profemur; head mostly of usual
shape with a gradually narrowing frons .........
................................ Cupedinae Laporte, 1836
– Procoxae conjoining, often without a flat fold
oriented anteriorly and covering protrochanter,
sometimes conically projecting; head often some-
what elongate and frequently with a rather wide
anterior part .....................................................
.................... Ommatinae Sharp et Muir, 1912
72
A.G. Kirejtshuk et al.
3. Subfamily Cupedinae Laporte, 1836
Type genus Cupes Fabricius, 1801
= Priacmini Crowson, 1962; type genus: Priacma
LeConte, 1861; currently not considered as separate tribe;
= Mesocupedini Ponomarenko, 1969; type genus:
Mesocupes Martynov, 1926; currently not considered as
separate tribe.
Notes on the taxonomy and systematics of
Cupedinae
This subfamily is sometimes subdivided into
tribes (Crowson, 1962; Ponomarenko, 1969;
Tan, Ren, 2006, 2009; etc.). However, the tribe
Priacmini has been shown to be impossible to
discriminate based on external structures (see
below). The unique structure of the male pre-
genital sclerites and aedeagus in Priacma, makes
this genus different from other genera of the
subfamily. However, this character is not appli-
cable for most fossils (Kirejtshuk et al., 2010).
The tribe Mesocupedini has been proposed for
some Mesozoic genera with flat (not overlap-
ping) abdominal ventrites and a weakly tuber-
culate dorsal surface of the head. Abdominal
ventrites, if they are tegular and/or showing a
transverse depression along the base following
Ponomarenko (1969), are frequently regarded
by paleocoleopterists as an important diagnos-
tic character for supraspecific taxa of the family.
The neontologists usually do not use them,
although sometimes these characters are men-
tioned in the species descriptions (Atkins, 1963)
or recognised as diagnostic for some groups
(Lawrence, 1999). This structural feature (artic-
ulation type of abdominal ventrites) could be
sometimes rather efficient for grouping the spe-
cies, but it is rather variable in larger groups of
related species. For example, the abdominal
ventrites of extant representatives are complete-
ly flat (apparently also in fossil species of Omma
and Tetraphalerus) while in the extant Para-
cupes brasiliensis Kolbe, 1898, they have both
a transverse depression at the base and a convex
roller behind the middle. On the other hand,
extant representatives of Priacma, Ascioplaga
Neboiss, 1984, stat.n., Prolixocupes Neboiss,
1960, and Rhipsideigma Neboiss, 1984, dem-
onstrate only a weak depression and a weak
roller behind it (usually scarcely traced). At the
same time, most recent species of Cupes have a
rather raised depression and a roller on each of
ventrites 2–4. However, among specimens from
a large series of modern Cupes capitatus Fabr-
icius, 1801, or C. mucidus Chevrolat, 1829, a
highly variable degree of expression of these
features can be observed. Their congeners from
Baltic amber also show variability from com-
pletely flat ventrites up to ventrites with a clear
transverse depression (groove) (Kirejtshuk,
2005). The abdominal ventrites in compression
fossils frequently show overlapping between
them, although in some cases they are not clear-
ly visible or absent. Therefore, many remains
cannot be diagnosed, based on these characters
of ventrites. Thus, using these features for group-
ing has caused some confusion in defining the
tribes and subfamilies (partly considered be-
low) and requires caution, taking into consider-
ation the above variations.
Elytral venation and the pattern of cells (see
above in the “Notes on structural peculiarities to
the family Cupedidae”) represent highly impor-
tant characters for dividing the subfamily into
groups. Many slender species can be grouped,
based on the expression of the differences be-
tween primary and secondary veins, the number
of veins, and the patterrn of conjunction of the
primary veins, in particular, the sequence in
their conjunction at the elytral apices. The most
outstanding fusions are, as in Ommatinae groups,
the sequential fusion of CuA with A1 and M,
sometimes a separate fusion of A1 and CuA and
a fusion of M to R. This group of structural
peculiarities is somehow associated with the
character of sloping of the sides (gentle or
subvertical along a flat median part of the elytral
disk), of the bending line and of the explanation
(flattening) of the lateral submarginal stripes of
the elytra. It is noteworthy that fusion of veins in
species with distinct primary veins is usually
quite distinct, whereas the fusions of veins in
forms with weakly raised primary veins are
vague or even quite variable. In cases when the
elytral veins are not elevated, the fusions of
veins at the apices are obscure or untraceable.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
73
Some forms from the Triassic (including Kir-
ghizocupes) have the elytra with weak primary
veins which look like reaching the apices with-
out fusion.
The degree of separation of antennal inser-
tions was used by Crowson (1962) for tribal
diagnostics in the subfamily Cupedinae. Later
reseachers followed him in the discrimination
of Cupedini and Priacmini. Application of this
character as a diagnostic group character intro-
duced a certain confusion because Priacma
serrata, has comparatively small eyes and the
distance between its antennal insertions is com-
parable to that in other Recent Cupedinae (in
Cupes mucidus and other species with compar-
atively larger eyes). The most important fact is
that many extant (including Priacma serrata)
and Cenozoic cupedines (except members of
Mesocupes (e.g. Cainomerga subgen.n.) and
Menatops gen.n.) show the antennal insertions
rather deep at their anterior edge, close to each
other and more or less covered by a strong
tubercle of the frons at their posterior edge.
Their scapes are also rather thick while those of
many Mesozoic members of the subfamily are
moderately thick to even rather thin and more
widely (or moderately) separated. Finally, Me-
sozoic cupedines usually have completely open
antennal insertions (not covered by tubercles
posteriorly). This character can be redefined,
taking into account that the antennal insertions
of most of the extant cupedines (including Pri-
acma serrata) are much more narrowly separat-
ed than the distance between the inner edges of
the eyes on the upper surface of the epicranium.
On the contrary, many Mesozoic species gener-
ally have somewhat different structures of the
head, with varying shapes of the scape, and
therefore they could be regarded as a set of taxa
distinct from modern cupedines with compara-
ble partial similarities to modern species of Cupes
and Priacma. It is necessary to mention that the
proportion of length of antennae to body length
(or even the comparative length of a particular
antennomere, sometimes used for group diag-
nostics) shows some variability as well, fre-
quently depending on body size and sex (Kirejt-
shuk, personal observation). The comparative
length of the entire antennae can be a reliable
character only in some exceptional cases, but
not in many supraspecific taxa as it has frequent-
ly been used by some paleocoleopterists.
An important feature of many cupedine
groups is structure of the upper integument of
the epicranium, i.e. configuration and degree of
development of convexities (tubercles) on the
frons and vertex. These structural peculiarities
are very easy to study in the modern taxa, but
often difficult to trace in compression fossils.
The structure of the tubercles over the antennal
insertions is particularly important if they are
raised. This feature is very characteristic of
modern cupedines, while many fossil members
have convexities only behind the antennal inser-
tions. In such cases, a frontal anterior depres-
sion (located in modern species just before the
anterior edge of the frons, in front of the tuber-
cles covering the antennal insertions) is shifted
posteriad behind the antennal insertions. This
structural change seems to have been frequently
associated with some convexity at the anterior
edge of the frons. Similar transformations can
also be observed among ommatines and triado-
cupedines. In some case the median space be-
tweentubercles (longitudinal convexities) looks
like a longitudinal depression and bears a more
or less clear sculptural line (or sometimes a
weak and narrow carina). Therefore, in many
fossils the depression shifted posteriad looks
like an expressed Y- or V-shaped depression
which sometimes is called “epicranial suture”
(Ponomarenko, 1969). This term has since been
in use even without quotation marks (e.g. Tan et
al., 2012; Jarzembowski, 2013a; etc.), as an
equivalent to the traditional term coined for the
suture on the epicranium of beetle larvae and
which also concerns both larvae and adults in
polyneopteran groups. Usage of this term for
coleopterous adults could easily create great
confusion, although rediments of epicranial su-
ture can be traced in different coleopterous
groups, including fossil and modern cupedines.
Some early Cretaceous species have been
attributed to the genus “Priacma” (Ponomaren-
ko, 1986, 1997, 2000; Soriano, Delclós, 2006),
even though those remains share many charac-
74
A.G. Kirejtshuk et al.
ters of a whole number of cupedine genera.
However, in fact it is impossible to be certain
about the generic placement of any of them. It
would be more reasonable at the current level of
our knowledge to regard these species of “Pri-
acma” as Cupedinae or Cupedidae without any
generic assignment (see below Chapter 8), be-
cause they fail to demonstrate any evidence of a
closer relationship to Priacma serrata than some
other Mesozoic cupedines do. Hieke & Pietrze-
niuk (1984) recorded the genus Priacma from
Baltic amber, but that finding was not accompa-
nied by a picture or any other piece of evidence.
As a result, the genus Priacma remains mono-
typic. Its type species, Priacma serrata, is char-
acterized by a unique structure of its aedeagus
(Priacma-type) among extant cupedids, although
species of the Mesozoic Gracilicupes Tan, Ren
et Shin, 2006, and some still undescribed fossils
show similar genital structures (personal com-
munication of A.G. Ponomarenko). The differ-
ence in structure of the male genitalia cannot be
used for separating most of the fossil genera
because their genitalia remain unknown. Never-
theless, the male genitalia of the few fossil
species from different genera known (Apriacma
gen.n., Latocupes Tan et Ren, 2006, and Meso-
cupes Martynov, 1926) certainly look more
similar to those of the Recent genera closely
related to Cupes (Cupes-type) than to those of
Priacma, albeit most of the other (external)
characters appear to be more similar to those in
Priacma serrata rather than to those of the other
modern genera. Traces of an aedeagus in the
holotype of Gracilicupes tenuicruralis Tan, Ren
et Shih, 2006, suggest that organ to have been be
at least considerably wider than in most cu-
pedines (or cupedids) with a rather narrow penis
trunk particularly at apex and with two pairs of
lateral lobes, and widely and separately curved
narrow parameres (Priacma-type — see de-
tailed description below). All known male gen-
italia of Cupedinae, except those in Priacma
serrata and Gracilicupes tenuicruralis, show a
trilobed Cupes-type of the male genital struc-
ture, including a tube-like penis trunk (median
lobe without lateral compression) and compar-
atively wide and usually subflattened parameres
(lateral lobes) with narrow appendages at base
and at apices. These exceptions (i.e., aedeagi of
Priacma serrata and Gracilicupes tenuicrura-
lis) demonstrate many characters occurring in
other holometabolan groups which could be a
symplesiomorphy for Priacma and Coleoptera
as a whole (see below Chapter 9).
4. Key to fossil and Recent genera and
subgenera of Cupedinae
(partly based on keys in Crowson, 1962; Atkins, 1963;
Ponomarenko, 1969; Neboiss, 1984)
1. Frons distinctly excavate along anterior part of
rather long and very large eyes, but convex
medially; transverse diameter of eyes nearly as
great as half the distance between rather long
eyes; antennal insertions rather narrowly sepa-
rated and covered by distinct tubercles on frons;
lateral rows of punctures confluent in distal half
of elytra; scales on elytra elongate (lanceolate)
and sparse. Fossil; late Cretaceous (USA, New
Jersey, Old Crossman’s Clay Pit). Figs 80–83
............................................... Cupopsis gen.n.
– Frons convex or flattened along anterior part of
eyes or with a Y-shaped depression in the middle
or a small triangular depression between anten-
nal insertions; transverse diameter of eyes clear-
ly less than half the distance between eyes (only
species of Menatops gen.n., Mesocupes (Caino-
merga) ponti sp.n. and M. (Mesocupes) spinosus
(Ponomarenko, 1964), comb.n. could be with
transverse eye diameter subequal to distance
between eyes); combination of remaining char-
acters different ................................................ 2
2. Pronotum about as long as wide to almost elon-
gate, arcuate on sides, gradually narrowing both
anteriorly and posteriorly, its anterior angles not
projecting (as on the photograph in Fig. X, 7
(page 97) but not in drawing in Fig. 5 (page 97)
in Ponomarenko, 2006); elytra with 10 regular
long longitudinal rows of comparatively small
suboval cells, but without clear veins; temples
behind small eyes longer than anterior part of
head before eyes; length of body ca. 6.5 mm.
Fossil; early Cretaceous (England, Early Pur-
beck) ....... Chalepocarabus Handlirsch, 1906
– Pronotum more or less transverse, variable in
configuration, but not arcuate on sides, its ante-
rior angles more or less distinctly projecting;
combination of remaining characters different
....................................................................... 3
3. Head with very long temples (longer than frons
before eyes) and subflattened; all primary veins
on elytra parallel, not fusing at apices; veins
Taxonomy of the reticulate beetles of the subfamily Cupedinae
75
equally weakly raised, but well expressed and
cells between them with a weak arrangement in
regular rows; pronotum strongly transverse (about
twice as wide as long) and subrectilinearly wid-
ening from base to distinct anterior angles; body
elongate; length of body 12.5 mm. Fossil; Mi-
ocene (Czech Republic, Pochlovice). Ponomar-
enko, 1973: 104, Figs a–c ..............................
...................... Miocupes Ponomarenko, 1973
– Head with short or moderately long temples (at
most as long as frons before eyes, although
Cupidium abavum Ponomarenko, 1968, has rath-
er small eyes located at a comparable distance
from both anterior edge and widest point of
temples); elytra with non-subparallel veins and
usually fusing at apices; combination of remain-
ing characters different ................................... 4
4. Head with a clear Y- or V-shaped depression on
frons at or behind antennal insertions, formed
because of paramedian convexities behind an-
tennal insertions .............................................. 5
– Head without Y-shaped depression on frons, at
most with a transverse emarginate depression
between antennal insertions or a subtriangular/
subsemicircular depression in anterior part of
frons before antennal insertions (although head
of Mesocupes (Mesocupes) bidens Ponomaren-
ko, 1964, has traces of a weak Y- or V-shaped
depression; in the comparatively large Priac-
mopsis adumbrata Ponomarenko, 1966, the sub-
triangular depression on the frons is very
large) .............................................................. 10
5. Prosternum before procoxae at most about as long
as procoxae; head with neck half as wide as head
at temples, without tubercles over antennal in-
sertions; elytra with primary and secondary veins
comparable in expression and with somewhat
transverse, large cells arranged in nine long
longitudinal rows; body more than three times as
long as wide; distance between antennal inser-
tions greater than half of the distance between
eyes .................................................................. 6
– Prosternum before procoxae much longer than
procoxae; head with neck half as wide as head at
temples and with more or less raised tubercles
over antennal insertions; combination of remain-
ing characters different ................................... 7
6. Procoxae narrowly separated (distance between
them less than their diameter); head with anten-
nal insertions completely open; body length ca.
9.2 mm. Fossil; early Cretaceous (China, Liaon-
ing). Figs 91–92 ..............................................
......................... Furcicupes Tan et Ren, 2006
– Procoxae widely separated (distance between them
greater than their diameter); head with antennal
insertions slightly covered with tubercles; body
length ca. 11.5 mm. Fossil; early Cretaceous
(China, Liaoning). Figs 77–79 .......................
.......................................... Cretomerga gen.n.
7. Elytra with nine long longitudinal rows of cells
and a very clear difference between main and
secondary veins; body at least three times longer
than wide; metaventrite about as wide as long;
anterior angles of pronotum projecting and sub-
acute; elytra with nine long longitudinal rows of
cells; primary veins of elytra better expressed
than secondary ones; elytra with disks flat and
subvertically sloping along M; pronotum about
1.5 times as wide as long and widening anterior-
ly; scape subequal in length and slightly thicker
than antennomeres 2–11 or 3–11; length of body
ca. 9.5–16.5 mm. Fossil; early Cretaceous (Chi-
na, Liaoning). Figs 73–75 ..............................
.............................................. Apriacma gen.n.
– Elytra at least with 10 long longitudinal rows of
cells and only a slight difference between main
and secondary veins; metaventrite markedly wider
than long; anterior angles of pronotum not pro-
jecting and rectangular or blunt to rounded; body
less than three times as long as wide (only in
Latocupes elytra with 9 longitudinal rows of
cells while in L. bellus comb.n. anterior angles
slightly subacute); combination of remaining
characters different ......................................... 8
8. Antennomere 3 longest (at least twice as long as
each of antennomeres 2 and 4); elytra with ap-
parently gently sloping sides and 10 long longi-
tudinal rows of large cells subparallel or only
slightly coverging at apices; primary veins, if
expressed, without fusion A1 and CuA; prono-
tum slightly transverse (less than 1.5 times as
wide as long) and with arcuate sides, its anterior
and posterior angles rounded; length of body ca.
5.8–10.0 mm. Fossil; late Jurassic (Kazakhstan,
Karatau), early Cretaceous (Transbaikalia, Bais-
sa; Mongolia, Bon Tsagan). Figs 95–102 ......
....................................................... Mesocupes
(Anaglyphites Ponomarenko, 1964, stat.n.)
– Antennomere 3 not longest; elytra with long longi-
tudinal rows of cells making it possible to trace
fusion of A1 and CuA; combination of remain-
ing characters different ................................... 9
9. Elytra with subvertically sloping sides and long
longitudinal rows of subquadrangular, subsquare
and sublongitudinal cells or at most slightly
transverse, much more than twice as long as wide
combined (usually about 2.5 times); pronotum
slightly transverse (less than 1.5 times as wide as
long) and not widening anteriorly; scape nearly as
thin as following antennomeres; length of a very
slender body ca. 4.0–7.5 mm. Fossil; Middle–late
Triassic (Kyrgizstan, Madygen). Figs 109–116
............... Kirghizocupes Ponomarenko, 1966
(= Mesocupoides Ponomarenko, 1969, syn.n.)
76
A.G. Kirejtshuk et al.
– Elytra with gently sloping sides and long longitu-
dinal rows of subquadrangular and, on disks,
strongly transverse cells (about twice as wide as
long), about 1.5 times as long as wide combined
(about twice); pronotum strongly transverse (1.5–
2.0 times as wide as long); scape markedly
thicker than following antennomeres; length of a
more robust body ca. 12–27 mm. Fossil; Middle
Jurassic (Inner Mongolia, Daohugou), early Cre-
taceous (China, Liaoning; Bejing Province, Lus-
hangfen). Figs 117–119 ..................................
........................... Latocupes Tan et Ren, 2006
(? = Pulchicupes Ren, 1995)
10. Body length about 30 mm; metacoxae very
slightly extended in mesal part, forming a gentle
curve along their posterior edge; metaventrite
about twice as wide as long; head with tubercles
over antennal insertions and at temples; tarsom-
ere 4 lobed; antennomere 2 (pedicel) much shorter
than antennomeres 1 (scape) and 3; long longitu-
dinal rows on elytra with small suboval to sub-
quadrate cells; pronotum about twice as wide as
long; head somewhat tuberculate. Fossil; early
Cretaceous (Transbaikalia, Baissa). Ponoma-
renko, 1969: 112, Fig. 61; Figs 120–121 ......
................... Priacmopsis Ponomarenko, 1966
– Body not longer than 20 mm; metacoxae moderate-
ly or strongly extended in mesal part and clearly
subtriangular; metaventrite markedly less than
twice as wide as long; combination of remaining
characters different ....................................... 11
11. Head with very large eyes and much wider than
pronotum with arcuate sides and nearly rounded
anterior angles; prosternum before procoxae not
longer than procoxae; elytra with primary and
secondary veins comparable in expression and
with 10 regular rows of large cells; head narrow-
ing behind very large eyes (along temples); scape
significantly thicker than following antenno-
meres, forming subfiliform antennae; abdominal
ventrite 1 much longer than each of ventrites 2–
4; body about 2.5 times as long as wide; eyes
with transverse diameter as great as third of
distance between eyes; femora and tibiae rather
thin and long; male genitalia of Priacma-type
(penis trunk with an acuminate apex, parameres
very narrow and strongly curved); length of body
ca. 10.8–11.4 mm. Fossil; Middle Jurassic (In-
ner Mongolia, Daohugou). Figs 93–94 ..........
............. Gracilicupes Tan, Ren et Shin, 2006
– Head more or less narrower than pronotum (head
subequal to or only slightly wider than pronotum
only in some Mesocupes s.lato, which have,
however, smaller eyes and antennae rather sub-
moniliform, and also in some Cupes, which
have, however, an always longer prosternum,
pronotal sides not arcuate, distinct anterior an-
gles and more or less smaller eyes); prosternum
before procoxae usually longer than procoxae
(although prosternum in some species of Caino-
merga subgen.n. and Mesocupes s.str. about as
long as procoxae); combination of remaining
characters different ....................................... 12
12. Part of prosternum before procoxae shorter than
procoxae; head triangular and with comparative-
ly small eyes located at a comparable distance
from both anterior edge and widest point of
temples; antennomeres 1–3 with comparable
lengths; pronotum about twice as wide as long,
somewhat shorter than head and with arcuate
sides; body length ca. 10.0 mm. Fossil; late
Jurassic (Kazakhstan, Karatau). Figs 107–108
...................... Cupidium Ponomarenko, 1968
– Part of prosternum before procoxae longer than
procoxae; combination of remaining characters
different ......................................................... 13
13. Elytra with nine rows of clear large cells or, if
with 10 rows, 1–2 rows at lateral edge not ex-
tending behing basal third of elytra; primary and
secondary veins of elytra very contrasting in
levels of development; elytral disks flat (hori-
zontal) and sides vertically slopping along M;
antennal insertions narrowly separated; distance
between them usually less than transverse diam-
eter of moderately large eyes; prothorax usually
with more or less expressed tarsal grooves ....
..................................................................... 14
– Elytra at least with 10 rows of cells of different
outlines, extending behind the middle; combina-
tion of remaining characters different; elytra
with regular rows of clear larger cells reaching
elytral apices or cells arranged in less clear rows
or very small; primary and secondary veins of
elytra similar in degree of development or, rare-
ly, with contrasting primary veins (in Paracu-
poides subgen.n.); antennal insertions usually
widely separated (in Paracupes s.str., Paracu-
poides subgen.n. and Priacma they are compar-
atively narrowly separated); eyes of different
size; prothorax without traces of tarsal grooves
..................................................................... 19
14. Prosternum with process about as wide as pro-
coxae (or slightly narrower than base of epipleu-
ra) and widely rounded at apex; elytra with
strongly transverse cells (about twice as wide as
long); head with two pairs of paramedian tuber-
cles (at antennal insertions and at base of eyes)
and strongly projecting temples; elytra with a
short anal vein isolating three cells at scutellum;
length of body ca. 16.1–18.1 mm. Fossil; late
Eocene (Baltic amber). Figs 121–122 ............
............................................... Taxopsis gen.n.
– Prosternum with process much narrower than
procoxae (or at most half as wide as base of
Taxonomy of the reticulate beetles of the subfamily Cupedinae
77
epipleura) and narrowly rounded to subacute at
apex; elytra with cells of different shape, but
never strongly transverse (only slightly wider
than long); combination of remaining characters
different ......................................................... 15
15. A1 fused with CuA and then with M, futher
curving and ending in distal portion of the suture
(not reaching lateral edges of elytra); all four
upper head tubercles obtuse, posterior ones dis-
tinctly much larger and longer than anterior
ones; scales on elytra suboval; tarsal grooves
anteriorly separated by full width of prosternum;
basal half of elytra with ten rows of punctures,
the 7th and 8th confluent and terminating before
the middle; length of body 10–12 mm. Recent;
Western Hemisphere species (Chile; Argentina;
USA: California, Arizona). Neboiss, 1960: 20,
Figs 3 ................. Prolixocupes Neboiss, 1960
– A1 fused with CuA and then reaching lateral edges
of elytra; one or two pairs of somewhat conical
head upper tubercles, the latter subequal in size;
subsemicircular or laterally flattened; combina-
tion of remaining characters different 16
16. Head with one pair of distinct upper paramedian
conical tubercles situated above bases of anten-
nae .................................................................. 17
– Head with 2–3 pairs of upper paramedian conical
tubercles, anterior pair situated above base of
antennae, posterior pairs along and behind eyes
..................................................................... 18
17. Elytra with punctures free of scales; male abdom-
inal tergite IX slender, distinctly constricted
before a rounded base; length of body ca. 6–16
mm. Recent; Australian species. See Neboiss,
1984: 469, 470, 475; Figs 85–88, 99 & 117 ..
Adinolepis (Adinolepis) Neboiss, 1984, s.str.
– Elytra covered with oval or elongate oval scales;
male abdominal tergite IX rather broad, gradual-
ly tapered towards base, middle of apex extend-
ed into an elongate, slightly spatulate lobe; length
of body ca. 9.0–14.5 mm. Recent; New Cale-
donian species. See Neboiss, 1984: 469 & 476,
Figs 89–90 & 118–119 ................ Adinolepis
(Ascioplaga Neboiss, 1984) stat.n.
18. Elytra with a dense cover of scales more or less
completely covering contours of elytral cells;
apices extended into an acute point; preapical
conjoining of main veins; A1 conjoining with
median vein at elytral apex, forming a strong
elevation, while CuA not reaching this conjoin-
ing; length of body ca. 11.0–20.5 mm. Recent;
East African (Tanzania) and Madagascar spe-
cies. See Neboiss, 1984: 470 & 476 Figs 91–94
& 120–123 ...... Rhipsideigma Neboiss, 1984
– Elytra with scales mostly on intervals, cell outlines
not obscured by scales; elytra with apices not
extended into an acute point; A1 fused with CuA
before subapical fusion with M; length of body
6.0–12.5 mm. Fossil, Palaeocene (Menat),
Eocene (French Oise and Baltic amber; Germa-
ny, Messel, Eckfeld), Pliocene (Germany, Will-
ershausen, Niedersachsen); Recent; Asian (Palae-
archiarctic Province (Primorsky Kray, Japan,
continental China, Philippines) and Indo-Ma-
layan Region, including the Papuan Province
(Kalimantan, New Guinea)), Australian, North
American (Canada, USA), Polynesian (Hawaii),
South African species. Figs 41–72, 169–172
.................................... Cupes Fabricius, 1801
(=Cupoides Motschoulsky, 1856; Tenomerga
Neboiss, 1984; Distocupes Neboiss, 1984)
19. Part of abdominal ventrite 1 exposed from under
metacoxae not longer than each of following
ventrites 2–4; gular sutures very widely separat-
ed, with a distance between them more than 2/3
as great as maximum width of head; male geni-
talia of Priacma-type (penis trunk with an acumi-
nate apex, parameres very narrow and strongly
curved); head with antennal insertions covered
by strong tubercles; elytra with irregularly trun-
cate apices and 10 rows of cells reaching elytral
apices; elytral lateral carina thickened as a rather
wide stripe with sharp tubercles; length of anten-
nomere 2 (pedicel) less than half the length of
antennomeres 1 (scape); distance between eyes
more than half the distance between eyes; body
rather slender, length ca. 10–17 mm. Recent;
USA (West of the Rocky Mountains: California,
Oregon, Washington, Idaho, Montana, British
Columbia). Figs 124–134 ...............................
.................................. Priacma LeConte, 1874
– Part of abdominal ventrite 1 exposed from under
metacoxae markedly longer than each of follow-
ing ventrites 2–4; gular sutures moderately widely
to narrowly separated, with a distance between
them about 1/3 as great as maximum width of
head or even less; male genitalia of Cupes-type
(usual cupedine trilobed structure); combination
of remaining characters different ................. 20
20. Head very short and strongly transverse, with
transverse diameter of eyes markedly more than
half the distance between eyes; pronotum 1.5–
2.0 times as wide as long; anterior angles of
pronotum with a distinct top; primary veins more
raissed than secondary ones; antennomere 2
(pedicel) about half as long as antennomere 3;
length of body 7.2–9.1 mm. Fossil; Palaeocene
(Menat). Figs 10–17, 165–168 .......................
.............................................. Menatops gen.n.
– Head elongate and at most slightly transverse, with
transverse diameter of eyes less than half the
distance between eyes; pronotum much less than
twice as wide as long; combination of remaining
characters different ....................................... 21
78
A.G. Kirejtshuk et al.
21. Elytra obliquely truncate at apices; head with
strong tubercles over antennal insertions; elytra
with subvertically slopping sides, narrowly ex-
planate along subreclilinear lateral edges. Re-
cent; South America .......................................
.......................... Paracupes Kolbe, 1898...22
– Elytra subacute or nearly rounded at apices; head
without raised tubercles over antennal inser-
tions; elytra with gently slopping sides and more
or less explanate along broadly rounded lateral
edges. Fossil; Mesozoic and Cenozoic ..........
.................... Mesocupes Martynov, 1926...23
22. Elytra with slight differences between primary
and secondary viens (both somewhat obliterat-
ed), a serrate apical edge and A1 subparallel to
suture; elytral integument covered by narrow
elongate (to elliptic) scales with truncate apices,
leaving cells partly open; elytral lateral carina
thickened as a rather wide stripe with sharp
tubercles; pronotum widening anteriorly; anter-
omedian gular plate slightly longer than wide
(1.2 times); length of body ca. 12–15 mm. Re-
cent; Brazilian species. Figs 84–86 ................
... Paracupes (Paracupes Kolbe, 1898), s.str.
– Elytra with distinct primary and weak secondary
viens, a smooth apical edge and A1 clearly
deviating from suture in both anterior and poste-
rior fifths; elytral integument thoroughly (in-
cluding cells) covered by subtriangular scales
(widening to a truncate apex); elytral lateral
carina simple; pronotum subparallel-sided; an-
teromedian gular plate almost twice longer than
wide (1.9 times); length of body 10.4 mm. Re-
cent; Ecuador species. Figs 87–90 .................
............ Paracupes (Paracupoides subgen.n.)
23. Cells in long longitudinal rows on elytra sub-
quadrate or somewhat transverse; elytra about or
somewhat more than twice as long as wide com-
bined; length of body 16.3–18.5 mm. Fossil;
Palaeocene (Menat). Figs 5–9, 18–40 ............
............. Mesocupes (Cainomerga subgen.n.)
– Cells in long longitudinal rows on elytra more or
less oval and sparser; elytra less than twice as
long as wide combined; body length ca. 4.5–8.4
mm. Fossil; Late Jurassic (Kazakhstan, Kara-
tau), Early Cretaceous (Spain, Las Hoyas). Figs
1–4, 103–106 ................................ Mesocupes
(Mesocupes Martynov, 1926), s.str.
5. Subfamily Cupedinae from Menat
5.1. Geological setting and locality infor-
mation for Menat specimens
The Middle Paleocene Menat fossil site
(Menat Basin, Puy-de-Dôme, France) is a vol-
canic maar containing a lake ca. 700 m in
diameter, which at present contains sedimenta-
ry rocks (spongo-diatomites) with remains of
diverse aquatic and terrestrial flora and fauna
(Piton, 1940; Nel, 1989, 2008; Wappler et al.,
2009; Nabozhenko, Kirejtshuk, 2013; etc.). This
locality has been well known since the begin-
ning of the 20th century (Fritel, 1903; Laurent,
1912; etc.) and a general preliminary overview
of its biota was published by Piton (1940).
According to Wappler et al. (2009), the plant
and insect diversity in the lacustrine outcrop of
Menat is comparatively high, confirming the
results of Piton (1940), who was the first to
describe the fossil flora and fauna. Recent field
investigations made by our team strongly sug-
gest that it is even higher than thought before.
The composition of faunal and floral remains
makes it possible to conclude that this lake was
surrounded by a paratropical evergreen forest
and the palaeoenvironment was very warm and
humid, supporting a high insect diversity. Fol-
lowing the pollen, paleomammalian stratigraphic
and radiometric K/Ar analyses, the age of Menat
was estimated at 59 Ma (Kedves, Russel, 1982;
Nel, 2008). However the newest estimates based
on macroflora postulate its age within 60–61
Ma (Wappler et al., 2009). Interestingly, bee-
tles dominate the animal biodiversity in terms of
species number and the abundance of speci-
mens, as they represent more than 80% of the
animals collected in this outcrop (after the col-
lection of ca. 1,200 animals made during the
years 2013–2015). Nevertheless, the most abun-
dant beetle group remains the Curculionoidea
while the Archostemata are represented by 32
fossils, and part of them will be considered in a
further paper.
5.2. Genus Mesocupes Martynov, 1926
Type species Mesocupes primitivus Mar-
tynov, 1926, by monotypy, fossil.
Composition. The genus includes 16 spe-
cies from the late Mesozoic and Paleocene [see
the list and comments in Chapter 8 and in the
catalogue by Kirejtshuk & Ponomarenko
(2016)]. The subgenera Mesocupes s.str., Ana-
glyphites stat.n. and Cainomerga subgen.n. rec-
ognized in this paper can be diagnosed after the
above key and, in addition, these groups have
Taxonomy of the reticulate beetles of the subfamily Cupedinae
79
Figs 1–4. Mesocupes (Mesocupes) primitivus Martynov, 1926 (neotype “PIN 2452/682”, photographed
under Leica MZ 9.0 with camera DFC290); Upper Jurassic, Oxfordian (or/and Kimmeridgian); Kazakhstan,
Karatau: 1 — print covered by alcohol; 2 — body, ventral, with broken lines for outlines of dorsal structures
of head and sutural edges of elytra; 3 — dry print; 4 — aedeagus. Length of specimen 6.4 mm.
Рис. 1–4. Mesocupes (Mesocupes) primitivus Martynov, 1926 (неотип “PIN 2452/682”, фотографии,
выполненные на микроскопе Leica MZ 9.0 с камерой DFC290); верхняя юра, оксфордский (или/и
кимериджский) яруса; Казахстан, Каратау: 1 — отпечаток, покрытый спиртом; 2 — тело, снизу;
прерывистой линией показаны очертания верхних структур головы и шовных краев надкрылий; 3 —
сухой отпечаток; 4 — эдеагус. Длина экземпляра 6,4 мм.
differences in the shape and venation of the
elytra, shape of the head and pronotum, sculp-
ture of the head, antennae and pronotum, and
also in the length of the prosternum (for more
details see also the Notes to the description of
the subgenus Cainomerga subgen.n.).
Diagnosis. Body elongate oval to rather
elongate of small to medium size (3.0–19.0
mm). Integument with moderately coarse sculp-
ture and punctation; large suboval, subpolygo-
nal or somewhat transverse cells of elytra ar-
ranged into ten or more long longitudinal rows
interspaced by weak veins of more or less sub-
equal development. Head usually subtriangular
(to subquadrangular or suboval), with more or
less projecting temples; dorsal surface with weak
or not raised tubercles and longitudinal convex-
ities (usually nearly obliterated), antennal inser-
tions open (not covered by tubercles) or slightly
covered by tubercles. Mandibles moderately to
80
A.G. Kirejtshuk et al.
rather strongly developed, and arcuately to an-
gularly curved at outer edge before stout bi-
dentate apex. Antennae subfiliform and moder-
ately long; antennomere 1 (scape) and other
antennomeres subcylindrical to slightly sub-
conical, various proportions or scape some-
times swollen. Pronotum diverse in shape, with
gently sloping to somewhat explanate sides,
anterior angles frequently not projecting, sur-
face usually tuberculate and with weak median
carina. Part of prosternum before procoxae usu-
ally much longer than procoxae, although it in
subgenus Mesocupes s.str. more or less shorter.
Elytra with more or less arcuate lateral edges,
subacute apices and usually gently sloping at
sides; A1 usually distinct along entire length,
other main veins, if distinct, mostly clear in
basal half and distally frequently becoming ob-
solete and without clear trace of fusion. Abdom-
inal ventrite 1 clearly longer than each of ven-
trites 2–4. Legs moderately thin, frequently com-
paratively long, with straight protibiae and
narrow tarsi. Aedeagus of Cupes-type with wide
flattened parameres.
Comparison. This genus is the most diverse
of the subfamily, but it can be distinguished
after the characters used in the above key to
genera and subgenera. The genus Mesocupes
s.lato is somewhat similar to Latocupes, but
differs from it in the comparatively narrowly
separated antennal insertions, not strongly trans-
verse pronotum without widely explanate sides,
not or only slightly transverse elytral cells, the
absence of fusion of primary veins at the elytral
apices, the lack of clear Y-shaped depression on
the frons and the short antennomere 2. The
comparatively large eyes of some Mesocupes
(particularly in M. (Mesocupes) spinosus
comb.n.) are known also in members of Cupop-
sis gen.n. and Menatops gen.n., however, the
mentioned species of Mesocupes differs from
Cupopsis svitkoi comb.n. in the greater trans-
verse, but not elongate diameter of eyes, not
concave frons along the anterior portion of the
eyes, the more widely separated antennal inser-
tions and the lack of pair of strong tubercles in
anterior part edge of frons partly covering an-
tennal insertions; it differs from Menatops in
the more or less smaller eyes with great distance
between them and the absence of fusion be-
tween of primary veins on the elytra, but also
from M. orbiculatus comb.n. in the not strongly
transverse head, not strongly transverse prono-
tum (less than twice as wide as long). In addi-
tion, Mesocupes s.lato can be distinguished
from Chalepocarabus — by the transverse
pronotum, the markedly longer antennae, the
usually wider body with the not strongly acumi-
nate elytral apices and also by the larger cells
arranged in the clearly expressed rows on the
elytra; from Cupidium — by the longer proster-
num before the procoxae and the protibia not
curved at apex; from Kirghizocupes — by the
longer antennae, the head with lack of both
raised tubercles over the antennal insertions and
Y-shaped depression on the frons, and also by
the gently sloping elytral sides; from Miocupes
— by the much shorter temples, not parallel
veins and the rows of cells on the elytra with the
larger cells; from Priacmopsis — by the much
smaller body size, the longer abdominal ventrite
1, weak (if raised) tubercles on the dorsal sur-
face of the head and the longer metacoxae; from
Gracilicupes — by the head wider than protho-
rax and with the lack of raised tubercles on the
dorsal surface, the usually longer prosternum
before the procoxae, the shorter sclerites of the
legs and particularly by the structure of aedea-
gus; from Priacma — by the longer abdominal
ventrite 1, weak primary veins on elytra without
fusion at apices and particularly by the structure
of the aedeagus; from Furcicupes — by the
usually much longer prosternum before procox-
ae; from Apriacma gen.n. and Cretomerga gen.n.
— by at least the ten long longitudinal rows of
cells and the absence of fusion of primary veins
on the elytra, the usually gently sloping elytral
sides and usually lack of Y- or V-shaped depres-
sion on the head, and from Cretomerga gen.n.
also by the longer prosternum before the nar-
rowly separated procoxae. Besides, this genus
is very distinct from the recent and fossil genera
and subgenera with the strongly raised primary
veins on the elytra (Adinolepis, Ascioplaga,
Cupes, Paracupoides subgen.n., Taxopsis
gen.n., Prolixocupes and Rhipsideigma) by the
widely separated antennal insertions, the ab-
sence of strongly raised upper tubercles on the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
81
head, the usually more robust body, the absence
of fusion of primary veins on the elytra and the
usually gently sloping elytral sides; also from
Adinolepis, Ascioplaga, Cupes, Prolixocupes,
Rhipsideigma and Taxopsis gen.n. — by the 10
long longitudinal rows of cells on elytra; also
from Paracupoides subgen.n. — by the sub-
acute or rounded (not truncate) elytral apices.
Finally, Mesocupes differs from Paracupes s.lato
in the weak tubercles (if raised) over the more
widely separated antennal insertions and the sub-
acute or rounded (not truncate) elytral apices.
Notes on variability. This genus is very
diverse and shows a mosaic distribution of dif-
ferent characters. Here it is regarded as divided
into three subgenera, but hiati between them are
not always clear. Besides, the head shows a
variabily in many characters analogous to those
in Cupes, but the upper tubercles on the head
usually are not so raised, or if the tubercles over
the antennal insertions or in the distal half are
present, they are apparently more or less weak.
The shape of the pronotum among representa-
tives of this genus is rather variable; however
the ground plan of this group is quite character-
istic of the Cupedinae with the rather convex
disk, the more or less raised median carina and
the gently sloping to (sub) explanate sides. The
length of the abdominal ventrite 1 is also vari-
able, although it is always more or less longer
than each of ventrites 2–4 and not more than
twice as long as ventrite 2. The elytra of the
congeners are usually gently sloping at sides but
in Mesocupes (Anaglyphites) capitatus (Pono-
marenko, 1966), comb.n., they demonstrate a
rather sharp inflection to the steep lateral slopes.
Subgenus Mesocupes s.str.
Composition. This subgenus includes, ex-
cept the type species (Mesocupes primitivus),
M. (Mesocupes) bidens Ponomarenko, 1964;
M. (M.) minor Ponomarenko, 1968; M. (M.)
paulus (Ponomarenko, 1964), comb.n. [Anag-
lyphites]; M. (M.) spinosus Ponomarenko, 1964;
M. (M.) zherikhini (Soriano et Delclós 2006),
comb.n. [Anaglyphites].
Locality and stratigraphy. For Martynov’s
and Ponomarenko’s species: Karatau; Mikhai-
lovka, Galkino, Tchokhaj, Kara-Bas-Tau, right
bank of Kashkar-Ata River valley, outcrops
near village Uspenovka (Uspenskoye), Kara-
Tau Range, Algabass District, Chimkent Re-
gion, Kazakhstan; late Jurassic, Oxfordian (or/
and Kimmeridgian), Karabastau Formation; for
Soriano’s and Delclós’ species: Spain, Cuenca,
4 km NE from village Cierva, Las Hoyas; early
Cretaceous, Barremian.
Diagnosis. Body elongate oval of medium
size (6.4–8.4 mm). Elytra with usually rather
large suboval to polygonal cells arranged into
10 or 11 long longitudinal rows (sometimes
rows are not clearly expressed) interspaced by
very weak veins partly fused at apex, at least A1
and CuA fused. Head usually subtriangular and
moderately narrowing anteriorly, without ex-
pressed Y- or V-shaped depression, well pro-
jecting temples. Antennae subfiliform and mod-
erately long (reaching body midlength); anten-
nomere 1 (scape) usually longer than following
antennomeres; antennomere 2 (pedicel) and fol-
lowing flagellomeres comparable in length.
Pronotum with arcuate sides, its anterior and
posterior edges slightly convex to subrectilin-
ear, surface more or less obliterated at disk,
anterior angles rounded. Prosternum before pro-
coxae about as long as procoxae or somewhat
longer. Elytra somewhat less than twice as long
as wide combined or slightly longer, with broadly
arcuate lateral edges, arcuately narrowing to
conjointly subacute apices and apparently gen-
tly sloping sides; Abdominal ventrite 1 clearly
longer than each of ventrites 2–4 and hypopy-
gidium as long as ventrite 1 or longer and
rounded at apex.
Comparison. See below the Comparison to
the subgenus Cainomerga subgen.n.
Mesocupes (Mesocupes) primitivus Mar-
tynov, 1926 Figs 1–4.
Mesocupes primitivus Martynov, 1926: 6; Ponomar-
enko, 1964: 58 (designation of neotype and redescription).
Neotype. “PIN-2452-682” designated by
Ponomarenko (1964: 58), male, print of dorsal
surface of the complete specimen with traced
outlines of the most ventral sclerites and the
exposed aedeagus, but with missing a part of
antennomeres, right eye, part of protibiae, pro-
tarsi, left mesotibia and mesotarsus, right distal
mesotarsomeres, and right posterior leg.
82
A.G. Kirejtshuk et al.
Notes. Since the holotype of this species had
been lost, in order to provide a reliable typifica-
tion, a neotype was designated and some correc-
tions to the description were made (Ponomaren-
ko, 1964). The holotype was somewhat larger
(length 7.5 mm) than the neotype. However,
Ponomarenko (1969) studied many specimens
of this species and mentioned its body size
ranging 6.6–7.5 mm. This species seems to have
shown some variability in shape of the prono-
tum. Therefore, the neotype is characterized by
the pronotum somewhat different from what
was drawn by Ponomarenko (1968: 59, fig. 4a,
1969: 108, fig. 58a). Ponomarenko also men-
tioned that the pronotum of this species was
twice as wide as long (Ponomarenko, 1969:
108), yet both in his drawing and in the drawing
of the original description the pronotum was
depicted not so wide.
Locality and stratigraphy. Karatau;
Mikhailovka, Galkino, Tchokhaj, Kara-Bas-
Tau, right bank of Kashkar-Ata River valley,
outcrops near village Uspenovka (Uspenskoye),
Kara-Tau Range, Algabass District, Chimkent
Region, Kazakhstan; late Jurassic, Oxfordian
(or/and Kimmeridgian), Karabastau Formation.
Comparison. This species is characterized
by the somewhat “generalized” appearance and
has a medium body size; the pronotum subqua-
drangular with somewhat arcuate sides, the dis-
tinct posterior angles and the rounded anterior
angles. It differs from Mesocupes (Mesocupes)
bidens Ponomarenko, 1964 in the narrower
pronotum with the widely rounded anterior an-
gles; from M. (M.) minor Ponomarenko, 1968 in
the larger (longer and wider) pronotum (mark-
edly wider than head) and the conjointly sub-
acute elytra apices; from M. (M.) paulus (Pono-
marenko, 1964), comb.n. [Anaglyphites] in the
longer pronotum with the distinct posterior an-
gles (not rounded) and the longer antennae;
from M. (M.) spinosus Ponomarenko, 1964 in
the widely rounded anterior angles of the prono-
tum and the shorter antennae; from M. (M.)
zherikhini (Soriano et Delclós, 2006), comb.n.
[Anaglyphites] in the longer antennae, the dis-
tinct posterior angles of the pronotum and the
longer abdominal ventrite 1.
Addition to the description. Length 6.4
mm. Shape of head and pronotum somewhat
different from the drawing in the redescription
by Ponomarenko (1964): head wider with larger
eyes (right eye missing and left eye shifted
posteriorly), temples gently narrowing posteri-
orly (not projecting outwards as in redescrip-
tion), dorsal surface slightly tuberculate, anten-
nal insertions apparently open and moderately
separated; pronotum about 1.5 times as wide as
long, with arcuate sides shallowly sinuate at
posterior angles, distinct posterior angles and
rounded. Elytra apparently with more than 10
long longitudinal rows of more or less oval cells.
Prosternum before procoxae about as long as
procoxae (procoxae located at middle of pro-
thorax). Abdominal ventrites overlapping. Ex-
posed aedeagus including heavily sclerotised
penis trunk with long subapical window and less
heavily sclerotised and rather wide flattened
parameres with apices curved inwards.
Cainomerga subgen.n.
Type species Mesocupes (Cainomerga)
brevicornis sp.n., fossil.
Etymology. The name of this new subgenus
is formed from the name of geological era Cen-
ozoic (= Cainozoic from Greek καινός kainos
“new”, and ζωή zoe “life”) and generic root
“merga” (“Bimerga”, “Merga”, “Tenomerga”,
“Cretomerga”). Gender feminine.
Composition. Except the type species, M.
(C.) immaculatus (Piton, 1940), comb.n., M.
(C.) fraternus sp.n., M. (C.) palaeocenicus sp.n.,
and M. (C.) ponti sp.n.
Olympus SCX9 с камерой Olympus; 6 — тело; снизу, прерывистой линией показаны очертания
шовных краев и жилок надкрылий; пунктиром показаны верхние структуры головы (бугорки на
голове); 7 — сухой отпечаток, сфотографированный на микроскопе Nikon SMZ1500 с Microscope
Eyepiece Camera 9.0MP Aptina Color CMOS MU900; 8 — голова, сфотографированная на растровом
электронном микроскопе (SEM); 9 — вершина брюшка, сфотографированная на растровом элект-
ронном микроскопе (SEM). Длина экземпляра 18,7 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
83
Figs 5–9. Mesocupes (Cainomerga) immaculatus (Piton, 1940), comb.n. (holotype “MNHN R07012”);
Paleogene, Paleocene; France, Menat: 5 — dry print, photographed under stereomicroscope Olympus
SCX9; 6 — body, ventral, with broken lines for outlines of sutural edges and veins of elytra, and dotted lines
outlines of dorsal structures of head (tubercles on head); 7 — dry print, photographed under stereomicroscope
Nikon SMZ1500 with Microscope Eyepiece Camera 9.0MP Aptina Color CMOS MU900; 8 — micrograms
of head, photographed under scanning electron microscope (SEM); 9 — micrograms of abdominal apex,
photographed under scanning electron microscope (SEM). Length of specimen 18.7 mm.
Рис. 5–9. Mesocupes (Cainomerga) immaculatus (Piton, 1940), comb.n. (голотип “MNHN R07012”);
палеоген, палеоцен; Франция, Мена: 5 — сухой отпечаток, сфотографированный на микроскопе
84
A.G. Kirejtshuk et al.
Diagnosis. Body elongate oval of compara-
tively large size (16.5–18.5 mm). Integument
with moderately coarse sculpture and puncta-
tion; large subquadrangular to somewhat trans-
verse, arranged into 10 or 11 long longitudinal
rows interspaced by weak secondary and slight-
ly raised primary veins; A1 apparenly rectilin-
early reaching elytral apex but without fusing
with CuA and other primary veins before apex.
Head suboval (transverse or elongate), with
moderately large eyes and temples; dorsal sur-
face without tubercles; antennal insertions mod-
erately widely separated and open. Mandibles
moderately or weakly raised and arcuately curved
at outer edge. Antennae subfiliform and com-
paratively short (reaching elytral shoulders) to
moderately long (reaching body midlength);
antennomere 1 (scape) and other antennomeres
various in shape and length. Pronotum rather
various in shape, its anterior and posterior edge
subreclilinear or convex, more or less obliterat-
ed at disk and with median elevated carina and
somewhat subexplanate at sides, anterior and
postgerior angles with distinct top and some-
times projecting or rounded. Part of prosternum
before procoxae about as long or slightly longer
than procoxae. Elytra about or somewhat more
than twice as long as wide combined, with
broadly arcuate lateral edges, arcuately narrow-
ing to conjointly subsubacute, gently sloping
sides and (sub) explanate along edges, primary
veins weakly raised, CuA and A1 fused at apex
and cells are more or less transverse. Abdomi-
nal ventrite 1 much longer than each of ventrites
2–4 or in M. (C.) fraternus sp.n. and M. (C.)
immaculatus comb.n. with ventrites 1–3 more
or less comparable in length and hypopygidium
as long as ventrite 1. Legs rather thin, compar-
atively long, apparently with straight protibia
and more or less widened tarsi.
Comparison. This new subgenus can be
diagnosed after the above key to genera and
subgenera. In stead of a considerable variability
of the included species in different structures,
they can be united in one taxon thanks to their
general appearance, the shape of the elytra and
the abdominal ventrites, and also thanks to the
character of the elytral venation and the cells. It
is rather similar to species of Mesocupes s.str.
by many characters, but differs from the latter in
the usually rather slender and larger body, and
the particularly subquadrangular to transverse
cells in the longitudinal rows of the elytra.
Cainomerga subgen.n. differs from the subge-
nus Anaglyphites stat.n. in the larger body with
the narrower elytra, the transverse or subqua-
drangular cells and the more clear primary veins
on the elytra, and also in the absence of Y-
shaped depression on the frons. Besides, spe-
cies of the new subgenus, in contrast to those in
other subgenera, are characterisedwith the com-
paratively long and narrow legs, and also all
members of Cainomerga subgen.n. have been
collected in outcrop from the Paleocene while
those of Mesocupes s.str. and Anaglyphites stat.n.
are known only from the early Cretaceous and
late Jurassic.
Some members of this subgenus seem to
have almost narrowest and longest legs among
cupedines (only the legs of Gracilicupes are
comparable) and differ from other cupedine
genera with 10 and more long longitudinal rows
of cells on elytra in the following characters:
– Chalepocarabus in the much larger body
with the less arcuate elytral sides, the expressed
veins and the transverse or subquadrate cells on
elytra, the shorter and somewhat transverse
pronotum;
– Cupidium in the much longer prosternum
before the procoxae and the conjointly subacute
elytral apices;
– Gracilicupes in the primary veins without
clear fusion at apices, the smaller eyes, the
absence of Y-shaped depression on the frons,
the more or less longer prosternum before the
procoxae, the pronotum markedly wider than
head and the distinct anterior and posterior
angles, and also in the usually much longer
abdominal ventrite 1;
– Kirghizocupes in the primary veins with-
out clear fusion at apices, the shorter head (not
so narrow and elongate), the absence of Y-
shaped depression on the frons, the pronotum
not so wider than head, and the not so acuminate
elytral apices which are conjointly ended (not
separately), and the frequently much longer
abdominal ventrite 1;
Taxonomy of the reticulate beetles of the subfamily Cupedinae
85
– Menatops gen.n. in the larger body, the
primary veins without clear fusion at apices, the
longer head with the larger eyes and the moder-
ately long anterior frontal part, the narrower and
longer elytra with the more arcuate sides and the
more narrowing apices, and also in the usually
much longer abdominal ventrite 1;
– Paracupes s.str. and Paracupoides
subgen.n. in the somewhat curved ouline of the
body, the primary veins without clear fusion at
apices, the absence of tubercles on the frons, the
open antennal insertions, the shorter proster-
num, the conjointly acute elytral apices (not
truncate) and the not curved protibiae;
– Priacma in the somewhat curved outline
of the body, the primary veins without clear
fusion at apices, the absence of convexities on
the frons, the open antennal insertions, the shorter
prosternum, the conjointly acute elytral apices
(not truncate), and the longer abdominal ven-
trite 1.
Cainomerga subgen.n. differs from the re-
cent taxa with nine long longitudinal rows on the
elytra, not only in the number of these rows, but
also in the somewhat curved ouline of the body,
the primary veins without clear fusion at apices,
the absence of very prominent convexities on
the frons, the frequently shorter prosternum
before the procoxae, and the gently sloping
elytral sides. Also Cainomerga subgen.n. dif-
fers from the fossil taxa with nine long longitu-
dinal rows on the elytra in the number of these
rows and also in the following characters:
– Apriacma gen.n., Cupes and Furcicupes in
the somewhat curved ouline of the body, the
primary veins without clear fusion at apices, the
frequently shorter prosternum before the pro-
coxae and the longer abdominal ventrites; and
also from from Apriacma gen.n. and Furci-
cupes in the absence of Y-shaped depression on
the frons; and also from Cupes also in the
absence of very prominent convexities on the
frons and the straight protibia;
– Cretomerga gen.n. in the primary veins
without clear fusion at apices, the more oval
head with the wider neck, the absence of Y-
shaped depression on the frons and the narrowly
separated procoxae;
– Cupopsis gen.n. in the head with the small-
er eyes and not concave along the anterior part
of their edges;
– Latocupes (? =Pulchicupes) in the more
slender body, the primary veins without clear
fusion at apices, the less transverse cells on the
elytral disk, and the absence of Y-shaped de-
pression on the frons;
– Miocupes in the more slender body, the
usually narrow and the not transverse head with
the not strongly projecting temples, the longer
pronotum, the shorter prosternum before the
procoxae, and also in the not parallel primary
veins and the more distinct cells on the disk of
the elytra;
– Taxopsis gen.n. in the smaller body-size,
the less transverse cells on the elytral disk,
narrow prosternal process, and the primary veins
without clear fusion at apices.
Finally, Cainomerga subgen.n. differs from
the forms with unclear number of the long lon-
gitudinal rows of cells, namely from Priacmop-
sis in the smaller and much more slender body,
and the usually shorter prosternum before the
procoxae, and from Cupidium in the more slen-
der body, and the conjointly acute elytral apices
(not separately rounded).
Mesocupes (Cainomerga) brevicornis sp.n.
Figs 18–26.
Holotype. “MNHN.F.A51119”, print and
counterprint. The holotype is with somewhat
smoothed outlines of sclerites (not clearly visi-
ble most sclerites), although the elytral sculp-
ture is more or less clear, with missing 3–11
segments of one antenna, one anterior and one
posterior tibiae, almost all tarsomeres, the apex
of one elytron and the apex of the abdomen; the
prothoracic segment with an angular break. The
rock has been collected by A. Nel in the histor-
ical site of the ancient quarry of “Noir
d’Auvergne” where Piton investigated in the
years 1935–1940. The piece of rock is compar-
atively hard and slightly becoming loose.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
86
A.G. Kirejtshuk et al.
Figs 10–11. Menatops orbiculatus (Kirejtshuk, Nel et Collomb, 2010), comb.n. (holotype “MNT-05-198”,
dry print and counterprint, photographed under Olympus SCX9 with camera Olympus); Paleogene,
Paleocene; France, Menat: 10 — print; 11 — counterprint. Length of specimen 9.1 mm.
Рис. 10–11. Menatops orbiculatus (Kirejtshuk, Nel et Collomb, 2010), comb.n. (голотип “MNT-05-198”,
сухие отпечаток и противоотпечаток; фотографии, выполненные на микроскопе Olympus SCX9 с
камерой Olympus); палеоген, палеоцен; Франция, Мена: 10 — отпечаток; 11 — противоотпечаток.
Длина экземпляра 9,1 мм.
Etymology. The epithet of this new species
is composed from the Latin “brevis” (short,
small) and “cornu” or “cornus” (beak, horn,
antler etc.) and referred to its short antennae.
Diagnosis. This new species is very well
characterized by its short antennae and the pe-
culiar pronotum with the subtruncate narrow
anterior edge and the arcuate sides. It can be
diagnosed among the species known from Menat
after the key to species of the subgenus (see
below). In addition to this key, M. (C.) brevicor-
nis sp.n. also differs from M. (C.) fraternus sp.n.
Рис. 12–17. Menatops orbiculatus (Kirejtshuk, Nel et Collomb, 2010), comb.n. (голотип “MNT-05-198”,
сухой отпечаток); палеоген, палеоцен; Франция, Мена: 12 — голова и проторакс, сфотографирован-
ные на микроскопе Olympus SCX9 с камерой Olympus; 13 — голова и переднеспинка, сверху;
прерывистой линией очерчен ментум; 14 — тело с усиками, сфотографированное на растровом
электронном микроскопе (SEM); 15 — метавентрит и средние бедра, сфотографированные на
растровом электронном микроскопе (SEM); 16 — тело без усиков, сфотографированное на растро-
вом электронном микроскопе (SEM); 17 — голова и переднегрудь, сфотографированные на растро-
вом электронном микроскопе (SEM). Длина экземпляра 9,1 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
87
Figs 12–17. Menatops orbiculatus (Kirejtshuk, Nel et Collomb, 2010), comb.n. (holotype “MNT-05-198”,
dry print); Paleogene, Paleocene; France, Menat: 12 — head and prothorax, photographed under Olympus
SCX9 with camera Olympus; 13 — head and pronotum, dorsal, with broken line outlined mentum; 14 —
micrograms of body with antennae, photographed under scanning electron microscope (SEM); 15 —
micrograms of metaventrite with mesofemora, photographed under scanning electron microscope (SEM);
16 — micrograms of body without antennae, photographed under scanning electron microscope (SEM);
17 — micrograms of head and prothorax, photographed under scanning electron microscope (SEM). Length
of specimen 9.1 mm.
88
A.G. Kirejtshuk et al.
Figs 18–21. Mesocupes (Cainomerga) brevicornis sp.n. (holotype “MNHN.F.A51119”, dry print,
photographed under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 18 —
print; 19 — body, ventral, with broken line outlined sutural edges of elytra, edges of laterosternites and
structures on upper surface of head (tubercles), and also dotted lines outlined traced edges of scape, procoxae,
protrochanter and tarsi; 20 — head and prothorax; 21 — left antenna. Length of specimen 18.3 mm.
Рис. 18–21. Mesocupes (Cainomerga) brevicornis sp.n. (голотип “MNHN.F.A51119”, сухой отпечаток,
сфотографированный на микроскопе Olympus SCX9 с камерой Olympus); палеоген, палеоцен;
Франция, Мена: 18 — отпечаток; 19 — тело, снизу; прерывистой линией очерчены шовные края
надкрылий, края латеростернитов, верхних структур головы (бугорки); пунктиром показаны просле-
живающиеся очертания скапуса, прококсы переднего вертлуга и лапок; 20 — голова и переднегрудь;
21 — левый усик. Длина экземпляра 18,3 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
89
Figs 22–26. Mesocupes (Cainomerga) brevicornis sp.n. (holotype “MNHN.F.A51119”); Paleogene,
Paleocene; France, Menat: 22 — body of dry counterprint, photographed under Olympus SCX9 with camera
Olympus; 23 — apex of left elytron of dry counterprint under Olympus SCX9 with camera Olympus; 24 —
head and prothorax of dry counterprint, photographed under Olympus SCX9 with camera Olympus; 25 —
micrograms of body of print, photographed under scanning electron microscope (SEM); 26 — micrograms
of head of print, photographed under scanning electron microscope (SEM). Length of specimen 18.3 mm.
Рпс. 22–26. Mesocupes (Cainomerga) brevicornis sp.n. (голотип “MNHN.F.A51119”); палеоген, палео-
цен; Франция, Мена: 22 — тело сухого противоотпечатка, сфотографированное под микроскопом
Olympus SCX9 с камерой Olympus; 23 — вершина левого надкрылья сухого противоотпечатка,
сфотографированная под микроскопом Olympus SCX9 с камерой Olympus; 24 — голова и переднег-
рудь сухого противоотпечатка, сфотографированные под микроскопом Olympus SCX9 с камерой
Olympus; 25 — тело отпечатка, сфотографированное на растровом электронном микроскопе (SEM);
26 — голова отпечатка, сфотографированная на растровом электронном микроскопе (SEM). Длина
экземпляра 18,3 мм.
90
A.G. Kirejtshuk et al.
in the sparse sculpture of the underside, the
shorter scape and particularly following anten-
nomeres, the shorter legs, apparently the longer
metacoxae and the much longer abdominal ven-
trites; from M. (C.) immaculatus comb.n. in the
rather narrowly separated mesocoxae, some-
what arcuate elytral sides, the longer all abdom-
inal ventrites (particularly the hypopygidium),
the much denser and finer sculpture of the un-
derside integument; from M. (C.) palaeoceni-
cus sp.n. in the oblique metacoxae, the longer
each of abdominal ventrites; from M. (C.) ponti
sp.n. in the longer prosternum before the pro-
coxae, the more narrowly separated mesocoxae,
the somewhat longer abdominal ventrite 1 and
hypopygidium.
Description. Body length (with elytra) 18.3
mm, width apparently 6.5 mm, elytral length 9.1
mm. Body elongate; apparently slightly to mod-
erately convex dorsally and subflattened ven-
trally. Integument of head and pronotum with
moderately large regular punctures separated
by about one puncture diameter or less; metaven-
trite and many other underside sclerites with
comparably fine and moderately sparse punc-
tures (interspaces between punctures about as
great as puncture diameter); elytra with large
transverse to somewhat subpolygonal cells ar-
ranged in at least 10 (probably 11) longitudinal
rows between veins.
Head subpentagonal (probably because of
fossilization) and without traces of upper tuber-
cles, somewhat longer than wide, apparently
widest at temples (rather projecting and suban-
gular), with moderately large oval eyes; anten-
nal insertions open. Mandibles apparently mod-
erately developed, slightly exposed beyond la-
brum, with gently curved outer edge. Antennae
comparatively short; scape longest and rather
thick; antennomere 2 (pedicel) subcylindrical,
somewhat shorter than flagellomeres, narrower
than scape and wider than flagellomeres;
flagellomeres subequal in length and ultimate
one acuminate. Prothorax moderately short;
pronotum about one and 1/3 as wide as long,
apparently slightly and gently convex, its sides
seemingly widely (sub) explanate and slightly
narrowing anteriorly, its sides arcuate, anterior
edge subconcave, anterior and posterior angles
stump and with distinct top; prosternum before
procoxae at least as long as procoxae, procoxae
apparently narrowly separated. Mesoventrite
apparently about as long as prosternum. Meso-
coxae moderately large and distance between
them about 1/4 of their diameter. Metaventrite
about half as long as wide at base. Metacoxae
rather oblique, about three times as wide as long
and their mesal parts moderately projecting
posteriorly. Abdomen with overlapping ven-
trites; ventrite 1 longest (nearly twice as long as
following ones), ventrites 2–4 subequal in length
and much shorter than hypopygidium (ventrite
5); hypopygidium apparently rounded at apex.
Elytra slightly wider than prothorax, with broadly
arcuate sides, together more than twice as long
as wide, apparently moderately convex; with
slightly expressed veins.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, about three times as long as wide. Tibiae
apparently somewhat shorter than femora and
very thin. Tarsi represented by short traces of
one fore and both intermediate legs.
Mesocupes (Cainomerga) fraternus sp.n.
Figs 139–141.
Holotype. “MNHN.F.A52773” (“NEL
1452”), print. The specimen has the more or less
clear outlines of its legs and abdominal ven-
trites, but the outlines of the thoracic sclerites
and antennomeres are rather obscure, and also
the terminal antennomeres and apical part of
abdomen are missing. The specimen originated
from a new, small outcrop near the south-east of
the village of Menat, the piece of rock is rather
soft and fragile.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The epithet of this new species
(Latin “brotherly”, “fraternal”) refers to a con-
siderable similarity of it to Mesocupes (Caino-
merga) brevicornis sp.n.
Diagnosis. This new species is character-
ised by the shortest frons within its consubgen-
ers, the densest sculpture of the underside integ-
ument, the comparatively long abdominal ven-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
91
trites 2–4 and the comparatively large eyes. It
has the similar outlines of the katepisternal
sutures and the metacoxae to those in M. (C.)
brevicornis sp.n., although the metacoxae of the
latter apparently are longer. However its under-
side integument is much more densely sculp-
tured, the legs and antennae are somewhat long-
er, the abdominal ventrite 1 is markedly shorter
and the pronotum is apparently widening ante-
riad. The particular differences of these species
are in the shape of their head, the temples and
size of eyes, although these differences could
partly be caused by peculiar circumstances in
fossilization. Mesocupes (C.) fraternus sp.n.
can be diagnosed after the below key to species,
Besides, this new species differs from M. (C.)
immaculatus comb.n. in the longer abdominal
ventrites, the gular sutures divergent in the pos-
terior part, the much finer and rather dense
sculpture of the underside; from M. (C.) palae-
ocenicus sp.n. in the much larger eyes, the
longer legs, the shorter metacoxae and the much
shorter abdominal ventrite 1; from M. (C.) ponti
sp.n. in the much larger eyes, the oblique and
shorter metacoxae.
Description. Body length (without apical
parts of abdomen and elytra) 14.6 mm, width
apparently 6.3 mm. Integument of underside
with rather fine and very dense regular (some-
where subcontiguous) punctures; metaventrite
with comparably fine and sparse punctures;
elytra with large transverse to subpolygonal
cells arranged longitudinal rows between veins.
Head transversally subtriangular (probably
because of fossilization) widely separated gular
sutures, apparently widest at temples (some-
what projecting and rounded), apparently with
comparatively large oval eyes; antennal inser-
tions apparently open. Mandibles comparative-
ly small, slightly exposed beyond labrum, with
gently curved outer edge. Antennae moderately
long; scape longest and rather thick; antenno-
mere 2 (pedicel) subcylindrical, somewhat short-
er than flagellomeres, narrower than scape and
wider than flagellomeres; flagellomeres 4-7
subequal in length, each of somewhat shorter
than scape and somewhat longer than each of
antennomeres 2 and 3. Prothorax rather short;
its sides widening anteriad, anterior prosternal
edge subconcave, anterior and posterior angles
rather acute and with distinct top; prosternum
before procoxae at least as long as procoxae.
Mesoventrite apparently about as long as pros-
ternum. Mesocoxae not visible. Metaventrite
about 2/3 as long as wide at base. Metacoxae
slightly oblique, their mesal parts modeartely
projecting posteriorly. Abdomen with overlap-
ping ventrites; ventrites 1–3 subequal in length.
Elytra somewhat wider than prothorax, appar-
ently with broadly arcuate sides, with slightly
expressed veins.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, about four times as long as wide. Tibiae
apparently somewhat shorter than femora and
very thin. Tarsi represented by moderately long
and widened tarsomeres.
Mesocupes (Cainomerga) immaculatus (Pi-
ton, 1940), comb.n.
Figs 5–9.
Zonabris immaculatus Piton, 1940: 194 (Meloidae
Gyllenhal, 1810).
Holotype. “MNHN R07012”, “Zonabris im-
maculatus Piton, TYPE”, male, print of the
entire body with a mixture of outlines of both
dorsal and ventral sclerites, and also with un-
clear remains of the sculpture of the integument
and the exposed aedeagus, but with missing
distal antennomeres, all tarsi (except right meta-
tarsus), anterior part of protibia, right mesotibia
and left metatibia. The specimen is entombed in
the rock collected by Louis Piton, in the histor-
ical site of the ancient quarry of “Noir
d’Auvergne” where Piton investigated in the
years 1935–1940, this piece of rock is compar-
atively smooth and rather consistent.
Notes. The holotype of this species demon-
strates all features of Cupedidae, but not Me-
loidae where it was initially put (Piton, 1940). It
is here considered as belonging to the Cenozoic
group of the genus Mesocupes. This interpreta-
tion is confirmed also by the structure of its
aedeagus similar to that of the type species of
this genus. The elytral cells of the holotype have
unclear and obliterated edges, although some
92
A.G. Kirejtshuk et al.
cells in the lateral longitudinal rows demon-
strate the subquadrate to transversely quadran-
gular outlines.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Diagnosis. This new species is very well
characterized by its comparatively robust body,
subpentagonal pronotum with rounded angles,
very long and rather oblique metacoxae, most
coarse and sparse sculpture of the underside.
The particular features of this species are the
more widely separated mesocoxae, and meta-
coxae nearly reaching posterior edge of abdom-
inal ventrite 1. It can be easily diagnosed among
the species known from Menat after the key to
species (see below). In addition to this key, M.
(C.) immaculatus comb.n. also differs from M.
(C.) brevicornis sp.n. in the subparallel elytral
sides and shorter all abdominal ventrites (par-
ticularly hypopygidium); from M. (C.) frater-
nus sp.n. in the markedly shorter head, shorter
abdominal ventrites, gular sutures convergent
in the posterior part; from M. (C.) palaeoceni-
cus sp.n. in the more widely separated gular
sutures, longer abdominal ventrites 2–4; from
M. (C.) ponti sp.n. in the apparently longer
prosternum before procoxae.
Redescription. Body length (with elytra)
18.7 mm, width 7.8 mm, elytral length 9.1 mm.
Body elongate; apparently slightly to moderate-
ly convex dorsally and sub-flattened ventrally.
Integument of dorsal and ventral sclerites with
moderately coarse sculpture including coarse
and dense oval punctures, interspaces between
them apparently coarsely microreticulate; elytra
with subquadrate to transversely quadrangular
cells arranged in rows between veins (visible
only rows at lateral edge of elytra).
Head subtriangular and with scarcely traced
upper tubercles but with paramedian longitudi-
nal lines, slightly wider than long, widest at
temples, with moderately large oval eyes; an-
tennal insertions open. Mandibles apparently
moderately developed, slightly exposed beyond
labrum, with gently curved outer edge. Anten-
nae apparently moderately long; scape slightly
longer and thicker than following antennomer-
es; antennomere 2 (pedicel) comparable in-
length with basal flagellomeres (preserved left
antennomeres 3 and 4). Prothorax moderately
short; pronotum about one and 1/3 as wide as
long, apparently slightly and gently convex, its
sides seemingly widely (sub) explanate and
slightly wider anteriorly; prosternum before
procoxae at least as long as procoxae, procoxae
apparently narrowly separated. Mesoventrite
apparently about as long as prosternum. Meso-
coxae moderately large and distance between
them about 2/3 of their diameter. Metepisterna
nearly four times as long as wide. Metaventrite
about half as long as wide at base. Metacoxae
moderately oblique, about twice as wide as long
and their mesal parts strongly projecting poste-
riorly. Abdomen with overlapping ventrites;
ventrite 1 longest and nearly as long as hypopy-
gidium and somewhat longer than each of ven-
trites 2–4; pygidium and hypopygidium widely
rounded at apex. Elytra much wider than pro-
thorax, with subparallel sides, together about
2.3 times as long as wide, apparently moderate-
ly convex; R and M somewhat more clear than
other main veins (may be as sequence of preser-
vation) but CuA and A1 very slightly traced.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, 2.5–3.0 times as long as wide, metafem-
ora somewhat longer than pro- and mesofemo-
ra. Tibiae apparently somewhat longer than
femora and very thin. Metatarsus slightly short-
er than metatibia.
Aedeagus with broken apex, slightly sclero-
tized, including of moderately long subparallel-
sided penis trunk and wide flattened parameres
markedly shorter than penis trunk.
Mesocupes (Cainomerga) palaeocenicus
sp.n. Figs 27–31.
Holotype. “MNHN.F.A51117”, “11/05/
1970”, print, collection of Philippe Olivier. The
print of the holotype is with somewhat smoothed
outlines of sclerites (not visible anterior part of
head except apices of the mandibles), with miss-
ing apical atennomeres of one antenna, apex of
one mesotibia, most part of tarsomeres and
Taxonomy of the reticulate beetles of the subfamily Cupedinae
93
Figs 27–31. Mesocupes (Cainomerga) palaeocenicus sp.n. (holotype “MNHN.F.A51117”, dry print,
photographed under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 27 —
body; 28 — mentum and mandibles, ventral; 29 — body, ventral, with broken line outlined sutural edges
of elytra, edges of laterosternites and median carina of pronotum, and also dotted lines outlined traced tarsi;
30 — part of body with elytron; 31 — head and prothoracic segment. Length of specimen 16.5 mm.
Рис. 27–31. Mesocupes (Cainomerga) palaeocenicus sp.n. (голотип “MNHN.F.A51117”, сухой отпечаток,
сфотографированный под микроскопом Olympus SCX9 с камерой Olympus); палеоген, палеоцен;
Франция, Мена: 27 — тело; 28 — ментум и мандибулы, снизу; 29 — тело, снизу; прерывистой линией
очерчены шовные края надкрылий, края латеростернитов, медиальный гребень переднеспинки;
пунктиром показаны прослеживающиеся очертания лапок; 30 — часть тела с надкрыльем; 31 —
голова и переднегрудь. Длина экземпляра 16,5 мм.
94
A.G. Kirejtshuk et al.
elytral apices. The rock has been collected by
Philippe Olivier in the historical site of the
ancient quarry of “Noir d’Auvergne” where
Piton investigated in the years 1935–1940. This
piece of rock is comparatively smooth and slight-
ly becoming loose.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The epithet of this new species
refers to time (epoch) of its origin.
Diagnosis. This new species is very well
characterized by its hexagonal pronotum and
very long antennomere 2. It can be easily diag-
nosed among the species known from Menat
after the key to species (see below). In addition
to this key, M. (C.) palaeocenicus sp.n. differs
from M. (C.) brevicornis sp.n. in the less slender
body, the longer antennae, the narrowly separat-
ed (subcontiguous) mesocoxae, the much short-
er ventrite 1 and hypopygidium; from M. (C.)
fraternus sp.n. in the much longer head with the
much smaller eyes, the shorter legs, the longer
metacoxae, the much longer abdominal ventrite
1 and markedly denser sculpture of the under-
side; from M. (C.) immaculatus comb.n. in the
more narrowly separated gular sutures, the nar-
rowly separated mesocoxae, the shorter abdom-
inal ventrites 2–4, the strongly denser and finer
sculpture of the underside; from M. (C.) ponti
sp.n. in the larger eyes, the longer prosternum
before the procoxae, the longer ventrite 1 and
hypopygidium.
Description. Body length (with elytra) 16.5
mm, width 4.6 mm, elytral length 12.8 mm.
Body elongate and rather slender; apparently
slightly to moderately convex dorsally and sub-
flattened ventrally. Integument of dorsal and
ventral sclerites with moderately coarse sculp-
ture including coarse and dense oval punctures,
very narrow interspaces between them (less
than one puncture diameter) apparently coarse-
ly microreticulate; metaventrite apparently with
somewhat obliterated punctation; elytra with
transverse to subpolygonal cells arranged in at
least 11 rows between veins.
Head subtriangular and with a pair of small
and round upper tubercles at the middle of eyes,
slightly wider than long, widest at angularly
projecting temples, with rather large oval eyes
and with distance between them about as great
as transverse eye diameter; antennal insertions
open. Mandibles apparently moderately devel-
oped and represented in print only by their
apices. Antennae comparatively long and reach-
ing the middle of abdominal ventrite 1; scape
not very thick; antennomere 2 (pedicel) some-
what longer than each of three following
flagellomeres and subequl in length to each of
six apical flagellomeres (antennomeres 6–11).
Gular sutures subparallel to slightly curved in
basal half of epicranium. Prothorax moderately
short; pronotum subhexagonal (widest at the
middle) with distinct anterior and posterior an-
gles and widest at the middle, straight anterior
and posterior edges, about 1.5 times as wide as
long, apparently slightly and gently convex, its
sides seemingly widely (sub) explanate and
slightly wider anteriorly; prosternum before
procoxae at least as long as procoxae and longi-
tudinal stripe along intire length (to apex of
prosternal process), procoxae moderately nar-
rowly separated. Mesoventrite somewhat short-
er than prosternum. Mesocoxae moderately large
and distance between them very narrow.
Metaventrite somewhat longer than half its width
at base. Metacoxae transverse, about twice as
wide as long and their mesal parts strongly
projecting posteriorly. Abdomen with ventrite 1
longest and more than three times as long as
each of ventrites 2–4 and somewhat longer than
hypopygidium (ventrite 5); pygidium and hy-
popygidium widely rounded at apex. Elytra
slightly wider than prothorax, with sub-parallel
sides, somewhat more than twice as long as wide
combined, primary and secondary veins equally
raised.
Femora comparable in shape and proportions
gently curved along anterior and posterior edges,
3.0 times or more as long as wide, metafemora
somewhat longer than pro- and mesofemora.
Tibiae more or less longer than femora and very
thin. Metatarsus comparatively wide.
Mesocupes (Cainomerga) ponti sp.n.
Figs 32–40.
Holotype. “MNHN.F.A51116”, “11/05/
1970, 63-Menat”, print and counterprint, col-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
95
Figs 32–36. Mesocupes (Cainomerga) ponti sp.n. (holotype “MNHN.F.A51116”, dry print and counterprint,
photographed under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 32 —
counterprint, body; 33 — counterprint, head, ventral, with broken line outlined scape; 34 — print, body,
dorsal, with broken line outlined edges of ventral sclerites; and dotted line apparent edges of compressed
head and traced mesotarsus; 35 — counterprint, head and prothoracic segment with appendages; 36 — print,
head and prothoracic segment with appendages. Length of specimen 18.5 mm.
Рис. 32–36. Mesocupes (Cainomerga) ponti sp.n. (голотип “MNHN.F.A51116”, сухие отпечаток и
противоотпечаток, сфотографированные под микроскопом Olympus SCX9 с камерой Olympus);
палеоген, палеоцен; Франция, Мена: 32 — тело противоотпечатка; 33 — голова противоотпечатка,
снизу; прерывистой линией очерчены края скапуса; 34 — тело отпечатка, сверху; прерывистой
линией очерчены нижние склериты; пунктиром возможные края сдавленной головы и прослежива-
ющиеся очертания мезотарсуса; 35 — голова и переднегрудной сегмент противоотпечатка c придат-
ками; 36 — голова и переднегрудной сегмент отпечатка c придатками. Длина экземпляра 18,5 мм.
96
A.G. Kirejtshuk et al.
Figs 37–40. Mesocupes (Cainomerga) ponti sp.n. (holotype “MNHN.F.A51116”, print and counterprint,
micrograms, photographed under scanning electron microscope (SEM)); Paleogene, Paleocene; France,
Menat: 37 — counterprint, body; 38 — counterprint, head; 39 — print, body; 40 — print, head and
prothoracic segment with appendages. Length of specimen 18.5 mm.
Рис. 37–40. Mesocupes (Cainomerga) ponti sp.n. (голотип “MNHN.F.A51116”, отпечаток и противо-
отпечаток, сфотографированные на растровом электронном микроскопе (SEM)); палеоген, палеоцен;
Франция, Мена: 37 — тело противоотпечатка; 38 — голова противоотпечатка; 39 — тело отпечатка;
40 — голова и переднегрудной сегмент отпечатка с придатками. Длина экземпляра 18,5 мм.
lection Philippe Olivier. The holotype has rath-
er smoothed outlines of sclerites (most of them
not clearly visible), and also rows of cells on the
elytra, but missing 4–11 segments of both anten-
nae, tibiae except one intermedial and one pos-
terior tibiae, almost all tarsomeres, apex of one
elytron and apex of abdomen. The rock has been
collected by Philippe Olivier in the historical
site of the ancient quarry of “Noir d’Auvergne”
where Piton investigated in the years 1935–
1940. This piece of rock is comparatively smooth
and slightly becoming loose.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The epithet of this new species
is devoted to Sylvain Pont who assisted the
authors in the studies of beetles with electronic
microscopy.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
97
Diagnosis. This new species is very well
characterized by its peculiar head and prono-
tum, the short prosternum and the transverse
metacoxae, making its identification quite easy
and reliable. It can be easily diagnosed among
the species known from Menat after the below
key to species. In addition to this key as well as
shape of head and prothorax, M. (C.) ponti sp.n.
differs from M. (C.) brevicornis sp.n. in the
longer basal antennomeres and shorter abdom-
inal ventrite 1 and hypopygidium; from M. (C.)
fraternus sp.n. in the longer frons, the somewhat
more sparse sculpture of the underside, the
comparatively short abdominal ventrites 2–4
and the comparatively small eyes; from M. (C.)
immaculatus comb.n. in the slender body, the
shorter and transverse metacoxae ended far from
the posterior edge of the abdominal ventrite 1,
the more narrowly separated mesocoxae, the
dense and finer sculpture of the underside; from
M. (C.) palaeocenicus sp.n. in the much shorter
basal antennomeres and the less narrowly sepa-
rated mesocoxae.
Description. Body length (with elytra) 18.5
mm, width 4.5 mm, elytral length 12.3 mm.
Body elongate; apparently slightly to moderate-
ly convex dorsally and subflattened ventrally.
Integument of dorsal and ventral sclerites with
moderately coarse sculpture including coarse
and very dense oval punctures, interspaces be-
tween them apparently coarsely microreticu-
late; metaventrite with somewhat larger and
shallower punctures; elytra with subpolygonal
to more or less transverse cells arranged in 10
(or could be 11) rows between veins.
Head elongate and with scarcely traced up-
per tubercles, slightly wider than long, widest at
gently rounded temples, with comparatively
small oval eyes at the middle of head; antennal
insertions open. Mandibles apparently well de-
veloped, well exposed beyond labrum, with
gently curved outer edge. Antennae represented
only by three basal antennomeres; scape rather
swollen, much thisker and somewhat longer
than each of antennomeres 2 and 3 with sub-
equal length. Prothorax moderately short; prono-
tum about one and 1/3 as wide as long, apparent-
ly slightly and gently convex, its sides seeming-
ly widely (sub) explanate and slightly wider
anteriorly, from the middle widened to acutely
projecting anterior angles, its anterior edge con-
vex; prosternum before procoxae at least as long
as procoxae, procoxae rather narrowly separat-
ed. Mesoventrite shorter than prosternum. Me-
socoxae moderately large and distance between
them about 1/3 of their diameter. Metaventrite
about half as long as wide at base. Metacoxae
transverse, about twice as wide as long and their
mesal parts strongly projecting posteriorly.
Abdomen with overlapping ventrites; ventrite 1
about as long as hypopygidium and about 1.5
times as long as each of ventrites 2–4; pygidium
and hypopygidium widely rounded at apex.
Elytra apparently slightly wider than prothorax,
with subparallel sides, about 2.3 times as long as
wide combined, apparently moderately con-
vex; Cu and R somewhat more clear than other
main veins.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, about three times as long as wide. Tibiae
apparently as long as femora and very thin.
Mesotarsus slightly shorter than mesotibia.
Mesocupes (Cainomerga) sp.1
Fig. 159.
Specimen examined. “MNHN.F.A.57518”
(“NEL 2200”), print with the incomplete spec-
imen (partly pronotum and distal part of body)
showing rather obliterated and unclear outlines,
deposited in the comparatively soft and fragile
rock. It originated from a new, small outcrop
near the south-east of the village of Menat.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. Body 12.2 mm long, and elytra 10.3
mm long. This specimen is characterised by the
not slender body, the pronotum somewhat sim-
ilar to that in M. (Mesocupes) primitivus with
rounded sides and acuminate posterior angles,
the elytra with longitudinal rows of transverse
subpolygonal cells and comparatively clear pri-
mary veins, and very acute apex of last abdom-
inal ventrite which somewhat longer than wide.
Other characters are not clear, although its meta-
98
A.G. Kirejtshuk et al.
coxae are apparently with transverse anterior
edges, ventrites 2–4 are comparable in length
and each of them slightly longer than ventrite 5.
The outlines of coxae and some of leg sclerites
are very obscure. The available characters are
very distinct from those in all consubgeners here
described, however preservation of the speci-
men and lack of head with appendages make
unreasonable description of a new additional
species after study of this alone specimen.
Mesocupes (Cainomerga) spp.
Figs. 148, 160–162.
Specimens examined. “MNHN.F.A.52769”
(“NEL 1579”), print of separate elytron with the
clear veins and cells deposited in the hard rock.
The specimen originated from a new, small
outcrop near the south-east of the village of
Menat. “MNHN.F.A.57519” (“NEL 2210”),
print and counterprint with the separate elytron,
deposited in the moderately soft and fragile
rock. The specimen originated from a new,
small outcrop near the south-east of the village
of Menat. “MNHN.F.A.57520” (“Oli 685”),
print of apical part of elytron with the clear veins
and cells deposited in the moderately soft and
fragile rock. The rock has been collected by
Philippe Olivier in the historical site of the
ancient quarry of “Noir d’Auvergne” where
Piton investigated in the years 1935–1940.
“MNHN.F.A.57521” (“Oli 314”), print of mid-
dle part of separate elytron, deposited in the
moderately soft and fragile rock. The rock has
been collected by Philippe Olivier in the histor-
ical site of the ancient quarry of “Noir
d’Auvergne” where Piton investigated in the
years 1935–1940.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. “MNHN.F.A. 52769”: Elytron 12.6
mm long and 3.5 mm wide. “MNHN.F.A.
57519”: Elytron 14.6 mm long and 3.9 mm
wide. The complete elytra of these specimens
have the primary veins weakly expressed, fu-
sion of A1 and CuA and 10 longitudinal rows of
distinctly transverse cells and gently sloping
sides. “MNHN.F.A.57520”: Fragment of elytron
4.5 mm long with weak primary vein and trans-
verse cells. “MNHN.F.A. 57521”: Fragment of
elytron 4.1 mm long with the primary veins
weakly expressed and transverse cells.
5.3. Key to species of the subgenus Cain-
omerga subgen.n. from Menat
1. Pronotum widest at strongly projecting anterior
angles; procoxae located at the middle of protho-
rax (i.e. prosternum before procoxae seemingly
as long as procoxae); head only slightly narrow-
ing before eyes; abdominal ventrite 1 about 1.5
times as long as each of ventrites 2–4; antenno-
meres 2 (pedicel) and 3 subequal in length and
somewhat shorter than antennomere 1 (scape);
hypopygidium nearly as long as wide; body
about three times as long as wide; length of body
18.5 mm. Figs 32–40 ....... M. (C.) ponti sp.n.
– Pronotum widest at the middle; procoxae located at
posterior edge of prothorax (i.e. prosternum be-
fore procoxae much longer than procoxae); head
moderately narrowing before eyes; abdominal
ventrite 1 about twice as long as each of ventrites
2–4; combination of remaining characters dif-
ferent ................................................................ 2
2. Pronotum subhexagonal and subangular at widest
place near the middle; antennomere 2 (pedicel)
longer than antennomere 1 (scape) and antenno-
mere 3; head markedly longer than wide; anteri-
or edge of metacoxae transverse (not oblique);
length of body 16.5 mm. Figs 27–31 .............
............................ M. (C.) palaeocenicus sp.n.
– Pronotum not subhexagonal and subangular at
widest place near the middle; antennomere 2
(pedicel) shorter than antennomere 1 (scape) and
antennomere 3; head not longer than wide; ante-
rior edge of metacoxae more or less oblique .
....................................................................... 3
3. Head strongly transverse (nearly twice as wide as
long) with very short frons before eyes; abdom-
inal ventrite 1 only slightly longer than each of
ventries 2 and 3; mandibles slightly exposed;
pronotum widening to acuminate anterior an-
gles; metacoxae short and slightly oblique; length
of body (without apices of elytra and abdomen)
14.6 mm. Figs 139–141 ..................................
.................................... M. (C.) fraternus sp.n.
– Head at least 1.5 times as wide as long; abdominal
ventrite 1 at least 1.5 times as long as each of
ventries 2–4; mandibles usually developed; com-
bination of other remaining characters different
....................................................................... 4
4. Pronotum subpentagonal with subrectilinear sides;
body more robust with abdomen less than 1.5
times as long as wide; head slightly transverse;
Taxonomy of the reticulate beetles of the subfamily Cupedinae
99
metacoxae very long and rather oblique, mesally
reaching posterior edge of abdominal ventrite 1;
hypopygidium transverse; length of body 18.7
mm. Figs 5–9 ..................................................
. M. (C.) immaculatus (Piton, 1940), comb.n.
– Pronotum not subtrapezoid and subarcuate at wid-
est place; body more slender with abdomen
about twice as long as wide; head about as long
as wide; metacoxae moderately long and ob-
lique, mesally reaching the middle of abdominal
ventrite 1; hypopygidium markedly longer than
wide; length of body 18.3 mm. Figs 18–26 ...
................................. M. (C.) brevicornis sp.n.
5.4. (?) Mesocupes (Cainomerga) spp.
(?) Mesocupes (Cainomerga) sp.1.
Fig. 142.
Specimen examined. “MNHN.F.A.52768”
(“Nel 1697”), print. The specimen is represent-
ed by a fragment of elytron with the length 7.0
mm and remains of distal part of the posterior
leg in the piece of comparatively hard rock. The
specimen originated from a new, small outcrop
near the south-east of the village of Menat.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. The print of the elytral apex exam-
ined demonstrates the clear transverse cells in
the longitudinal rows as characteristic of spe-
cies attributed to the subgenus Cainomerga
subgen.n. Nevertheless this elytral apex can
scarcely be associated with any species of this
subgenus because it represents only a third of
the entire elytron (7.0 mm) and respectively the
length of the complete specimen having this
elytral fragment should be at least 25 mm.
Besides, the apex of this elytron is not so narrow
as in the members of Cainomerga subgen.n.
and, therefore, it can be supposed that the beetle
with this elytron could be much more robust
than the cupedines included into this taxon.
(?) Mesocupes (Cainomerga) sp.2.
Fig. 164.
Specimen examined. “MNHN.F.A. 57522”
(“Oli 218”), print. The specimen in lateral posi-
tion is represented by the body with detached
and turned abdomen without clear remains of
antennae and legs, deposited in the piece of
moderately soft and fragile rock. The rock has
been collected by Philippe Olivier in the histor-
ical site of the ancient quarry of “Noir
d’Auvergne” where Piton investigated in the
years 1935–1940.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. The print with the general length of
the specimen in curved position is 12.1 mm and
with elytron 10.0 mm long. It demonstrates the
clear gently sloping elytral sides, weakly raised
primary veins, comparatively large and slightly
transverse cells in the longitudinal rows (appar-
ently ten rows or even 11), head with rather large
eyes and tubercle(s) on its upper surface, com-
paratively wide base of pronotum gradually
narrowing anteriorly, abdomen with subequally
long ventrites and with ventrite 5 slightly trans-
verse and arcuate at apex. The many accessible
features of this print are characteristic of spe-
cies attributed to the subgenus Cainomerga
subgen.n. and gives some emagination on the
lateral appearance of the species of this genus.
Nevertheless the not strongly transverse ely-
tral cells and large eyes raise doubts in such an
attribution.
(?) Mesocupes (Cainomerga) sp.3
Fig. 163.
Specimen examined. “MNHN.F.A. 57523”
(“Oli 674”), print. The specimen represents the
separate elytron. The rock has been collected by
Philippe Olivier in the historical site of the
ancient quarry of “Noir d’Auvergne” where
Piton investigated in the years 1935–1940.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. The elytron 6.4 mm long clearly
shows at least ten longitudinal rows of rather
strongly transverse cells and weakly raised pri-
mary veins. Partly these characters are corre-
spondent to those in members of Cainomerga
subgen.n. However the small size of the elytron
examined make it possible to doubt in such an
attribution.
100
A.G. Kirejtshuk et al.
5.5. Menatops gen.n.
Type species: Cupesorbiculatus Kirejtshuk,
Nel et Colomb, 2010, fossil.
Etymology. The name of this new subgenus
is formed from the name of the locality (Menat)
and an abbreviated generic root “ops” (from the
Greek “opsis”, meaning “view, sight, appear-
ance”). Gender masculine.
Composition. The type species and M. bar-
tenevi sp.n.
Diagnosis. Body elongate oval, of medium
size (9.1 mm). Integument with moderately
coarse sculpture and punctation; large subqua-
drangular to slightly transverse on disk to polyg-
onal or suboval cells along sides of elytra,
arranged into 10 long longitudinal rows inter-
spaced by weak secondary and distinctly raised
primary veins; A1 deviating at scutellum and
rectilinearly reaching elytral apex and fusing
with CuA before apex, and then with M and R.
Head strongly transverse, very short before very
large eyes (with transverse diameter as great as
half of the distance between them), with temples
narrowing behind eyes; upper surface without
tubercles; antennal insertions narrowly separat-
ed and open. Mandibles moderately raised and
arcuately curved at outer edge. Antennae subfil-
iform and moderately long (reaching body
midlength); antennomere 1 (scape) subequal in
length with flagellomeres or slightly longer and
somewhat thicker than other antennomeres.
Pronotum transverse, subhexagonal or subpen-
tagonal, its anterior and posterior edges slightly
convex, more or less smoothed (not tuberculate)
at disk or with a slight median elevated carina
and subexplanate on sides, anterior and poste-
rior angles blunt and not projecting. Part of
prosternum before procoxae markedly longer
than procoxae, the latter located near posterior
edge of pronotum. Elytra about twice as long
as wide combined or 1 and 3/5 as long as wide
combined, with broadly arcuate lateral edges,
arcuately narrowing to nearly conjointly round-
ed apices and steeply sloping sides. Abdomi-
nal ventrite 1 much longer than each of ven-
trites 2–4 and hypopygidium as long as ventrite
1. Legs moderately thin, moderately long and
comparatively stout, apparently with a straight
protibia.
Notes. This genus is characterized by the
rather steeply sloping lateral sides of elytra
which can be traced due to the strongly raised R
and the sloping stripe of one of elytron well
visible in the holotype M. bartenevi sp.n. The
differences of both congeners of this new genus
are very great (anterior edge of head and struc-
ture of antennae), althouth at the moment it is
not necessary to separate them into two genera
or subgenera.
Comparison. Using the above key to genera
and subgenera, this new genus can be diagnosed
by the extraordinary structure of the head with
very large eyes and a very short anterior part of
the frons, also by the peculiar shape of the
pronotum. The very distinct R in the elytra of
Menatops gen.n. finds an analogy only in the
venation of Cainocups gen.n., known only from
a fragment of an elytron which, in contrast to the
elytra of Menatops gen.n., shows R reaching the
elytral top and its consequent fusion with M
(first), CuA (second) and A1 (third). Besides
this, in addition to the above-mentioned charac-
ters in structure of the head, pronotum and
elytral venation, it differs from other cupedine
genera showing 10 or more long longitudinal
rows of cells on the elytra in the following
features:
– Anaglyphites stat.n. in the clear primary
veins and the fusion of A1 and CuA on the
elytra;
– Cainomerga subgen.n. in the robust body
with more rectilinear elytral sides, wider and
shorter elytra, somewhat shorter and thicker
legs;
– Chalepocarabus in the more slender body
with less strongly arcuate elytral sides, expressed
veins on the elytra, the shorter and transverse
pronotum, the wider and shorter elytra, the
raised primary veins, and the very short anten-
nomere 2;
– Cupidium in the longer prosternum before
the procoxae and the conjointly subacute elytral
apices;
– Gracilicupes in the longer prosternum, the
pronotum markedly wider than the head, the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
101
thicker antennae with a longer and thicker scape,
the shorter pronotum, the wider and shorter
elytra, and the very short antennomere 2;
– Kirghizocupes in the more robust body,
the pronotum not so much wider than the head,
the wider and shorter elytra, and not so acumi-
nate elytral apices which are ended conjointly,
not separately;
– Mesocupes s.str. in the more robust body,
the better expressed primary veins and the
clear fusion of A1 and CuA on the elytra, the
shorter pronotum, and the very short antenno-
mere 2;
– Paracupes s.str. and Paracupoides gen.n.
in the more robust body, the absence of tuber-
cles on the frons, the open antennal insertions,
the shorter pronotum, the wider and shorter
elytra, the conjointly acute elytral apices (not
truncate) and the not curved protibiae;
– Priacma in the more robust body, the ab-
sence of convexities on the frons, the open anten-
nal insertions, the shorter pronotum, the wider
and shorter elytra, the conjointly acute elytral
apices, and the longer abdominal ventrite 1.
In addition to its extraordinary structure of
the head with very large eyes and a very short
anterior part of the frons, Menatops gen.n. dif-
fers from the modern taxa with nine long longi-
tudinal rows on the elytra not only in the number
of these rows, but also in the more robust body
with a shorter and hexagonal pronotum, the
much wider elytra, and the gently sloping elytral
sides. Menatops gen.n., in addition to its head
with very large eyes and a very short anterior
part of the frons, differs from the fossil taxa with
nine long longitudinal rows on the elytra in the
number of these rows and also the following
characters:
– Apriacma gen.n., Cupes and Furcicupes in
the more robust body, the strongly transverse
pronotum with non-projecting anterior angles;
– Cretomerga gen.n. in the more robust
body, the longer prosternum before narrowly
separated procoxae, and the strongly transverse
pronotum with non-projecting anterior angles;
– Cupopsis gen.n. in the more robust body;
– Latocupes (? =Pulchicupes) in the less
transverse cells on the elytral disk;
– Miocupes in the more distinct cells on the
disk of the elytra;
– Taxopsis gen.n. in the smaller body, the
less transverse cells on the elytral disk, the
narrow prosternal process, the strongly trans-
verse pronotum with non-projecting anterior
angles.
Finally, Menatops gen.n., in addition to its
head with very large eyes and a very short
anterior part of the frons, differs from the forms
with an unclear number of long longitudinal
rows of cells, namely, Priacmopsis in the small-
er and more robust body, and from Cupidium in
the more robust body, the longer prosternum
before the procoxae, the non-arcuate pronotal
sides, and the conjointly acute elytral apices
(not separately rounded).
Note. This new genus looks very similar to
Latocupes, particularly in its general appear-
ance, the shape of the pronotum and the pecu-
liarities of the cells on the elytral disk.
Menatops orbiculatus (Kirejtshuk, Nel et
Colomb, 2010), comb.n.
Figs 10–17.
Cupes orbiculatus Kirejtshuk, Nel et Colomb, 2010:
217.
Holotype. “MNT-05-198” (MTM), print
and counterprint. The specimen was collected
in a new, small outcrop near the south-east end
of the village of Menat and entombed in a
comparatively smooth, somewhat fragile and
loosened rock.
Notes. As the scales for illustrations used in
the original description of this species (Kirej-
tshuk et al., 2010) were wrong, the true mea-
surements of the holotype body are actually as
given in the corrections to the description be-
low. After an additional study, particularly us-
ing electron microscopy, some characters in
shape of the head and pronotum have also been
clarified. In the original description, this species
was assigned to the genus Cupes, albeit its
placement in this genus was questioned.
Locality and stratigraphy. Paleocene,
Menat (see Materal and methods).
Diagnosis. This species is very distinct from
the another subgener (M. bartenevi sp.n.) in the
102
A.G. Kirejtshuk et al.
wider body, narrower antennae and much small-
er antennomere 2 (pedicel), apparently narrow-
er anterior edge of head and lack of median line
on upper surface of head, shape of pronotum
and its elytra less than twice as long as wide
combined (while the elytra of M. bartenevi sp.n.
are more than twice as long as wide combined).
Corrections to the description. Body length
9.1 mm, head 1.3 mm long and 1.5 mm wide,
body width 3.7 mm. Surface of head without
prominent upper tubercles, median part of frons
and anterior part of occiput apparently elevated,
at antennal insertions along posterior half of
eyes and at temples with distinct grooves. Eyes
particularly large, with diameter more than half
of distance between eyes. Elytra with at least 10
(probably 11) long longitudinal rows of large
and somewhat transverse cells, clearly raised
veins CuA and A1 fusing together; intermediate
vein between veins M and CuA (also joining
with CuA and A1) and short anal vein at scutel-
lum more or less distinct. Mesocoxae oval and
subconjoining (wrongly drawn in original de-
scription). Metaventrite slightly longer than mes-
oventrite. Metepisterna about three times as long
as wide. Metacoxae slightly oblique and only
slightly longer than mesocoxae in mesal parts.
Menatops bartenevi sp.n.
Figs 165–168.
Holotype. “MNHN F.A. 57524” (“NEL
2947”). The print and counterprint of the holo-
type represent comparatively complete speci-
men (the print with traced most part of body,
antennae, femora, pro- and mesotibiae, and also
mesotarsus; the counterprint with exposed dis-
tal dart of body — pterothorax with elytra). The
specimen has been collected by A. Nel in a new,
small outcrop near the south-east of the village
of Menat, rock comparatively smooth and be-
coming loose.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The new species is named after
Alexander F. Bartenev, professor in entomolo-
gy and invertebrate zoology at Kharkov State
(National) University, close friend of the first
author from the student time, who recently passed
away (end of November 2015).
Diagnosis. See the diagnosis of the preced-
ing species.
Description. Body length (with elytra) 7.2
mm, width apparently 2.1 mm, elytral length 4.4
mm. Body elongate; apparently slightly to mod-
erately convex dorsally and subflattened ven-
trally. Integument of head and pronotum appar-
ently with dense and coarse punctures; elytra
with large suboval or subquadrate to somewhat
subpolygonal cells arranged in at least 10 longi-
tudinal rows between veins (and probably 11th
row externally from Sc).
Head suboval and without traces of clear
upper tubercles, suboval and nearly transverse,
widest at extremely large eyes and apparently
separated by half of transverse diameter of eye,
temples gently narrowing posteriorly, antennal
insertions apparently open. Mandibles and la-
brum not visible. Antennae comparatively short
and rather thick; scape longest and rather thick;
antennomere 2 (pedicel) subcylindrical, nearly
as long as flagellomeres, markedly narrower
than scape and slightly wider than flagellomer-
es; flagellomeres subequal in length and ulti-
mate one acuminate. Prothorax moderately short;
pronotum about one and 1/4 as wide as long,
apparently slightly and gently convex, its sides
seemingly widely (sub) explanate and slightly
widening anteriorly, its sides rectilinear, anteri-
or edge convex at the middle, anterior and
posterior angles stump; prosternum before pro-
coxae rather longer than procoxae, procoxae
apparently narrowly separated. Mesoventrite
markedly longer than prosternum. Mesocoxae
moderately large and distance between them
about 1/5 of their diameter. Metaventrite very
short. Metacoxae transverse, about three times
as wide as long and their mesal parts moderately
projecting posteriorly. Abdomenal ventrites 2–
5 subequal in length, hypopygidium (ventrite 5)
rounded to subangular at apex. Elytra slightly
wider than prothorax, with very broadly arcuate
sides, together more than twice as long as wide,
steeply sloping along sides; with moderately
expressed veins.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
Taxonomy of the reticulate beetles of the subfamily Cupedinae
103
edges, about three times as long as wide. Tibiae
apparently somewhat shorter than femora and
very thin. Mesotarsus represented much longer
than mesotibia.
5.6. Genus Cupes Fabricius, 1801
Type species Cupes capitatus Fabricius,
1801, by monotypy, recent.
= Cupoides Motschulsky, 1856: 27 (type species
Cupoides tesselatus Motschulsky, 1856, by monotypy,
fossil);
= Tenomerga Neboiss, 1984: 448 (type species Cupes
mucidus Chevrolat, 1829, by original designation, recent);
= Distocupes Neboiss, 1984: 457 (type species Cupes
varians Lea, 1902, by original designation, recent).
Composition. The genus includes 15 spe-
cies from deposits beginning from the Pale-
ocene and 16 species are known in the extant
fauna (Neboiss, 1984; Ge, Yang, 2004). The
fossil species are listed in Chapter 8 and in the
catalogue by Kirejtshuk & Ponomarenko (2016).
Diagnosis and comparison (after Pono-
marenko, 1969; Neboiss, 1984; Kirejtshuk,
2005). It is necessary to note that, in addition to
the notes in Kirejtshuk et al. (2010a), the spe-
cies of this genus are characterized by the rather
slender body with the subparallel-sided elytral
edges, subvertically sloping elytral sides, the
tuberous dorsal surface of the head with the
strongly raised tubercles over each antennal
insertions. Besides, the venation of elytra of
Cupes species is represented by the strongly
raised primary veins looking like ribs, which
show a more or less clear difference in the level
of development between the primary and second-
ary veins, and also cells between the veins are
very large and subpolygonal (not transverse).
The genus Cupes can be easily diagnosed
after the above key to genera and subgenera.
The rather detailed diagnostics of the modern
species of Cupes among modern representa-
tives of other cupedines was published by
Neboiss (1984). This genus is rather similar to
the fossil groups of the subfamily with nine long
longitudinal rows of cells on the long elytra,
although, some modern representatives of Cupes,
except the short prescutellar row of cells, have
also a short row at base of the lateral edge of
elytra (Neboiss, 1984). It is most similar and
probably most closely related to Taxopsis gen.n.,
differing from the latter only in the characters
listed in the mentioned key (body size, features
of the prosternum, outline of the elytral cells).
By its moderately long prosternum before the
procoxae and the raised tubercles of the frons
covering the narrowly separated antennal inser-
tions Cupes can be distinguished from Cre-
tomerga gen.n. and Furcicupes with the very
short prosternum before the procoxae, from
which it is also distinct in the moderately narrow
neck (neck of Cretomerga gen.n. and Furci-
cupes is very narrow) and from the first, in
addition, in the narrowly separated procoxae
(procoxae of Cretomerga gen.n. are widely sep-
arated). The Cupes differs from the remainder
extinct groups with nine rows of cells on elytra:
— Apriacma gen.n. in the absence of Y-shaped
depression on the head and the narrowly sepa-
rated antennal insertions covered by the tuber-
cles of the frons; — Miocupes Ponomarenko,
1973 in the longer and narrower head with the
moderately narrow head and the moderately
large eyes, the primary veins fused at elytral
apices (primary veins of Miocupes are parallel
along the entire length); — Cupopsis gen.n. in
the moderately long head with the not large eyes
and the absence of concavities along the anteri-
or half of the inner edge of the eyes and the
steeply (subvertically) sloping elytral sides; —
Latocupes in the more slender body, the narrow-
ly antennal insertions covered by the tubercles
of the frons (absence of Y-shaped depression),
the usually narrower pronotum (less than twice
as wide as long, although the pronotum of C.
groehni Kirejtshuk, 2005 is as wide as in the
species of Latocupes) and the elytra with the
subvertically sloping sides, the strongly raised
primary veins and the less transverse cells (less
than twice as wide as long) on the elytra.
Another group of genera is united by the
groups with 10 or 11 long longitudinal rows of
cells on elytra [Chalepocarabus, Gracilicupes,
Kirghizocupes, Menatops gen.n., Mesocupes
s.lato, Paracupes s.lato (including Paracupo-
ides subgen.n.) and Priacma]. However, Cupes,
except the number of the long longitudinal rows
of cells, differs from:
104
A.G. Kirejtshuk et al.
Figs 41–44. Cupesmanifestus Kirejtshuk, Nel et Collomb, 2010 (holotype “MNT-06-902”, print, micrograms,
photographed under scanning electron microscope (SEM)); Paleogene, Paleocene; France, Menat: 41 —
body; 42 — head and prothoracic segment with appendages; 43 — scales on elytra; 44 — scales on pronotum.
Length of specimen 6.1 mm.
Рис. 41–44. Cupes manifestus Kirejtshuk, Nel et Collomb, 2010 (голотип “MNT-06-902”, отпечаток,
сфотографированный на растровом электронном микроскопе (SEM)); палеоген, палеоцен; Франция,
Мена: 41 — тело; 42 — голова и переднегрудной сегмент с придатками; 43 — чешуйки на
надкрыльях; 44 — чешуйки на переднеспинке. Длина экземпляра 6,1 мм.
– Chalepocarabus in the subparallel-sided
body, the larger eyes, the transverse pronotum,
the elytra with the well raised primary veins
clearly fused at apex, subverticaly sloping sides
and the dense cells in the long longitudinal
rows;
– Gracilicupes in the markedly wider prono-
tum usually with the rectiliner sides (not arcu-
ate) and the projecting anterior angles; the mod-
erately large eyes (prothorax of Gracilicupes
much narrower than its head with the very larger
eyes), the longer prosternum, and the antennal
insertions covered by the tubercles of the frons;
– Kirghizocupes in the not elongate head,
the absence of Y-shaped depression on the frons,
the narrowly separated antennal insertions cov-
ered by the tubercles of the frons and the prono-
tum markedly narrower than elytra;
Taxonomy of the reticulate beetles of the subfamily Cupedinae
105
– Menatops gen.n. in the more slender body,
the moderately long head with the moderately
large eyes, only three long primary veins (A1,
CuA and M) expressed on the elytral disk (elytra
of Menatops gen.n. with the clear R fusing A1),
the antennal insertions covered by the tubercles
of the frons;
– Mesocupes s.lato in the more slender body,
the well raised primary veins of the elytra, the
subverticaly sloping elytral sides and the nar-
rowly separated antennal insertions covered by
the tubercles of the frons, and in the clear fusion
of the primary veins at elytral apices;
– Paracupes s.str. in the elytra with well
raised primary veins, the subverticaly sloping
sides, the simple lateral epipleural carina and
the subacute apices, and also in the protibiae
straight or slightly curved at apex;
– Paracupoides subgen.n. in the elytra with
the subacute apices, the subparallel long anal
veins (A1) and the elongate (lanceolate) scales
on the dorsum, and also in the protibiae straight
or slightly curved at apex;
– Priacma in the elytra with the well raised
primary veins, the subverticaly sloping sides,
the simple lateral epipleural carina and the sub-
acute apices, the lack of preapical closing fold
on the below side of the elytra, the narrowly
separated gular sutures, the longer ventrite 1;
– Priacmopsis in the much smaller and slen-
der body with the subparallel-sided elytra, the
comparatively larger eyes, the longer metacox-
ae and the longer abdominal ventrite 1.
Lastly the genus Cupes differs from Cupi-
dium with unclear number of the long longitudi-
nal rows of cells in the more slender body, the
usually not arcuate pronotal sides and the longer
prosternum before the procoxae and the con-
jointly subacute elytral apices.
Notes. All species of Cupes found in Menat
are characterised by the rather wide anterior
part of the head with only one pairs of the
convexities covering the most length of their
epicranium and separated only a very narrow
median furrow, the very thick scapes, the not
outlined tarsal grooves in the anterior part of
their prosternum (if they are present at all) and
the pronotum subrectilinearly widening anteri-
ad. The studies of the specimens under electron
microscope provided with a chance to observe
scales on the elytral integument at least in two
examined species.
Cupes manifestus Kirejtshuk, Nel et Co-
lomb, 2010
Figs 41–44, 146, 147, 169–172.
Cupes manifestus Kirejtshuk, Nel et Colomb, 2010:
218.
Holotype. MNT-06-902-A and MNT-06-
902-B (MTM), print and counterprint. The spec-
imen has been collected in a new, small outcrop
near the south-east of the village of Menat, the
rock comparatively hard and consistent.
Additional specimens. “MNHN.F.A.
52771” (“NEL 2144”), print and counterprint of
an incomplete specimen showing very obliterat-
ed and unclear outlines of sclerites and deposit-
ed in the comparatively soft and fragile rock.
Most details of this specimen are not traceable
or missing. “MNHN.F.A.52770” (“NEL 801”),
print with the incomplete specimen showing
very obliterated and unclear outlines of sclerites
and deposited in the comparatively soft and
fragile rock. Most details of this specimen are
also not traceable or missing, but five segments
of the left antenna, the right anterior leg, the
metacoxae and the posterior legs are visible.
“MNHN F.A. 57525” (“NEL 1701”), print with
incomplete specimen with unclear outlines of
exposed sclerites is deposited in very hard rock.
The specimen demonstrates the body with right
antenna, subtriangular characteristic head, meso-
and metacoxae, traces of profemora, intermedi-
ate and posterior legs. The additional specimens
“MNHN.F.A.52771”, “MNHN.F.A.52770” and
“MNHN.F.A.57525” originated from a new,
small outcrop near the south-east of the village
of Menat. “MNHN F.A. 57526” (“Oli 186”),
print with incomplete specimen with coarsely
fragmented organic remains and unclear out-
lines of exposed sclerites is deposited in moder-
ately hard rock. The specimen demonstrates the
body with basal segments of both antennae,
characteristic outline of the head, slightly visi-
ble prothorax with procoxae, pterothorax with
metacoxae, traces of profemora and right protib-
106
A.G. Kirejtshuk et al.
ia. “MNHN F.A. 57527” (“056”), print with
incomplete specimen with clear outlines of ex-
posed sclerites is deposited in soft rock. The
specimen demonstrates the body with most seg-
ments of both antennae (preapical ones of left
antennae are somewhat obliterated), character-
istic outline of the head, obliterated place of
rock with slightly visible outline of pronotum
with clear sculpture, pterothorax with metacox-
ae, traces of all femora and protibiae. The spec-
imens “MNHN F.A. 57526” and “MNHN F.A.
57527” have been collected by Philippe Olivier
in the historical site of the ancient quarry of
“Noir d’Auvergne” where Piton investigated in
the years 1935–1940.
Notes. As the scales used in the original
description of this species (Kirejtshuk, et al.,
2010) were given wrong, the true measurements
of the holotype body are actually as given in the
corrections and addition to the redescription
below. After an additional study, particularly
using electron microscopy some characters in
shape of the head and the pronotum were clari-
fied.
Locality and stratigraphy. Paleocene,
Menat (see Materials and methods).
Diagnosis. This new species can be easily
diagnosed among the species from Menat after
the below key to species. As all of them are
rather similar, the diagnosis of C. distinctissi-
mus sp.n. (see below) combining with the diag-
nosis published by Kirejtshuk et al. (2010) and
this key can be used for a reliable diagnosis of
the species under consideration.
Corrections and addition to the descrip-
tion of the holotype. Body 6.2 mm long, head
2.2 mm long and 2.5 mm wide, body width 2.8
mm. Elytra with 9 rows of large polygonal cells.
Dorsum covered with dense lanceolate scales,
which are arranged in accordance with veins on
elytra. Head with truncate anterior edge of frons,
weak but clear tubercles over close antennal
insertions, elongate paramedian convexities
along frons and vertex and on each temple a
characteristic and weakly raised tubercle clear-
ly visible. Prosternum somewhat longer than
mesoventrite and somewhat shorter than
metaventrite.
Variations (additional specimens).
“MNHN.F.A.52771”: Body 5.6 mm long, head
1.7 mm long and 1.6 mm wide, body width 2.3
mm. In spite of the fact that many structural
features of the specimens examined are unclear,
it can be visible that its temples are short and not
prominent, and its pronotum is widening anteri-
ad. Therefore, it fits Cupes manifestus better
than other congeners from Menat, although the
head of this specimen seems to have been some-
what turned and deformed. “MNHN.F.A.
52770”: Body 7.8 mm long, head 1.3 mm long
and 1.1 mm wide, body width 2.3 mm. Many
structural features of the specimens examined
are unclear, although general outlines of visible
sclerites are similar to those in C. manifestus
better than other congeners from Menat. Never-
theless its comparatively short head with larger
eyes and apparently arcuate pronotal sides are
somewhat different from those in C. manifestus.
“MNHN F.A. 57525”: body 8.9 mm long with-
out antenna and 2.0 mm wide. This specimen
can be regarded as C. manifestus because of
characteristic many proportions of the individ-
ual sclerites (particularly head and right anten-
na), although the general outline of its body is
more slenger and elytra are subparallelsided.
“MNHN F.A. 57526”: 7.8 mm long without
antennae. This specimen has hardly traceable
most characters, but those of them, which are
visible, more or less correspond to the typical
characters of C. manifestus, but its general out-
lines of elytra look like somewhat more arcuate
and temples of its head are not so prominent.
“MNHN F.A. 57527”: body 7.2 mm long with-
out antenna. This specimen seems to have a
most correspondence to the holotype.
Cupes distinctissimus sp.n.
Figs 45–56.
Holotype. “MNHN.F.A.51118”, collected
by A. Nel. The print and counterprint of the
holotype is deposited in rather soft and fragile
rock, and represented by the mixture of visible
outlines from dorsal and ventral sclerites (the
print with more exposed ventral sclerites and
the counterprint with more exposed dorsal scler-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
107
Figs 45–48. Cupes distintissimus sp.n. (holotype “MNHN.F.A.51118”, dry print and counterprint,
photographed under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 45 —
print, body; 46 — counterprint, body; 47 — print, head and prothorax with appendages; 48 — counterprint,
pronotum. Length of specimen 7.3 mm.
Рис. 45–48. Cupes distintissimus sp.n. (голотип “MNHN.F.A.51118”, сухие отпечаток и противоотпе-
чаток, сфотографированные на микроскопе Olympus SCX9 с камерой Olympus); палеоген, палеоцен;
Франция, Мена: 45 — тело отпечатка; 46 — тело противоотпечатка; 47 — голова и переднегрудь
отпечатка с придатками; 48 — переднеспинка противоотпечатка. Длина экземпляра 7,3 мм.
ites), although the print shows some damage in
the prothoracic segment and the counterprint is
with missing posterior part of elytra, missing or
unclear few antennomeres, two tibiae, many
tarsomeres. The specimen has been collected by
A. Nel in a new, small outcrop near the south-
east of the village of Menat, in the rock compar-
atively smooth and becoming loose.
108
A.G. Kirejtshuk et al.
Figs 49–52. Cupes distintissimus sp.n. (holotype “MNHN.F.A.51118”, print and counterprint); Paleogene,
Paleocene; France, Menat: 49 — print, ventral, with broken line outlined sutural edges and primary veins
of elytra, with dotted lines probable outlines of temples; 50 — counterprint, dorsal; 51 — print, micrograms
of scales on elytra, photographed of print under scanning electron microscope (SEM); 52 — idem, with
another magnification. Length of specimen 8.3 mm.
Рис. 49–52. Cupes distintissimus sp.n. (голотип “MNHN.F.A.51118”, отпечаток и противоотпечаток);
палеоген, палеоцен; Франция, Мена: 49 — отпечаток, снизу; прерывистой линией очерчены шовные
края и первичные жилки надкрылий; пунктиром показаны вероятные очертания висков; 50 —
противоотпечаток, сверху; 51 — отпечаток, чешуйки надкрылий, сфотографированные на растровом
электронном микроскопе (SEM); 52 — то же, при другом увеличении. Длина экземпляра 8,3 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
109
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The epithet of this new species
is the superlative of “distinctus” (separate, dis-
tinct; definite, lucid)
Diagnosis. This new species is easily diag-
nosed among the species from Menat after the
below key to species. At the same time because
these species are rather similar and probably
closely related, the diagnosis of this new species
can be used for other species from Menat. Cupes
distinctissimus sp.n. demonstrates most simi-
larity with fossil C. rohdendorfi Iablokoff-
Khnzorian, 1960 differening from it in the long-
er head with only one pair of long convexities,
the much smaller eyes, the shorter antennae with
the thicker antennomeres. In addition to the
characteristic tubercles (convexities) on the head,
and also the strongly projecting and acute ante-
rior angles of pronotum, this new species differs
from other fossil congeners:
– C. eckfeldensis (Tröster, 1993), comb.n. in
the subquadrangular head with the large scape
and the subquadrangular prosternum;
– C. groehni Kirejtshuk, 2005 in the smaller
eyes and the comparatively shorter antennae;
– C. hoffeinsorum Kirejtshuk, 2005 in the
smaller eyes, the wider pronotum and the much
more stout antennomeres;
– C. kerneggeri Kirejtshuk, 2005, C. komis-
sari Kirejtshuk, 2005 and C. ponomarenkoi
Kirejtshuk, Nel et Colomb, 2010 in the smaller
eyes;
– C. tesselatus (Motschulsky, 1856) in the
smaller eyes, the shorter antennae with the thicker
antennomeres and the markedly narrower prono-
tum;
– C. weitschati Kirejtshuk, 2005 in the shorter
antennae with the thicker antennomeres and the
wider pronotum.
Cupes distinctissimus sp.n. is distinct from
C. messelensis (Tröster, 1993), comb.n. repre-
sented only by the pterothorax, abdomen and
elytra in the more widely rounded apex of the
hypopygidium. This new species can be distin-
guished from C. praeglacialis Gersdorf, 1976
described with the rather short sharacteristics
and bad pictures at least in the smaller body and
the much thicker antennomeres. Finally, the
new species under consideration differs from
the modern members of the genus in the some-
what longer head with only one pair of parame-
dian convexities located along the most part of
dorsal surface of the head and the more or less
shorter antennae, the pronotum strongly wid-
ened anteriad and with the rather projecting and
acuminate anterior angles.
Description. Body length (with elytra) 8.3
mm, width apparently 2.5 mm, head 1.2 mm
long and 1.4 mm wide, elytral length 5.6 mm.
Body elongate; apparently moderately to rather
convex dorsally and subflattened ventrally. In-
tegument of head and pronotum with moderate-
ly large regular punctures separated by much
less than one puncture diameter or less; lower
surface of head and prosternum with somewhat
larger and shallower punctures; pterothoracic
ventral sclerites and abbomen with somewhat
finer and rather dense punctures; elytra with
large subpolygonal cells arranged in apparently
nine rows between veins. Dorsum covered with
dense lanceolate scales, which are arranged in
accordance with veins on elytra.
Head subquadrangular, with only clear tu-
bercles at antennal insertions, somewhat longer
than wide, apparently widest at subangular tem-
ples, with moderately large oval eyes (moder-
ately wide dorsally and rather narrow ventrally),
their elongate diameter nearly as great as length
of frons before eyes and somewhat less than
length of temples; gular sutures moderately sep-
arated. Mandibles not visible. Mentum moder-
ately small, subquadrangular and transverse.
Antennae comparatively short; scape about as
long as flagellomeres and almost twice longer
than antennomere 2 (pedicel); scape rather swol-
len, somewhat wider than pedicel and basal
flagellomeres, other flagellomeres gradually
becoming narrower to ultimate flagellomere
with pointed apex. Prothorax subtrapezoid and
moderately short; pronotum about 1.5 times as
wide as long and markedly wider than head;
apparently slightly and gently convex, its sides
seemingly widely (sub) explanate and widening
anteriorly in S-curve, its sides shallowly emar-
ginate, anterior edge subbi-sinuate, anterior and
110
A.G. Kirejtshuk et al.
Figs 53–56. Cupes distintissimus sp.n. (holotype “MNHN.F.A.51118”, print and counterprint, micrograms
under scanning electron microscope (SEM)); Paleogene, Paleocene; France, Menat: 53 — print, anterior part
of head; 54 — counterprint, head and pronotum; 55 — counterprint, distal part of elytra; 56 — counterprint,
integument on pronotum. Length of specimen 8.3 mm.
Рис. 53–56. Cupes distintissimus sp.n. (голотип “MNHN.F.A.51118”, отпечаток и противоотпечаток,
сфотографированные на растровом электронном микроскопе (SEM)); палеоген, палеоцен; Франция,
Мена: 53 — передняя часть головы отпечатка; 54 — голова и переднеспинка противоотпечатка; 55 —
дистальная часть надкрылий противоотпечатка; 56, поверхность переднеспинки противоотпечатка.
Длина экземпляра 8,3 мм.
posterior angles with distinct top, anterior ones
rather projecting, its disk with median carina
and narrow furrow along lateral edge, inter-
space between carina and furrow rather swol-
len; prosternum before procoxae about 1.5 times
as long as procoxae and probably slightly de-
pressed along propleura (but without tarsal
groove along anterior edge); procoxae appar-
ently narrowly separated. Mesoventrite appar-
ently about as long as prosternum before pro-
coxae and somewhat shorter than metaventrite.
Mesocoxae moderately large and distance be-
tween them extremely narrow. Metaventrite
about 2/3 as long as wide at base. Metacoxae
medially projecting anteriad, somewhat wider
than long and their mesal parts moderately pro-
jecting posteriorly. Abdomen with visible bound-
aries between last three ventrites (without trace
Taxonomy of the reticulate beetles of the subfamily Cupedinae
111
of overlapping); hypopygidium about 1.5 times
as long as ventrite 4 and rounded at apex. Elytra
slightly wider than prothorax, with slightly arc-
uate sides, slightly more than twice twice as long
as wide combined, apparently moderately con-
vex; with well expressed main veins and clear
conjoining of CuA and A1.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, about 2.5 times as long as wide. Tibiae
apparently somewhat shorter than femora and
moderately thin. Tarsi represented by moder-
ately wide and somewhat lobed basal tarsomer-
es.
Cupes simillimus sp.n.
Figs 57–67.
Holotype. “MNHN.F.A5 1120”, collected
by A. Nel, print and counterprint. The holotype
is deposited in very hard rock and represented
by the somewhat obliterated outlines of scler-
ites, its counterprint is with missing posterior
part of body, most antennomeres, two femora,
four tibiae, many tarsomeres. The specimen has
been collected by A. Nel in a new, small outcrop
near the south-east of the village of Menat. The
rock is rather smooth.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Etymology. The epithet of this new species
is the superlative of “similis” (like, similar,
resembling).
Diagnosis. This new species is easily diag-
nosed among the species from Menat after the
below key to species. As all of them are rather
similar, the diagnosis of C. distinctissimus sp.n.
(see below) combining with this key can be used
for a reliable diagnosis of the species under
consideration.
Description. Body length (with elytra) 6.8
mm, width apparently 2.0 mm, head 1.6 mm
long and 1.2 mm wide, elytral length 4.6 mm.
Body elongate; apparently moderately to rather
convex dorsally and subflattened ventrally. In-
tegument of head and pronotum with compara-
tively large regular punctures separated by much
less than one puncture diameter or less; pterotho-
racic ventral sclerites and abbomen with some-
what larger punctures; elytra with large subpo-
lygonal cells arranged in nine rows between
veins. Dorsum without lanceolate scales visible
under electronic microscope.
Head suboval and widest at arcuate temples,
with only clear tubercles at antennal insertions
and apparently on temples, about as long as
wide, with moderately large oval eyes moder-
ately wide and strongly close to anterior edge of
frons. Mandibles not visible. Mentum moder-
ately small, subquadrangular and transverse.
Antennae represented by only scapes (rather
thick and slightly longer than wide). Prothorax
subtrapezoid and moderately short; pronotum
about 1.5 times as wide as long and markedly
wider than head; apparently slightly and gently
convex, its sides seemingly widely (sub) ex-
planate and rectilinearly widening anteriorly, its
sides shallowly emarginate, anterior edge sub-
bi-sinuate, anterior and posterior angles with
distinct top, anterior ones rather projecting, its
disk with median carina and narrow furrow
along lateral edge, interspace between carina
and furrow rather swollen; prosternum before
procoxae apparently 1.5 times as long as pro-
coxae. Mesoventrite looking like very short
(probably as sequence of peculiar fossiliza-
tion). Mesocoxae moderately large and distance
between them extremely narrow. Metaventrite
apparently twice longer than prosternum before
procoxae and about 2/3 as long as wide at base.
Metacoxae transverse, about 1.5 times as wide
as long and their mesal parts moderately pro-
jecting posteriorly. Abdomen with long over-
lapping, ventrite 1 somewhat longer than each
of ventrites 2–5, which are subequal in length;
hypopygidium rounded at apex. Elytra slightly
wider than prothorax, nearly subparallel-sided,
slightly more than twice as long as wide com-
bined, apparently moderately convex; with well
expressed main veins.
Femora comparable in shape and propor-
tions, gently curved along anterior and posterior
edges, about three times as long as wide. Tibiae
apparently somewhat shorter than femora and
moderately thin. Tarsi represented by remains
of one rather narrow mesotarsus.
112
A.G. Kirejtshuk et al.
Figs 57–61. Cupes similliimus sp.n. (holotype “MNHN.F.A5 1120”, print and counterprint, photographed
under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 57 — print, body;
58 — print, body, dorsal, with broken outlines of ventral sclerites head and thorax; 59 — counterprint, body;
60 — counterprint, head and prothorax; 61 — print, anterior part of body. Length of specimen 6.8 mm.
Рис. 57–61. Cupes similliimus sp.n. (голотип “MNHN.F.A5 1120”, отпечаток и противоотпечаток,
сфотографированные на мироскопе Olympus SCX9 с камерой Olympus); палеоген, палеоцен; Фран-
ция, Мена: 57 — тело отпечатка; 58 — тело отпечатка, сверху; прерывистой линией очерчены нижние
склериты головы и груди; 59 — тело противоотпечатка; 60 — голова и переднегрудь противоотпе-
чатка; 61 — передняя часть тела отпечатка. Длина экземпляра 6,8 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
113
Figs 62–67. Cupes simillimus sp.n. [holotype “MNHN.F.A5 1120”, print and counterprint, micrograms,
photographed under scanning electron microscope (SEM)]; Paleogene, Paleocene; France, Menat: 62 —
print, body; 63 — counterprint, metaventrite and base of abdomen; 64 — print, elytra; 65 — print, elytral
apices; 66 — counterprint, head and prothorax; 67 — counterprint, body, ventral. Length of specimen 6.8 mm.
Рис. 62–67. Cupes simillimus sp.n. [голотип “MNHN.F.A5 1120”, отпечаток и противоотпечаток,
сфотографированные на растровом электронном микроскопе (SEM)]; палеоген, палеоцен; Франция,
Мена: 62 — тело отпечатка; 63 — метавентрит и основание брюшка противоотпечатка; 64 —
надкрылья отпечатка; 65 — вершины надкрылий отпечатка; 66 — голова и переднегрудь противо-
отпечатка; 67 — тело противоотпечатка, снизу. Длина экземпляра 6,8 мм.
114
A.G. Kirejtshuk et al.
5.7. Key to species of Cupes from Menat
1. Head subquadrangular and with subangular tem-
ples; anterior edge of frons at distance from
anterior edge of eyes nearly subequal to elongate
diameter of eyes; pedicel and basal flagellomer-
es only slightly narrower than scape; elytra with
gently arcuate sides; body length ca. 8.3 mm.
Figs 45–56 ................ C. distinctissimus sp.n.
– Head suboval and with rounded temples; combina-
tion of remaining characters other ................. 2
2. Eyes with usual distance from anterior edge of
frons; head widest at eyes; temples only slightly
longer than eyes; elytra with gently arcuate sides;
body length ca. 5.6–8.9 mm. Figs 41–44, 169–
172 ...................................................................
C. manifestus Kirejtshuk, Nel et Colomb, 2010
– Eyes very close to anterior edge of frons; head
widest at rounded temples; temples about twice
as long as head; elytra with subparallel lateral
edges;body length ca. 6.8 mm. Figs 57–67 ....
........................................... C. simillimus sp.n.
5.8. Genus incertus sp.1
Figs 142–144.
Specimen examined. MNHN “R 63906”,
print. The specimen includes the badly pre-
served and somewhat charry coleopterous re-
mains, rather fragile and unclear; almost com-
plete body have no traces of outlines of thoracic
sclerites, and its appendages are represented by
fragments of anterior legs, feeble contour of one
middle femur, one antenna with seven antenno-
meres and another with six antennomeres. This
piece of rock is comparatively smooth and slight-
ly becoming loose.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. Body length (with elytra) 12.6 mm,
width 4.4 mm. The outlines of the accessible
structures (shape of head, pronotum, elytra).
The small pronotum of this species has a typical
outline for many cupedines, however its propor-
tion to other sclerites is similar to that in the
Cretaceous Gracilicupes, it cannot be put in this
taxon because of different shape of its head and
pronotum. The elytra look like having gently
sloping sides with (sub) explanate edges (as in
Mesocupes), however it is impossible to associ-
ate this species with members of this genus
because of the very small pronotum. If it is
possible to admit that the elytra of the specimen
examined were spread as a result of fossiliza-
tion, this form could be an aberrant member of
the Cupes taking into consideration some exter-
nal similatity with C. manifestus. Nevertheless,
at the moment this species can be scarcely
linked with any group of the subfamily, but it
could have some relation with the two following
specimens remained without generic and spe-
cies attribution.
5.9. Genus incertus sp.2.
Fig. 149–150.
Specimens examined. “MNHN.F.A.52794”
(“NEL 2293”), print and counterprint with the
incomplete specimen showing rather obliterat-
ed and unclear outlines, deposited in the com-
paratively soft and fragile rock. It originated
from a new, small outcrop near the south-east of
the village of Menat.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. Body 14.4 mm long, head 2.4 mm
long and elytra 11.3 mm long. This specimen is
characterised by the rather slender body, the
narrow pronotum (narrower than head), the very
large eyes, the very dense and fine sculpture on
the underside, the elytra with apparently nine
longitudinal rows of transverse or subpolygonal
cells, not clear primary veins and the subverti-
cally sloping sides which are somewhat (sub)
explanate along the lateral edges. Its head seems
to be compressed and additionally spread be-
cause a feeble Y-shaped dorsal carina on its
dorsal surface is shifted aside. As a result, the
head demonstrates very large eyes which are
difficult to interprete. Thus this specimen could
belong to a separate genus which is impossible
to define in the terms comparable with those
used for other generic taxa of the subfamily.
5.10. Genus incertus sp.3.
Fig. 151–152.
Specimen examined. “MNHN.F.A.52772”
(“NEL 786”), print and counterprint with the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
115
incomplete specimen showing rather smoothed
and unclear outlines, deposited in the compara-
tively soft and fragile rock. It originated from a
new, small outcrop near the south-east of the
village of Menat.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. Body 13.2 mm long, head 2.0 mm
long and 1.8 mm wide, body width 3.5 mm,
length of elytra 9.8 mm. The head seems to have
been deformed because the eyes look like shift-
ed; posterior part of head with traces of the
paramedian convexities. Many characters of
this specimen are somewhat similar to those of
Mesocupes species from Menat, including the
transverse cells in the longitudinal rows of the
elytra. However the specimen under consider-
ation is smaller and has the narrow pronotum
and the gular sutures divergent in both anterior
and posterior parts. Nevertheless, the set of
accessible characters in it could not be used for
both generic and species diagnostics.
5.11. Genus incertus sp.4.
Fig. 173–174.
Specimens examined. “MNHN.F.A.57529”
(“Oli 137”), print of separate elytron with well
preserved chitin remains and clear traces of
imprinted venation and sculpture, deposited in
the moderately soft and fragile rock. “MNHN.
F.A.57530” (“Oli 856”), print of separate elytron
with few remains of chitin and rather obliterated
traces of imprinted of venation and sculpture,
deposited in the moderately soft and fragile
rock. The specimens have been collected by
Philippe Olivier in the historical site of the
ancient quarry of “Noir d’Auvergne” where
Piton investigated in the years 1935–1940.
Locality and stratigraphy. Paleocene,
Menat (see Materals and methods).
Notes. The separate elytra of “MNHN.F.A.
57529” with length 5.9 mm and “MNHN.F.A.
57530” with length 6.2 mm could belong to the
same genus and probably to the same species.
They also have significant similarity with those
in the modern Priacma and Paracupes in the
comparatively weak primary veins, 10 longitu-
dinal rows of cells, gentle sloping of elytral
sides and their blunt apices. Nevertheless the
disposition of veins and longitudinal rows of
cells are different from those in both mentioned
taxa. It seems to be better to leave them without
naming till new materials from Menat will be
found and studied.
6. New genera and species of Cupedinae
6.1. Apriacma gen.n.
Figs 73–75.
Type species. Priacma tuberculosa Tan,
Ren et Shin, 2006, fossil.
Etymology. The name of this new genus is
formed from Greek negative prefix “a” and
generic name “Priacma”. Gender feminine.
Composition. Four species previously de-
scribed from the early Cretaceous (Tan, Ren et
Shin, 2006) as members of Priacma: A. clavata
(Tan, Ren et Shin, 2006), comb.n., A. renaria
(Tan, Ren et Shin, 2006), comb.n., A. tubercu-
losa comb.n., and (?) A. latidentata (Tan, Ren et
Shin, 2006), comb.n.
Specimens examined. Holotype of Priac-
ma clavata “CNU-C-LB2006001”; holotype of
P. renaria “CNU-C-LB2006002”; holotype of
P. tuberculosa; “CNU-C-LB2005011”; holo-
type of P. latidentata “CNU-C-LB2005010”.
Locality and stratigraphy. All congeners
originated from Yixian; Beipiao City, Chao-
midian village, Liaoning Province, China; Early
Cretaceous, Barremian-Aptian, Yixian Forma-
tion.
Diagnosis. Body elongate of medium size
(9.5–12.3 mm). Integument with moderately
coarse sculpture and punctation; large subqua-
drangular (transverse) to polygonal cells of elytra
arranged into nine long longitudinal rows and a
short prescutellar row interspaced by weak sec-
ondary and more raised primary veins; A1
straight, almost reaching elytral apex and fusing
with CuA before apex. Head usually subtrian-
gular, with rather narrow neck (3/4–3/5 as wide
as head at temples), with more or less expressed
Y- or V-shaped depression, well projecting tem-
ples; dorsal surface with weakly raised tuber-
116
A.G. Kirejtshuk et al.
Figs 68–72. Species of of the genus Cupes (photos from S. Wedmann (SFNFM) and H. Lutz (LNNR)): 68 —
C. messelensis (Troester, 1993), comb.n., holotype (SFNFM “MeI 2806”); Paleogene, Middle Eocene;
Germany, Messel; length of specimen 6.0 mm; 69–70 — C. eckfeldensis (Troester, 1993), comb.n., holotype
(LNNR “1992/461”); Paleogene, Middle Eocene; Germany, Eckfelder Maar, print and counterprint, length
of specimen 8.8 mm; 71 — C. concolor Westwood, 1835; recent; posterior part of body, lateral (ZIN, “Am.
Sept.”), length of specimen 9.5 mm; 72 — idem, abdomen, ventral.
Рис. 68–72. Виды роды Cupes (фотографии от С. Ведманн (S. Wedmann, SFNFM) и Г. Луца (H. Lutz,
LNNR)): 68 — C. messelensis (Troester, 1993), comb.n., голотип (SFNFM “MeI 2806”); палеоген,
средний эоцен; Германия, Мессель; длина экземпляра 6,0 мм; 69–70 — C. eckfeldensis (Troester, 1993),
comb.n., голотип (LNNR “1992/461”); палеоген, средний эоцен; Германия, Экфельд; отпечаток и
противоотпечаток; длина экземпляра 8,8 мм; 71 — C. concolor Westwood, 1835; современный; задняя
часть тела, сбоку (ZIN, “Am. Sept.”), длина экземпляра 9,5 мм; 72 — то же, брюшко, снизу.
Рис. 73–76. Ископаемые виды сем. Cupedidae: 73 — Apriacma tuberculosa (Tan, Ren et Shih, 2006),
comb.n., голотип (“CNU-C-LB 2005011”, отпечаток, сфотографированный на микроскопе Leica MZ
12.5 с камерой Nikon Digital Camera DXM1200C); нижний мел, формация исянь; Китай, Ляонинь,
Исянь; тело сухого отпечатка; 74 — то же, тело; прерывистой линией очерчены вентральные
склериты головы, груди и брюшка; пунктиром прослеживающиеся очертания верхних структур
головы (бугорки) и переднеспинки (медиальное вздутие и отогнутые края); 75 — то же, тело
отпечатка под спиртом; длина экземпляра 9,8 мм; 76 — Cainocups aixensis sp.n., голотип (MNHN “Aix
18”, сфотографированный на микроскопе Olympus SCX9 с камерой Olympus); палеоген, верхний
олигоцен; Франция, Жа-де-Буффан, Экс-ан-Прованс; дистальная часть надкрылья; длина отпечат-
ка 6,3 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
117
Figs 73–76. Fossil species of the family Cupedidae: 73 — Apriacma tuberculosa (Tan, Ren et Shih, 2006),
comb.n., holotype (“CNU-C-LB 2005011”, print, photographed under Leica MZ 12.5 with camera Nikon
Digital Camera DXM1200C); Lower Сretaceous, Yixian Formation; China, Liaoning, Beipiao; body on dry
print; 74 — idem, body with broken outlines of ventral sclerites head, thorax and abdomen; and dotted line
traced outlines of dorsal structures of head (tubercles) and pronotum (median convexity and explanate sides);
75 — idem, body on print with alcohol; length of specimen 9.8 mm; 76 — Cainocups aixensis sp.n., holotype
(MNHN “Aix 18”, photographed under Olympus SCX9 with camera Olympus); Paleogene, latest Oligocene;
France, Jas de Bouffan, town Aix-en-Provence; distal part of elytron; length of print 6.3 mm.
118
A.G. Kirejtshuk et al.
cles (paramedian longitudinalconvexities along
eye edge to temples), antennal insertions mod-
erately widely separated and apparently not
covered by tubercles. Mandibles rather strong
and arcuately curved at outer edge before stout
bidentate apex. Antennae subfiliform and mod-
erately long; antennomere 1 (scape) subequal in
length with flagellomeres or slightly longer than
antennomere 2 (pedicel) and more or less thick-
er. Pronotum subtrapezoid and rectilinearly
widening anteriad, with widely explanate sides,
projecting and subacute anterior angles, usually
tuberculate surface and weak median elevated
stripe. Part of prosternum before procoxae mark-
edly longer than procoxae. Elytra 2.2–2.5 times
as long as wide combined, with subrectiliner
lateral edges, subrounded to subacute apices
and apparently steeply (subvertically) sloping
sides. Abdominal ventrites articulated without
clear overlapping, ventrite 1 subequal to clearly
longer than each of ventrites 2–4 and hypopy-
gidium about as long as ventrite 1 or longer and
rounded at apex. Legs moderately thin, fre-
quently comparatively long; with protibia
straight and narrow tarsi. Aedeagus of cupedid-
type, rather long and with wide flattened
parameres.
Comparison. This new genus is rather sim-
ilar to the recent members of the genera similar
and apparently closely related to Cupes with
nine long longitudinal rows of cells on elytra.
The main important characters to distinguish it
among the forms with the slender body having
such number of rows of cells and expressed
primary veins of elytra (Adinolepis, Asciopla-
ga, Cupes, Cupopsis gen.n., Prolixocupes, Rhip-
sideigma, etc.) are the clear Y- or V-shaped
depression on the frons, and this characters have
also the slender species of Cretomerga gen.n.
and Furcicupes, although Apriacma gen.n. dif-
fers from them in the longer prosternum before
the procoxae. The new genus under consider-
ation differs from Kirghizocupes with the Y-
shaped depression on the head and 10 (or 11)
long longitudinal rows of cells on elytra not only
in the number of these long rows of cells, but
also in the shorter head and pronotum widening
anteriorly. Apriacma gen.n. differs from Taxop-
sis gen.n. in the not transverse or slightly trans-
verse cells in the long rows on the elytra, smaller
and the more slender body with the narrower
metaventrite, the clear Y- or V-shaped depres-
sion on the frons and the narrower tarsi. It differs
from Latocupes in the much more slender body
with the subparallel-sided elytra having the steep-
ly (subvertically) sloping sides and the more
raised primary veins. From other genera with 10
or more long longitudinal rows of cells on the
elytra (Cainomerga subgen.n., Gracilicupes,
Menatops gen.n., Mesocupes s.lato, Paracupes
s.str., Paracupoides subgen.n., Priacma, and
Latocupes) Apriacma gen.n. differs not only in
the number of the rows of cells, but also in the
clear Y- or V-shaped depression (although sub-
genus Anaglyphites stat.n. also demonstrates a
more or less expressed Y- or V-shaped depres-
sion on the frons) and the pronotum strongly and
subrectilinearly widening anteriad. Besides, it
also differs from Gracilicupes in the smaller
eyes, the wider pronotum with the sides widen-
ing anteriad and with the prominent anterior
angles (not with arcuate sides narrowing both
anteriad and posteriad) and the Cupes-type of
aedeagus. It differs from Anaglyphites stat.n.
and Mesocupes s.str. in the slender body, the
subrectilinear elytral sides, the well expressed
primary veins with the fusion of A1 and CuA at
apices; from Cainomerga subgen.n. in the small-
er and less slender body; from Cupidium in the
slender body, the longer prosternum before the
coxae and the conjointly subacute elytral api-
ces; from Menatops gen.n. in the much slender
body, the longer head with the smaller eyes and
the not strongly transverse pronotum; from Para-
cupes s.lato and Priacma in the apparently ver-
tically sloping sides of the elytra, which show
the simple edge (not thickened and tuberculate),
the narrower tarsi and the more or less open
antennal insertions, and also from Gracilicupes
and Priacma in the Cupes-type of aedeagus.
Finally it differs from Paracupoides subgen.n.
in the narrower tarsi, the antennomere 2 (pedicel)
only slightly shorter than scape and the follow-
ing antennomeres, the more or less open and
more separated antennal insertions. Apriacma
gen.n. is rather distinct from Chalepocarabus in
Taxonomy of the reticulate beetles of the subfamily Cupedinae
119
the presence of nine long longitudinal rows of
cells, the clearly expressed primary veins and
the markedly larger cells on elytra, the steeply
sloping sides and subrectilinear lateral edges of
the elytra, the slender body, the transverse prono-
tum widening anteriad, and the narrower
metaventrite. It differs from Miocupes Pono-
marenko, 1973 in the more slender body, the
narrower and elongate head, the primary veins
of elytra fused at apices, the larger elytral cells
and the narrower metaventrite. Finally, the new
genus differs from Priacmopsis in the much
smaller and slender body, the larger antenno-
mere 2 (pedicel) in comparison with the previ-
ous and folloving ones, the narrower metaven-
trite.
Notes. All four species included into this
new genus were described as members of the
genus Priacma, however, they have, in contrast
to Priacma serrata, the rather clear Y-shaped
depression on frons, the steeply sloping sides
and the simple apices (without closing fold) of
the elytra and the Cupes-type of aedeagus (vis-
ible in the holotype of A. clavata comb.n.). All
these characters were ignored in their original
descriptions, and for the number of the long
longitudinal rows of cells on the elytra there was
noted 10 (Tan et al., 2006). However the holo-
types of A. tuberculosa comb.n. (Fig. 73) and
(?) A. latidentata comb.n. clearly have nine
rows of cells on the elytra. The prints of the other
specimens do not allow calculate a precise num-
ber of rows. Three species of the new genus
above mentioned have their appearance rather
similar to that in the recent members of the
Cupes and related genera, but (?) A. latidentata
comb.n. seems to be less slender, with the elytra
apparently somewhat gently sloping at sides
and comparatively widely explanate along their
lateral edges, the less expressed primary veins,
and apparently the very weak depression on the
frons. Nevetheless, because of many other sim-
ilarities in the structure of the head, the anten-
nae, the general body outline, and the shape of
elytra this species is temporaly also transferred
in the genus Apriacma gen.n. The holotype of
one of the species included in this new genus (A.
clavata comb.n.) has exposed aedeagus, which
is clearly different from that in Priacma serrata,
supporting the distinctness of the new genus
from Priacma.
6.2. Cretomerga gen.n.
Type species Priacmopsis subtilis Tan et
Ren, 2006.
Etymology. The name of this new genus is
formed from the name of the geological period
Cretaceous and generic root “merga” (“Bimer-
ga”, “Merga”, “Tenomerga”, “Cainomerga”).
Gender feminine.
Composition. The type species only.
Diagnosis. Body elongate of medium size
(11.5 mm). Integument with moderately coarse
sculpture and punctation; large subquadrangu-
lar (transverse) to polygonal cells of elytra ar-
ranged into nine long longitudinal rows inter-
spaced by weak secondary and well raised pri-
mary veins; A1 slightly deviating at scutellum
and almost completely rectilinearly reaching
elytral apex, unclearly fusing with CuA and then
with M before apex. Head looking like equilat-
eral triangle, with rather narrow neck (about
half as wide as head at temples), with expressed
Y-shaped depression, well projecting temples;
dorsal surface with weakly raised tubercles (para-
median longitudinal convexities along eye edge
to temples), antennal insertions moderately wide-
ly separated and apparently uncovered by tuber-
cles. Mandibles rather strong and arcuately
curved at outer edge before stout bidentate
apex. Antennae subfiliform and moderately long;
antennomere 1 (scape) subequal in length with
flagellomeres or much longer than antennomere
2 (pedicel). Pronotum subpentagonal and recti-
linearly widened anteriad, with smooth disc and
apparently widely explanate sides, arcuate ante-
rior angles and strongly convex anterior edge.
Part of prosternum before procoxae shorter than
procoxae. Elytra about 2.2 times as long as wide
combined, with broadly arcuate lateral edges,
subrounded to subacute apices and apparently
steeply (subvertically) sloping sides. Abdomi-
nal ventrites with a curved groove behind artic-
ulation, ventrite 1 subequal to clearly longer
than each of ventrites 2–4 and hypopygidium
120
A.G. Kirejtshuk et al.
somewthat longer than ventrite 1 and rounded at
apex. Legs moderately thin, frequently compar-
atively long; with protibia straight.
Comparison. The widely separated pro-
coxae of Cretomerga gen.n. are characteristic
of the members of the subfamily Triadocupedi-
nae, however this new genus can be scarcely put
in this group because nearly all its other charac-
ters are shared by other Cupedinae rather than
Triadocupedinae. It pertains to its slender body,
the elytra with nine long longitudinal rows of
cells, the steeply sloping sides and the very
narrow epipleura while the triadocupedines have
the wide and usually rather robust body, the
elytra with 11 rows of very large cells, the
widely explanate sides and the very wide epi-
pleura. It is necessary to note that Cretomerga
subtilis comb.n., in addition to the peculiar
procoxal cavities, has the distinctly raised pri-
mary veins characteristic of many taxa in Cupe-
dinae and Ommatinae, while most Triadocupe-
dinae demonstrate only the rows of cells (but not
well expressed veins). This new genus shares
most similarities with Furcicupes, differing from
the latter only in the structure of the prothoracic
segment, the shape of pronotum, and the widely
separated procoxae. The character to distin-
guish it among the taxa with the slender body
having the same number of the long longitudinal
row of cells and the expressed main veins of the
elytra (Adinolepis, Ascioplaga, Cupes, Cupop-
sis gen.n., Prolixocupes, Rhipsideigma, etc.) is
the clear Y-shaped depression on frons. This
last character is also present in Apriacma gen.n.,
Furcicupes, and Kirghizocupes, although Cre-
tomerga gen.n., except the widely separated
procoxae, differs from the first of these three
genera in the shorter prosternum before the
procoxae, and from the third in the pronotum
widening anteriad. The new genus under con-
sideration differs from Cupidium with unclear
number of the primary veins on the elytra and
the short prosternum before the procoxae at
least in the slender body with the rectilinear
pronotal sides and the conjointly subacute ely-
tral apices. Besides, this new genus has many
diagnostic features also shared by Apriacma
gen.n. (see the Diagnosis of the latter).
Note. Cretomerga gen.n. differs from Furci-
cupes mostly in the structure of the prothoracic
segment (see above), however other distinguish-
ing characters (shape of pronotum and widely
separated procoxae) seem to be also rather im-
portant to consider “Priacmopsis subtilis” as a
separate genus.
Cretomerga subtilis (Tan et Ren, 2006),
comb.n. Figs 77–79.
Priacmopsis subtilis Tan et Ren, 2006: 2655.
Specimen examined. Holotype “Priacmop-
sis subtilis” (“CNU-C-LB 2006003”, print and
counterprint).
Locality and stratigraphy. Beipiao City,
Chaomidian village, Liaoning Province, China;
early Cretaceous, Barremian-Aptian, Yixian
Formation.
Comments to the description. This species
was described without any mention of the char-
acters important to attribute it to any genus,
although many of them are quite clear in the
pictures accompaining the original desciption.
Рис. 77–83. Ископаемые виды подсем. Cupedinae: 77 — Cretomerga subtilis comb.n., голотип (“CNU-
C-LB 2006003-1”, отпечаток, сфотографированный на микроскопе Leica MZ 12.5 с камерой Nikon
Digital Camera DXM1200C); нижний мел, формация исянь; Китай, Ляонинь, Бейпяо; тело на сухом
отпечатке; 78 — то же, тело; прерывистой линией очерчены вентральные склериты головы, груди,
брюшка и прослеживающиеся контуры ног, не видные сверху; пунктиром Х-образное вдавление на
голове и медиальная выпуклость на переднеспинке; 79 — то же, тело отпечатка под спиртом; длина
экземпляра 10,3 мм; 80 — Cupopsis svitkoi (Lubkin, 2003), comb.n., голотип [“CUPC#1339”, сфотог-
рафированный на растровом электронном микроскопе (SEM), по Lubkin (2003)]; нижний мел, турон;
США, Нью Джерси, глины Олд Кроссмана; голова, антеродорсально; 81 — то же, спереди; 82 —
Cupopsis svitkoi (Lubkin, 2003), comb.n., дополнительный экземпляр [CUPC, без номера, сфотогра-
фированный на растровом электронном микроскопе (SEM), по Lubkin (2003)]; нижний мел, турон;
США, Нью Джерси, глины Олд Кроссмана; фрагмент обугленного надкрылья, латеродорсально; 83 —
то же, чешуйки на верхней поверхности надкрылий.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
121
Figs 77–83. Fossil species of the subfamily Cupedinae: 77 — Cretomerga subtilis comb.n., holotype
(“CNU-C-LB 2006003-1”, print, photographed under Leica MZ 12.5 with camera Nikon Digital Camera
DXM1200C); Lower Сretaceous, Yixian Formation; China, Liaoning, Beipiao; body on dry print; 78 —
idem, body with broken outlines of ventral sclerites head, thorax, abdomen and traced contours of legs
invisible dorsally; and dotted line X-shaped concavity on head and mediuan convexity on pronotum; 79 —
idem, body on print with alcohol; length of specimen 10.3 mm; 80 — Cupopsis svitkoi (Lubkin, 2003),
comb.n., holotype [“CUPC#1339”, microgram under scanning electron microscope (SEM), after Lubkin
(2003)]; Upper Cretaceous, Turonian; USA, New Jersey, Old Crossman’s Clay Pit; head, anterodorsal;
81 — idem, anteral; 82 — Cupopsis svitkoi (Lubkin, 2003), comb. n., additional specimen [CUPC, without
number, microgram under scanning electron microscope (SEM), after Lubkin (2003)]; Upper Cretaceous,
Turonian; USA, New Jersey, Old Crossman’s Clay Pit; fragment of a charcoalified elytron, laterodorsal;
83 — idem, scales on upper surface of elytra.
122
A.G. Kirejtshuk et al.
Most of these characters are indicated in the
above key to genera and subgenera and also in
the above diagnosis. The body size was men-
tioned correctly in the original description (Tan,
Ren, 2006: 11.5 mm), but not in the later mono-
graph (Tan, Ren, 2009: 10.27 mm). The print
and counterprint of the holotype demonstrate
nine long longitudinal rows of cells on the elytra
with only three clear rows externaly from M [not
“10” as written in the original description (Tan,
Ren, 2006) and in Tan & Ren (2009)], proster-
nal process between widely separated procoxae
without extending behind the procoxae (not
“extending behind coxae” as written in the orig-
inal description) and the scarcely visible gula
(not “rectangular” as written in the original
description). Besides, the upper surface of the
head of Cretomerga subtilis comb.n. has the
paramedian convexities along the eyes forming,
together with the anterior triangle of the frons,
the characteristic Y-shaped depression; the
pronotum with a median carina in the posterior
half and with a median triangular convexity in
the anterior one (these structures were misinter-
pretated in the drawings of the descriptors, see
Tan & Ren, 2006: 2656, Fig. 1, B, C); and the
elytra with reduced epipleura.
6.3. Cupopsis gen.n.
Type species Paracupes svetkoi Lubkin,
2003, fossil.
Etymology. The name is formed from the
generic name Cupes and Greek “opsis” (view,
sight, appearance). Gender feminine.
Composition. The type species only.
Diagnosis. Body apparently elongate of
medium size (around 10 mm). Integument with
moderately coarse sculpture and punctation;
large polygonal or oval cells of elytra arranged
into long longitudinal rows interspaced by weak
veins of subequal development. Head subtrian-
gular, with moderately projecting temples, very
large, rather wide and elongate eyes with trans-
verse diameter as a half of distance between
eyes; dorsal surface in the middle slightly con-
vex, with well raised tubercles over antennal
insertions and a deep depression between them,
and also concave along distal part of eyes.
Gently sloping elytral sides bearing three longi-
tudinal rows of cells in distal half of elytra.
Comparison. This new genus is well char-
acterized by the raised tubercles over the nar-
rowly separated antennal insertions and the con-
cave frons along the anterior part of the large
elongate eyes. The first feature and also the
elongate scales on the elytra make it possible to
suppose some close relation between it and
genera with steeply sloping elytral sides repre-
sented in the Recent fauna and apparently relat-
ed to the genus Cupes (Adinolepis, Ascioplaga,
Paracupes s.str., Paracupoides subgen.n., Pro-
lixocupes, and Rhipsideigma), however its very
large and elongate eyes together with the con-
cavity along the anterior part of the eyes clearly
separate this new genus from all others. The
mentioned structure of the head allows to distin-
guish Cupopsis gen.n. from the other genera of
the subfamily. The distal part of elytra with
three rows of cells along the sloping stripe
makes it possible to suppose that the complete
elytron should have only nine (but not 10) long
longitudinal rows of cells at least in the distal
half. The probable comparatively gentle slop-
ing elytral sides and apparently not strongly
expressed difference between at least the lateral
primary and the secondary veins provide an
opportunity to separate this new genus from
most probable relatives with such number of the
long longitudinal rows (genera Adinolepis, Apri-
acma gen.n., Ascioplaga, Cretomerga gen.n.,
Cupes, Furcicupes, Prolixocupes, Rhipsideig-
ma, Taxopsis gen.n., and probably Miocupes).
Note. This new genus represents some evi-
dence that a probable “ancestor” of most mem-
bers of the Cenozoic representatives of the
subfamily with tubercles over the antennal in-
sertions could appear not later than the late
Cretaceous (Turonian). Two features of the
recent “Cupes-complex” are rather crucial: sub-
vertical sloping sides of the elytra and close
(narrowly separated) antennal insertions. How-
ever the second could be a more reliable trace of
“ancestry” in comparison of the first, which
could be rather a sequence of homoplasy (see
below the Notes to Paracupoides subgen.n.).
Taxonomy of the reticulate beetles of the subfamily Cupedinae
123
Besides, it has also the medially convex frons
reaching a very short triangular depression be-
fore its anterior edge. This character seems to be
an autapomorphy of this genus.
The type species of this new genus cannot be
put in the genus Paracupes s.lato not only be-
cause of the characteristic structure of head, but
also because of the presence of only nine long
longitudinal rows of cells in the distal half of the
elytra while both Paracupes (Paracupes) bra-
siliensis and Paracupes (Paracupoides) ascius
bears 10 such rows of cells along the whole
length of its elytra.
Cupopsis svetkoi (Lubkin, 2003), comb.n.
Figs 80–83.
Paracupes svetkoi Lubkin, 2003: 190.
Material (after Lubkin, 2003). Holotype, a
three-dimensional fusainized (charcoalified)
beetle head, CUPC#1339; isolated cupedid
elytra are also present in the same sediments,
which can be preliminarily considered as re-
mains of the same species.
Locality and stratigraphy. Old Crossman’s
Clay Pit, South Amboy Fire Clay, Middlesex
County, New Jersey, USA; late Cretaceous Tu-
ronian; Raritan Formation.
6.4. Genus Paracupes Kolbe, 1898
Paracupoides subgen.n.
Type species Paracupes ascius Neboiss,
1989, recent.
Etymology. The name is formed from the
generic name “Paracupes” and Greek “ides” or
“eidos” (appearance, image, shape). Gender
masculine.
Composition. The type species only.
Diagnosis. Body elongate of medium size
(10.4 mm). Integument with moderately coarse
sculpture and punctation; large polygonal or
suboval cells of elytra arranged into 10 long
longitudinal rows interspaced by well raised
primary and weak secondary veins. Dorsum
with large scales: elongate (lanceolate) on head,
suboval on prothorax and subtriangular (widen-
ing towards truncate apex) on elytra. Scales on
elytra of different color (brownish and whitish),
forming on light field a pattern of three pairs of
oblique dark infuscations. Underside with scales
of different shape (elongate to oval or triangu-
lar) intermixed on different sclerites (median
ones with wider and lateral ones with scales).
Head subtriangular, with strongly projecting
temples; dorsal surface with a pair of elongate
and wide slightly elevated paramedian convex-
ity along most part of head and raised tubercles
over antennal insertions. Antennae subfiliform
and moderately long; antennomere 1 (scape)
longest and moderately thick (thickest), anten-
nomere 2 (pedicel) shortest and subconical
(slightly thicker than flagellomeres) and other
antennomeres (flagellomeres) subcylindrical.
Mandibles well developed, with arcuate outer
edge three-dentate apices. Maxillary palpi sub-
cylindrical, moderately long and slightly curved.
Mentum of usual shape and more than twice as
wide as long. Gular sutures moderately separat-
ed and subparallel-sided, anteromedian gular
plate almost twice longer than wide (1.9 times).
Pronotum nearly twice as wide as long, with
subtruncate base, bisinuate anterior edge, sub-
parallel and widely explanate sides, rather acumi-
nate anterior angles, a pair of suboval slightly
elevated paramedian convexities in anterior half
continuing distad as not wide and slightly ele-
vated median convex stripe. Part of prosternum
before procoxae much longer than procoxae.
Elytra more than 2.5 times as long as wide
combined, with subparallel lateral edges, sub-
vertically sloping sides along M; A1 curved in
anterior fifth (subparallely to short anal vein)
and in posterior fifth (at conjoining with CuA
and before at conjoining of A and CuA); apices
obliquely truncate and forming a clear sutural
angle; lateral edge simple and without sharp
teeth. Abdominal ventrite 1 clearly longer than
each of ventrites 2–4. Posterior fold of procoxae
directed anteriad and subtriangular. Legs mod-
erately thin, protibia curved at apex, tarsomeres
1–4 widely lobed.
Comparison. This subgenus can be distin-
guished after the above key to the genera. In
addition to this key Paracupoides subgen.n.
differs from the nominative Paracupes s.str. in
124
A.G. Kirejtshuk et al.
Figs 84–86. Paracupes (Paracupes) brasiliensis Kolbe, 1908 (MNHN, “Brasil, (?) Mur” “ex Museo
Mniczech” “coll. R. Oberthur”, photographed under Olympus SCX9) with camera Olympus; recent: 84 —
body, dorsal; 85 — pro- and mesothoraces, lateroventral; 86 — elytral apex of high magnification with clear
scales, dorsal. Length of specimen 16 mm. Abbreviation: Pcx. l. — procoxal lobe.
Рис. 84–86. Paracupes (Paracupes) brasiliensis Kolbe, 1908 (MNHN, “Brasil, (?) Mur” “ex Museo
Mniczech” “coll. R. Oberthur”, сфотографированный на микроскопе Olympus SCX9) с камерой
Olympus; современный: 84 — тело, сверху; 85 — передне- и среднегрудь, лптеровентрально; 86 —
вершина надкрылья при большом увеличении с отчетливыми чешуйками, сверху. Длина экземпляра
16 мм. Сокращение: Pcx. l. — прококсальная лопасть.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
125
Figs 87–90. Paracupes (Paracupoides) ascius Neboiss, 1989, holotype (MNHN, “Equateur Or., de Baсos
а Canelos, M. de Mathan, IX-X.1894”, photographed under Olympus SCX9 with camera Olympus); recent:
87 — body, dorsal; 88 — head and thorax, ventral; 89 — elytral apices, dorsal; 90 — elytral apex of high
magnification with clear scales, dorsal. Length of specimen 10.4 mm.
Рис. 87–90. Paracupes (Paracupoides) ascius Neboiss, 1989, голотип (MNHN, “Equateur Or., de Baсos
а Canelos, M. de Mathan, IX-X.1894”, сфотографированный на микроскопе Olympus SCX9 с камерой
Olympus); современный: 87 — тело, сверху; 88 — голова и грудь, снизу; 89 — вершины надкрылий,
сверху; 90 — вершина надкрылья при большом увеличении с отчетливыми чешуйками, сверху.
Длина экземпляра 10,4 мм.
the less developed mandibles [in Paracupes
(Paracupes) brasiliensis they are longer and
with a subangular outer edge], the surface of
pronotum [in P. (P.) brasiliensis it has a median
narrow and slightly elevated stripe], difference
in the shape of scales on the head, pronotum and
elytra [dorsum of P. (P.) brasiliensis is uniform-
ly covered with narrow lanceolate scales (partly
with truncate apices — see Figs 84–86 and
Vulcano & Pereira, 1975) intermixed by thick
hairs], the elytral venation (besides the weak
primary veins, A1 of P. (P.) brasiliensis is
126
A.G. Kirejtshuk et al.
almost subparallel to the suture and only slightly
deviating in the anterior fifth, the short anal vein
at the smaller distance from the suture), the
smaller cells in four long longitudinal rows
along the elytral sides and the larger cells in six
rows on the disk; the posterior fold of procoxae
[it in P. (P) brasiliensis has subtrapezoid shape,
but not subtriangular]. The pattern of infusca-
tion on the dorsum of P. (P.) brasiliensis is quite
variable and such pattern in P. (Paracupoides)
ascius fits in the range of pattern variation of the
first. The legs of these species are rather similar,
including the level of curvation of protibiae.
Besides, Paracupoides subgen.n. differs
from the other genera of the subfamily in the
truncate elytral apices, A1 not parallel to the
suture in the proximal and distal fifths and
triangular scales on elytra.
It also differs from:
– Adinolepis and Ascioplaga in the not par-
allel long anal veins (A1), 10 complete long
longitudinal rows of cells on the elytra and the
lack of tarsal grooves on the ventral surface of
the prothorax;
– Apriacma gen.n. in the smaller mandibles,
10 complete long longitudinal rows of suboval
cells on the elytra, A1 not rectilinear before the
elytral apex, the antennal insertions covered by
tubercles, the protibia curved at apex and the
wide tarsi;
– Chalepocarabus in the well raised primary
veins, the subparallel-sides elytral edges, the
larger cells on elytra, the transverse pronotum,
the antennal insertions covered by the tubercles,
the longer antennae with much larger scape and
the larger eyes;
– Cretomerga gen.n., Furcicupes and Gra-
cilicupes in the tubercles over the antennal in-
sertions (antennal insertions of Cretomerga
gen.n., Furcicupes, and Gracilicupes species
are open), the longer prosternum before procox-
ae, not rectilinear A1 before the elytral apex, the
curved apex of the protibiae and the wide tarsi;
and also from two first groups in the wide neck
and the lack of Y- or V-shaped median depres-
sion on the frons and 10 long longitudinal rows
of cells on the elytra; from the third in the
smaller eyes and the wider pronotum with the
rectilinear sides and the projecting anterior an-
gles (pronotum of Gracilicupes species is nar-
rower than head and arcuate at sides); and also
from Gracilicupes in the Cupes-type aedeagus;
– Cupes in the pair of longitudinal convex-
ities along the vertex and frons, 10 complete
long longitudinal rows of cells on the elytra, not
parallel long anal veins (A1) and the lack of
antennal grooves and the rather curved apex of
protibiae (protibiae of Cupes species are straight
or slightly curved at apex);
– Cupopsis gen.n. in the pair of longitudinal
convexities on the head, the subhemispherical
eyes and the not concave frons along the anteri-
or part of the eyes and 10 complete long longi-
tudinal rows of cells on the elytra;
– Cupidium in the somewhat more slender
body with the rectilinear pronotal sides and the
conjointly subacute elytral apices;
– Kirghizocupes in the much shorter head
with the larger eyes, the narrower pronotum
with the median carina, not rectilinear A1 be-
fore elytral apex;
– Latocupes in the slender body with the
subparallel elytral lateral edges, not rectilinear
A1 before elytral apex, polygonal cells on the
elytra (not transverse as usually in Latocupes
species), the antennal insertions covered by the
tubercles and the lack of Y- or V-shaped median
depression on the frons, the protibia curved at
apex;
– Mesocupes in the antennal insertions cov-
ered by the tubercles, the subparallel and steep-
ly sloping sides of elytra, not rectilinear A1
before elytral apex, the shorter and more stout
legs with the curved protibiae;
– Miocupes in the much smaller head with
the larger eyes, the markedly longer elytra with
a great difference in development of the primary
and secondary veins, and also the fusion of A1
with CuA before apex;
– Priacma in the elytra with the well ex-
pressed primary veins and the steeply sloping
sides, the absence of the preapical closing fold on
below sides of the elytra, the narrowly separated
gular sutures, the longer abdominal ventrite 1, the
curved protibiae and the Cupes-type aedeagus;
– Priacmopsis in the much smaller and slen-
der body, the head with the larger eyes, the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
127
larger and more projecting temples, and the
protibiae curved at apex;
– Prolixocupes in 10 complete long longitu-
dinal rows of cells on the elytra, A1 curved
along the short row of cells at scutellum, the
head convexities not divided into tubercles over
the antennal insertions and the posterior ones,
the separated gular sutures, the unexpressed
tarsal grooves on prothorax and the protibia
curved at apex;
– Rhipsideigma in 10 complete long longitu-
dinal rows of cells on the elytra, the very weak
tubercles (convexities) on the head, the anal
long vein deviating in the anterior and posterior
fifths, the fusion of A1 and CuA, the lack of
sharp process at the place of fusion of veins;
– Taxopsis gen.n. in the smaller and more
slender body, the narrower prosternal process,
the not strongly transverse cells on the elytra, 10
complete long longitudinal rows of cells on the
elytra, not rectilinear A1 before elytral apex and
the protibiae curved at apex.
Note. Paracupes (Paracupoides) ascius
seems to be closely related to P. (Paracupes)
brasiliensis. The taxonomic separation is pro-
posed to reach a balanced partition of recent
representatives of the subfamily, because the
characters separating the mentioned species are
as important as those between the other genera
of the subfamily. Moreover, differences be-
tween some genera used for recent forms are
even much less expressed in comparison with
those between Paracupes s.str. and Paracu-
poides subgen.n., reducible to the unique pres-
ence versus absence of scales on the dorsum, the
shape of pregenital sclerites of the male, and the
geographic distribution (Neboiss, 1984; Kirej-
tshuk, 2005). On the other hand, this case of two
close relatives in the Recent fauna manifesting
very expressive “generic” differences can be
compared with the presence of different similar
forms in the past, beginning from the Middle
Triassic. One relative [Paracupes (Paracupes)
brasiliensis] has great similarities with recent
Priacma and with other taxa that are already
extinct (for instance, members of Mesocupes
s.str. and Cainomerga subgen.n. with 10 long
longitudinal rows of cells on the elytra com-
pared to those in members of Latocupes with
nine rows of cells), particularly in the gently
sloping sides and the not regular venation on the
elytra. Also the thickened and serrate elytral
sides of P. (P.) brasiliensis are rather similar to
those in Priacma than in any other cupedines.
Another relative [Paracupes (Paracupoides)
ascius] sharing morphological features with this
P. (P.) brasiliensis (head and appendages, ex-
cept the gula and the level of development of
mandibles, the truncate elytral apices, the shape
of most thoracic sclerites, legs, etc.), although at
the same time its important (subgeneric) pecu-
liarities are shared by the members of the genera
near Cupes with the more steeply sloping elytral
sides (general appearance, and particularily the
elytral structure, viz. the strongly raised primary
veins, and subverticaly sloping sides). Finally,
Paracupes (Paracupoides) ascius is character-
ised by the unique structure of scales on its
integument. At the same time the male genitalia
of P. (P.) brasiliensis can be regarded as Cupes-
type with the rather short appendages at base of
parameres (Vulcano, Pereira, 1975) which could
be regarded as plesiotypic features among clos-
er relatives of Cupes and somewhat reminiscent
homologous structures in the male genitalia of
Priacma, these processes demonstrate a great
variability among other species of the Recent
fauna (Neboiss, 1984).
Paracupes (Paracupoides) ascius (Neboiss,
1989) Figs 87–90.
Holotype. “Paracupes ascius sp. n., det.
Neboiss 1988, HOLOTYPE”, $, “Equateur Or.,
de Baños à Canelos, M. de Mathan, IX-X.1894”
(MNHN).
Addition to the description. Elytra thor-
oughly covered with subtriangular scales (wid-
ened apically) but also bearing short and fine
hairs along the sutural and lateral edges; dorsal
integument of prothorax and head covered with
similar scales which are somewhat intermixed
by more or less elongate (lanceolate) and usual-
ly apically truncate scales, along anterior edges
of pronotum and head, as well as on labrum and
mandibles with long and thick hairs; underside
128
A.G. Kirejtshuk et al.
covered partly with subtriangular scales, partly
with elongate ones, but epipleura covered com-
pletely with very long and dilated hairs (scales);
appendagesmostly with thickandcontrast hairs,
although femora and proximal part of tibiae
with clear elongate scales. Cells disposed in six
long longitudinal rows on elytral disk markedly
larger than those in four rows along sloping
sides. Primary veins (A1, CuA and M) well
raised. Anteromedian gular plate almost twice
as long as wide (1.9 times). Antennal grooves of
prothorax rather shallow and scarcely outlined.
Procoxal plates trapezium-like and covering
only proximal part of protrochanter. Abdominal
ventrites with tegular articulation and showing
clear roller at apex and following transverse
depression at base of ventrites.
6.5. Taxopsis gen.n.
Type species Cupes motschulskyi Kirejtshuk,
2005, fossil.
Etymology. The name of this new genus is
formed from the Greek “taxis” (row, order) and
“opsis” (view, sight, appearance). Gender fem-
inine.
Locality and stratigraphy. Baltic amber;
probably Kaliningrad Region, Russia; late
Eocene, Bartonian/Priabonian, Prussian For-
mation.
Diagnosis. Body elongate of medium size
(16.1–18.1 mm). Integument with moderately
coarse sculpture and punctation; large subqua-
drangular transverse cells of elytra, which are
twice as wide as long, arranged into nine long
longitudinal rows interspaced by weak second-
ary and well raised primary veins; A1 slightly
deviating at scutellum and almost completely
rectilinearly reaching elytral apex, clearly fus-
ing with CuA and then with M before apex.
Head transverse and triangular, with moderate-
ly wide neck (about 3/4 as wide as head at
temples), without expressed Y-shaped depres-
sion, well projecting temples; dorsal surface
with two pairs of weakly raised paramedian
tubercles (at antennal insertions and at level of
posterior edge of eyes), antennal insertions
moderately narrowly separated and covered by
tubercles. Mandibles moderately strong and ar-
cuately curved. Antennae subfiliform, moder-
ately long and thin; antennomere 1 (scape) sub-
equal in length with flagellomeres and very
swollen, antennomere 2 (pedicel) shortest and
nearly 1/3 as long as antennomere 3. Pronotum
subtrapezoidal, anterior and posterior edges
curved, lateral sides subangular, widening from
base to middle and subparallel-sided in anterior
half, with smoothed disc divided by median
carina and apparently widely explanate sides,
anterior and posterior angles with distinct top.
Part of prosternum before procoxae about 2.5
times as long as procoxae, apex of its intercoxal
process widely rounded and about as wide as
procoxae. Elytra about twice as long as wide
combined, with broadly arcuate lateral edges,
subrounded to subacute apices and sides steeply
(subvertically) sloping along M. Hypopygidi-
um about twice as long as each of ventrites 2–4,
somewhat longer than wide and narrowly round-
ed to angular at apex. Legs moderately thin,
frequently comparatively long; with protibia
nearly straight.
Comparison. The genus Taxopsis gen.n.
can be easily diagnosed after the above key to
genera and subgenera. It shows so great similar-
ity with Cupes that the comparison of the latter
somehow fit also to this new genus. Neverthe-
less, this genus has the very prominent charac-
ters making it possible to regard it as a separate
genus with a distinctness equal to that of other
genera. The rather wide prosternal process of T.
motschulskyi comb.n. with the widely rounded
apex is the unique character among cupedines,
although widely separate coxae are also known
in Cretomerga gen.n. (Cupedinae) and all taxa
of the subfamily Triadocupedinae. The width of
procoxae in Taxopsis gen.n. is also nearly the
greatest among the cupedines. The strongly
transverse subquadrangular cells of elytra rep-
resent another important feature of T. motschul-
skyi comb.n., which is intrinsic only for species
of the genus Latocupes among cupedines, how-
ever the latters have the clear Y-shaped depres-
sion on the subtriangular head (which is not so
tranverse as in Taxopsis gen.n.). The elytra of
some species of Cainomerga subgen.n. and
Taxonomy of the reticulate beetles of the subfamily Cupedinae
129
Cupes also show a tendency to transform shape
of cells in a transverse type, however these
species have cells only slightly wider than long
(not twice as wide as long). Also the elytron of
Cainocups aixensis gen. et sp.n. (a taxon with
unclear position, see above) has transverse oval
cells, however the pattern of its veins is com-
pletely different.
7. Subfamily incertae sedis
7.1. Cainocups gen.n.
Type species Cainocups aixensis sp.n., fossil.
Etymology. The name of this new genus is
formed from the geological term Cainozoic or
Cenozoic (Greek καινός kainos “new”, and ζωή
zoe “life”) and abbreviation of the generic name
“Cupes”. Gender masculine.
Diagnosis. Distal part of elytron with 12
long longitudinal rows of transversely oval cells
interspaced by the weak secondary and the well
raised primary veins; R and M fusing first and
joint vein fusing at first with CuA and then with
A1 (Fig. 76). Elytral side apparently steeply
(subvertically) sloping without lateral expla-
nation.
Comparison. The sequence of the fusion of
the primary veins in Cainocups aixensis gen. et
sp.n. is very unusual and does not occur not only
among members of the subfamily Cupedinae,
but also among other two subfamilies of Cupe-
didae. The generalized type of fusion of primary
veins of the family includes the fusion of A1
with the arcuately curved CuA at elytral apices
and in many cases to this join vein (after fusion
of A1 and CuA) arcuately curved M can be
attached or fused (most genera of all subfami-
lies). However among many species of Noto-
cupes (Ommatinae) this type of fusion is also
observed and a tendency for independent fusion
of A1 with CuA and also the fusion of M and R
can be traced. Ponomarenko (2006) reconstruct-
ed the ground-plan of the elytron for Zygadenia
Handlirsch, 1906 in which he drew a sequence
of the fusion of the primary veins at elytral apex
including the first fusion of M and CuA and the
second fusion of this join vein and A1 (Pono-
marenko, 2006: 96, Fig. 4).
The number of the long longitudinal rows of
cells on the elytra of Cainocups gen.n. (12) is
unique among the cupedines. The number of
these rows of cells in the family varies between
nine and 11, although the usual ones are nine or
10 (see above the Notes on the structural pecu-
liarities of the family).
The pattern of the elytral venation of Cain-
ocups gen.n. makes it possible to certainly de-
fine the attribution of this genus to the family
Cupedidae, as the fusion of the primary veins at
elytral apices is characteristic of this family.
However because of uniqueness of the vein
pattern on elytra this new genus cannot be put in
any subfamily with confidence. Nevertheless
some analogical features in the elytral venation
of Cainocups gen.n. can be noticed in that of
Menatops gen.n. (see above), particularly in
well raised R and M, fusion of M with A1 after
the fusion of M with CuA (Figs 10–11). This
similarity could be used for putting the new
genus under consideration into the subfamily
Cupedinae. The apparently steeply sloping of
the elytral side along R in this new genus is also
the unique feature (usually such sloping occurs
along M) and gives no argumentation for a more
precise attribution of this new genus. The next
feature which could be considered in this con-
text is the type of sloping of elytral side. The
steep (subvertical) sloping is observed in both
Cupedinae and Ommatinae, however among the
latter there are many forms combining the sub-
vertical sloping and the lateral explanation of
elytral sides. The very diverse genus Notocupes
includes representatives with somewhat similar
patterns of the elytral venation, although they
also show the clear lateral explanation along the
lateral edge of the elytra [for example, N. eumera
(Tan, Ren et Shih, 2006), N. porrectus (Tan, Ren
et Shih, 2006), and N. stabilis (Tan, Ren et Liu,
2005) with independent fusion of A1 and CuA as
well as fusion of M and R; etc.]. Thereby the
position of the genus Cainocups gen.n. should be
interpreted as uncertain among the Cupedidae.
Taking into consideration the length of the
fragment of the elytron of the holotype of Cain-
ocups aixensis gen. et sp.n., the probable length
of the whole beetle, to which it belonged, can be
estimated as approximately 15 mm.
130
A.G. Kirejtshuk et al.
Cainocups aixensis sp.n.
Fig. 76.
Holotype. Specimen MNHN-F-A 51230
(“Aix 18, Nel leg”). The specimen is represent-
ed by a fragment of the elytron in the piece of
comparatively hard rock.
Locality and stratigraphy. Palaeogene,
Latest Oligocene; France, Jas de Bouffan, town
Aix-en-Provence, Bouches-du-Rhône.
Note. This fossil fragment of the elytron is
the unique specimen of this species among 10
thousands of beetles collected in the same lay-
ers. Cainocups aixensis sp.n. seems to have
been an extremely rare taxon in deposits of the
latest Oligocene (only known in its type deposit
and known by only one specimen among 50000
fossil insects).
Description. Length of print of elytron 6.3
mm. R almost gradually arcuate, meeting with
sutural edge just at elytral apex and dividing
elytron into a flat disk and a steeply sloping side
(Fig. 76). Eight long longitudinal rows on ely-
tral disk consisting of transversely elliptic cells
more than twice as wide as long, interspace
between them in rows about as great as length of
cells; cells on sloping part of elytron not so
strongly transverse to oval, somewhat smaller
and apparently shallower.
8. Lists of fossil taxa re-examined
The generic and subgeneric taxa mentioned
in the below lists include only those that are
possible to interpret after study of the specimens
and original descriptions with informative illus-
trations. In cases when the authors of this paper
used only the descriptions without checking
specimens the corresponding publications cited
by indication of the authors’ name(s) and year
after the Latin names of taxa are given in the
References. Some taxa are omitted in this ap-
pendix because of difficulties of interpretation
on the available data, particularly those pro-
posed after isolated elytra or described with
considerable deficiences. Some information on
the taxa that remained without consideration
can be obtained in the catalogue by Kirejtshuk
& Ponomarenko (2016).
8.1. List of fossil genera and subgenera of
Cupedinae
Apriacma gen.n.
(see above Diagnosis of this genus)
– Apriacma clavata (Tan, Ren et Shih, 2006),
comb.n. [Priacma]; early Cretaceous, Barremi-
an-Aptian, Yixian Formation; China, Liaoning,
Beipiao.
– (?) Apriacma latidentata (Tan, Ren et
Shih, 2006), comb.n. [Priacma]; early Creta-
ceous, Barremian-Aptian, Yixian Formation;
China, Liaoning, Beipiao.
– Apriacma renaria (Tan, Ren et Shih, 2006),
comb.n. [Priacma]; early Cretaceous, Barremi-
an-Aptian, Yixian Formation; China, Liaoning,
Beipiao.
– Apriacma tuberculosa (Tan, Ren et Shih,
2006), comb.n. [Priacma]; early Cretaceous,
Barremian-Aptian, Yixian Formation; China,
Liaoning, Beipiao.
Genus Chalepocarabus Handlirsch, 1906
(after Ponomarenko, 2006)
This genus was proposed in the short notes
of the monograph of Handlirsch, 1906 (1906–
1908), and later Crowson (1962) and Ponomar-
enko (1969, 2006) re-defined its position. The
interpretation of the last publication (Ponomar-
enko, 2006) is recognized in this paper. The
main features of this genus are the very long
pronotum and comparatively small cells dis-
posed in 10 long longitudinal rows between
unexpressed veins on elytra. Besides, this genus
has the gently sloping elytral sides, the some-
what tuberculate upper surface of the head, and
the rather separated antennal insertions. Some-
how this group seems to be close to Mesocupes
(particularly could be close to Anaglyphites stat.n.)
differing from the latter mostly in the shape of its
long pronotum, the very obliterated elytral integ-
ument and the rather small cells on the elytra.
– Chalepocarabus elongatus (Brodie, 1845)
[Carabus]; early Cretaceous (unless Late Juras-
sic), England, Wiltshire, Early Purbeck, Vale of
Wardour.
Cretomerga gen.n.
(see the above Diagnosis of this genus)
– Cretomerga subtilis (Tan et Ren, 2006),
comb.n. [Priacmopsis]; early Cretaceous, Bar-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
131
remian-Aptian, Yixian Formation; China, Lia-
oning, Beipiao.
Genus Cupes Fabricius, 1801
(= Cupoides Motschoulsky, 1856; Tenom-
erga Nebois, 1984; Distocupes Neboiss, 1984)
(see the above Diagnosis of this genus)
– Cupes distinctissimus sp.n.; Paleogene,
Paleocene; France, Puy-de-Dome, Menat.
– Cupes eckfeldensis (Tröster, 1993),
comb.n. [Tenomerga]; Paleogene, Middle
Eocene; Germany, Rheinland-Pfalz, Rhine Pa-
latine, Eckfelder Maar.
– Cupes groehni Kirejtshuk, 2005; Paleo-
gene, late Eocene; Baltic See coast, Russia, near
Kaliningrad (formerly Königsberg), Jantarny,
Baltic amber.
– Cupes hoffeinsorum Kirejtshuk, 2005;
Paleogene, late Eocene; Baltic See coast, Rus-
sia, near Kaliningrad (formerly Königsberg),
Jantarny, Baltic amber.
– Cupes kerneggeri Kirejtshuk, 2005; Pa-
leogene, late Eocene; Baltic See coast, Russia,
near Kaliningrad (formerly Königsberg), Jan-
tarny, Baltic amber.
– Cupes komissari Kirejtshuk, 2005; Paleo-
gene, late Eocene; Baltic See coast, Russia, near
Kaliningrad (formerly Königsberg), Jantarny,
Baltic amber.
– Cupes manifestus Kirejtshuk, Nel et Co-
lomb, 2010; Paleogene, Paleocene; France, Puy-
de-Dome, Menat.
– Cupes messelensis (Tröster, 1993),
comb.n. [Tenomerga]; Paleogene, Middle
Eocene; Germany, 9 km NE Darmstadt, Messel.
– Cupes ponomarenkoi Kirejtshuk, Nel et
Colomb, 2010; Paleogene, earliest Eocene;
France Paris Basin, Le Quesnoy, Oise amber.
– Cupes praeglacialis Gersdorf, 1976; Neo-
gene, late Pliocene; Germany, Niedersachsen,
Harz, Willershausen.
– Cupes rohdendorfi Iablokoff-Khnzorian,
1960; Paleogene, late Eocene; Baltic See coast,
Russia, near Kaliningrad (formerly Königsberg),
Jantarny, Baltic amber.
– Cupes simillimus sp.n.; Paleogene, Pale-
ocene; France, Puy-de-Dome, Menat.
– Cupes tesselatus (Motschoulsky, 1856)
[Cupoides]; Paleogene, late Eocene; Baltic See
coast, Russia, near Kaliningrad (formerly
Königsberg), Jantarny, Baltic amber.
– Cupes weitschati Kirejtshuk, 2005; Paleo-
gene, late Eocene; Baltic See coast, Russia, near
Kaliningrad (formerly Königsberg), Jantarny,
Baltic amber.
Genus Cupidium Ponomarenko, 1968
This group, on the one hand, is similar to
Cretomerga gen.n., Gracilicupes and Furci-
cupes by its very short prosternum before the
procoxae, but differs from all of them in the
more robust body with the elytra, which are
somewhat less than twice as long as wide com-
bined, widely and separately rounded at apices
and gently sloping at sides, and the compara-
tively small eyes; moreover Cupidium differs
also from the first in the narrowly separated
procoxae; from the second in the not so narrow
prothorax (which is about as wide as head) and
neck; and from the third in the arcuate sides of
pronotum and the not so narrow neck. On the
other hand, Cupidium abavum could be consid-
ered close of the genus Mesocupes, having the
general appearance and the subparallel-sided
elytra as those in Mesocupes s.str. Finally the
single species of the genus demonstrates the
separately rounded elytral apices forming a com-
paratively deep sutural angle which make this
genus quite distinct from all mentioned taxa.
– Cupidium abavum Ponomarenko, 1968;
late Jurassic, Oxfordian (or/and Kimmeridgian);
Kazakhstan, Chimkent Region, Kara-Tau Range,
Karatau.
Cupopsis gen.n.
(see above Diagnosis of this genus)
– Cupopsis svitkoi (Lubkin, 2003), comb.n.
(Paracupes); late Cretaceous, Turonian; USA,
New Jersey, Middlesex County, Old Cross-
man’s Clay Pit.
Genus Furcicupes Tan et Ren, 2006
This genus is characterized by the very slen-
der body, the head with the clear Y-shaped
depression and the rather narrow neck, the very
short prosternum before the procoxae and ap-
parently nine rows of cells on the elytra. The
132
A.G. Kirejtshuk et al.
Figs 91–94. Fossil species of the subfamily Cupedinae, photographed under Leica MZ 12.5 with camera
Nikon Digital Camera DXM1200C: 91 — dry print of holotype of Furcicupes raucus Tan et Ren, 2006
(“CNU-C-LB2006004”); Lower Сretaceous, Yixian Formation; China, Liaoning, Beipiao; length of body
9.2 mm; 92 — idem, covered by alcohol; 93 — dry print of holotype of Gracilicupes tenuicruralis Tan, Ren
et Shin, 2006 (“CNU-C-LB2005002”); Middle Jurassic, Jiulongshan Formation; Inner Mongolia; Chifeng
City, Daohugou; dry print, length of body 11.4 mm; 94 — idem, covered by alcohol. Abbreviation: aed —
aedeagus.
Рис. 91–94. Ископаемые виды подсем. Cupedinae, сфотографированные на микроскопе Leica MZ 12.5
с камерой Nikon Digital Camera DXM1200C: 91 — сухой отпечаток голотипа Furcicupes raucus Tan
et Ren, 2006 (“CNU-C-LB2006004”); нижний мел, формация Исянь; Китай, Ляонинь, Бейпяо; длина
тела 9,2 мм; 92 — то же, покрытый спиртом; 93 — сухой отпечаток голотипа Gracilicupes tenuicruralis
Tan, Ren et Shin, 2006 (“CNU-C-LB2005002”); средняя юра, формация Джулиньшан; Внутренняя
Монголия; Чифен, Даохугоу; сухой отпечаток, длина теля 11,4 мм; 94 — то же, покрытый спиртом.
Сокращение: aed — эдеагус.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
133
neck of this genus is the narrowest among cu-
pedines, although it can be somewhat compared
with that in Gracilicupes. The very short pros-
tenum before the procoxae is known also for
genera Cupidium, Cretomerga gen.n. and Gra-
cilicupes, although Furcicupes differs from the
first in the more slender body, the longer elytra
with the steeply sloping sides, the larger eyes;
from the second in the narrowly separated pro-
coxae and the open antennal insertions; and
from third in the wider prothorax. Tan & Ren
(2006) regarded that the type species of this
genus has eight long longitudinal rows of cells
on its elytra, however, the re-examimation of
this specimen showed that there are three rows
of cells located externally from M (but not two:
Fig. 91).
– Furcicupes raucus Tan et Ren, 2006; early
Cretaceous, ? Barremian-Aptian, Yixian For-
mation; China, Liaoning, Beipiao.
Genus Gracilicupes Tan, Ren et Shin, 2006
This genus was proposed for two species
having a rather slender body; the large head with
the comparatively narrow neck, very large eyes,
long antennae; very narrow pronotum with the
arcuate sides (markedly narrower than head),
not projecting anterior and posterior angles;
very short part of prosternum before the procox-
ae, the elytra with 10 long longitudinal rows of
cells, the well raised primary veins different in
expression from the secondary veins, and the
steeply sloping sides. The most distinct charac-
ter of this genus is the prothorax, which is much
narrower than head . The very short prosternum
is characteristic of not only this genus, but also
of Cretomerga gen.n., Cupidium, and Furci-
cupes (see above the characteristics of these
genera and the above key to genera and subgen-
era). This genus demonstrates some similarities
to the structures of the “complex of genera”
closely related to Cupes with the slender body
and nine long longitudinal rows of cells on the
elytra.
– Gracilicupes crassicruralis Tan, Ren et
Shin, 2006; Middle Jurassic, Jiulongshan For-
mation; Inner Mongolia; Chifeng City, Dao-
hugou.
– Gracilicupes tenuicruralis Tan, Ren et
Shin, 2006 (=tenuocruralis: Tan et Ren, 2009);
Middle Jurassic, Jiulongshan Formation; Inner
Mongolia; Chifeng City, Daohugou.
Genus Kirghizocupes Ponomarenko, 1966
(=Mesocupoides Ponomarenko, 1969,
syn.n.)
This genus represents the oldest group of the
cupedines in the fossil record, with the rather
slender and very convex body and the steeply
sloping sides of the elytra. It is somehow more
similar to Priacma because of 10 long longitu-
dinal rows of cells on the elytra, which are
apparently as flat as in Cupes, and, in addition to
these rows of cells, one additional row is visible
just between Sc and edge of elytra. This genus is
characterized by the very peculiar elongate head
with a pair of the long paramedian convexities
and the open antennal insertions, the rather wide
pronotum with the very widely (sub)explanate
sides, the comparatively medium body-size
(length 5.0–7.5 mm), and particularly with the
slightly different primary and secondary veins
gradually converging at apex, but without clear
fusion.
The type species of both Kirghizocupes (K.
cellulosus) and Mesocupoides (M. proporeius)
are certainly rather closely related and mostly
distinguished by the characters of the shape of
the head and pronotum, the length of antennae
and also in the different distances between
procoxae (a little greater in Kirghizocupes and
a little smaller in Mesocupoides). Therefore
they can be scarcely considered as separate
genera. Kirghizocupes could be placed among
the members of Cupedinae but not Triadocupe-
dinae, although the distance between the pro-
coxae in Kirghizocupes cellulosus is not very
narrow (Ponomarenko, 1966a: 60). The elytra
of the holotypes of the type species of both
“genera” have steep subvertical sides with sharp
inflection externally from seventh long rows of
cells (Figs 109–111), although some specimens
demonstrate not so sharp inflection or even
smooth but with distinct long longitudinal rows
of cells on the rather wide sloping lateral stripes
of the elytra (Fig. 116).
134
A.G. Kirejtshuk et al.
This genus unites some species from the
same localities and some of them were de-
scribed in the genera here synonymized and,
perhaps, partly in genera Asimma and Ptero-
cupes (see below). Some species probably relat-
ed to this group and deposited in the collection
of PIN still remain undescribed. Among mem-
bers of this group two subgroups can be defined
according to the length of the ultimate abdomi-
nal ventrite (hypopygidium) which looks in some
cases shorter, but in other cases divided into two
parts (two sclerites).
– Kirghizocupes cellulosus Ponomarenko,
1966; ? Middle–late Triassic, Ladinian-Kar-
nian; Kyrgyzstan, Osh Region, Djailaucho
(Dzhayloucho), Madygen.
– Kirghizocupes proporeius (Ponomaren-
ko, 1969), comb.n. [Mesocupoides]; ? Middle–
late Triassic, Ladinian-Karnian; Kyrgyzstan,
Osh Region, Djailaucho (Dzhayloucho), Mady-
gen.
– Kirghizocupes indistinctus (Ponomaren-
ko, 1969, comb.n. [Mesocupoides]; ? Middle–
late Triassic, Ladinian-Karnian; Kyrgyzstan,
Osh Region, Djailaucho (Dzhayloucho), Mady-
gen.
Genus Latocupes Tan et Ren, 2006
(? = Pulchicupes Ren, 1995)
The name Latocupes was synonymized by
Kirejtshuk et al. (2010) to Priacmopsis, howev-
er it is possible to distinguish the latter and
Latocupes by the V- or Y-shaped depression on
frons, which is lacking in Priacmopsis and, in
addition to this, Latocupes has the very charac-
teristic transverse cells on the elytra. These
characters combining with the comparatively
robust body and the widely transverse pronotum
could make it possible to synonymize it with
Pulchicupes described as a member of the fam-
ily Taldycupedidae (Ren et al., 1995). Howev-
er, the drawing in the original description of
Pulchicupes jiensis (Ren et al., 1995) shows 10
long longitudinal rows of transverse cells on
elytra, but the photographs accompanying it do
not allow verifying the exact number of rows of
cells. Therefore synonymy of these generic
names is regarded as preliminary and should be
clarified after a further re-examination of the
type specimen of Pulchicupes jiensis. Due to
the considered characters the species from Dao-
hugou described as members of Mesocupes
were transferred into the Latocupes because at
least in ‘Mesocupes collaris’ the V- or Y-de-
pression on the head and other characters of this
genus can be easily traced and according to the
original descriptions this character was observed
in other species described as Mesocupes from
Daohugou. Besides, Latocupes is characterized
by nine long longitudinal rows of cells on the
elytra (clearly visible in the holotypes of Lato-
cupes angustilabialis comb.n., L. bella and L.
fortis) within the primary veins, which some-
times seem to be slightly expressed. Tan & Ren
(2006) indicated 10 long longitudinal rows for
the species described by them in the composi-
tion of the genus Latocupes and later Tan et al.
(2007) mentioned nine rows for species de-
scribed by them as members of the genus “Me-
socupes”. In the latter paper there are some
photographs and drawings of cells at the elytral
sides which are not quite strongly transverse,
but the cells on the elytral discs in these speci-
mens are about twice as wide as long.
– (?) Pulchicupes jiensis Ren, 1995; early
Cretaceous, Lushangfen Formation; China,
Western Beijing Province, Jingxi Basin, Lush-
angfen village.
– Latocupes bella Tan et Ren, 2006; early
Cretaceous, Barremian-Aptian, Yixian Forma-
tion; China, Liaoning, Beipiao;
– Latocupes fortis Tan et Ren, 2006; early
Cretaceous, Barremian-Aptian, Yixian Forma-
tion; China, Liaoning, Beipiao;
– Latocupes angustilabialis (Tan, Huang et
Ren, 2007), comb.n. (Mesocupes); Middle Ju-
rassic, Jiulongshan Formation; Inner Mongolia;
Chifeng City, Daohugou.
– Latocupes collaris (Tan, Huang et Ren,
2007), comb.n. (Mesocupes); Middle Jurassic,
Jiulongshan Formation; Inner Mongolia;
Chifeng City, Daohugou.
– Latocupes latilabialis (Tan, Huang et Ren,
2007), comb.n. (Mesocupes); Middle Jurassic,
Jiulongshan Formation; Inner Mongolia;
Chifeng City, Daohugou.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
135
Figs 95–100. Fossil species of the subgenus Anaglyphites, photographed under Leica MZ 9.0 with camera
DFC290: 95–96 — print of holotype of Mesocupes (Anaglyphites) capitatus (Ponomarenko, 1966), comb.n.
(“PIN 1989/2993”): 95 — dry print, 96 — print covered by alcohol; Lower Cretaceous, Middle Neocomian;
Russia, Transbaikalia, Buryatya, Baissa; length of body 5.8 mm; 97–98 — counterprint of holotype of M.
(A.) capitatus comb.n. (“PIN 1989/1869”: 97 — dry counterprint, 98 — counterprint covered by alcohol);
99 — dry print of holotype of M. (A.) mongolicus (Ponomarenko, 1997), comb.n. (“PIN 3559/1608”); Lower
Cretaceous, ? Lower Aptian; Mongolia, Bon Tsagan; length of body 6.3 mm; 100 — dry print of paratype
of M. (A.) mongolicus comb.n. (“PIN 3790/277”), length of body 6.3 mm.
Рис. 95–100. Ископаемые виды подрода Anaglyphites, сфотографированные на микроскопе Leica MZ
9.1 с камерой DFC290: 95–96 — отпечаток голотипа Mesocupes (Anaglyphites) capitatus (Ponomarenko,
1966), comb.n. (“PIN 1989/2993”): 95 — сухой отпечаток, 96 — отпечаток, покрытый спиртом;
нижний мел, средний неоком; Россия, Забайкалье, Бурятия, Байса; длина теля 5,8 мм; 97–98 —
противоотпечаток голотипа M. (A.) capitatus comb.n. (“PIN 1989/1869”): 97 — сухой противоотпеча-
ток, 98 — противоотпечаток, покрытый спиртом; 99 — сухой отпечаток голотипа M. (A.) mongolicus
(Ponomarenko, 1997), comb.n. (“PIN 3559/1608”); нижний мел, ? нижний апт; Монголия, Бон Цаган;
длина тела 6,3 мм; 100 — сухой отпечаток паратипа M. (A.) mongolicus comb.n. (“PIN 3790/277”),
длина тела 6,3 мм.
136
A.G. Kirejtshuk et al.
Menatops gen.n.
(see above Diagnosis of this genus)
– Menatops orbiculatus (Kirejtshuk, Nel et
Colomb, 2010) [Cupes], comb.n.; Paleogene,
Paleocene; France, Puy-de-Dome, Menat.
– Menatops bartenevi sp.n.; Paleogene, Pa-
leocene; France, Puy-de-Dome, Menat.
Genus Mesocupes Martynov, 1926
(see above Diagnosis of this genus)
Subgenus Mesocupes Martynov, 1926, s.str.
– Mesocupes (Mesocupes) bidens Ponomar-
enko, 1964; late Jurassic, Oxfordian (or/and
Kimmeridgian); Kazakhstan Chimkent Region,
Kara-Tau Range, Karatau.
– Mesocupes (Mesocupes) primitivus Mar-
tynov, 1926; late Jurassic, Oxfordian (or/and
Kimmeridgian); Kazakhstan, Chimkent Region,
Kara-Tau Range, Karatau.
– Mesocupes (Mesocupes) minor Ponomar-
enko, 1968; late Jurassic, Oxfordian (or/and
Kimmeridgian); Kazakhstan, Chimkent Region,
Kara-Tau Range, Karatau.
– Mesocupes (Mesocupes) paulus (Pono-
marenko, 1964), comb.n. [Anaglyphites]; late
Jurassic, Oxfordian (or/and Kimmeridgian);
Kazakhstan, Chimkent Region, Kara-Tau Range,
Karatau.
– Mesocupes (Mesocupes) spinosus Pono-
marenko, 1964; late Jurassic, Oxfordian (or/and
Kimmeridgian); Kazakhstan, Chimkent Region,
Kara-Tau Range, Karatau.
– Mesocupes (Mesocupes) zherikhini (Sori-
ano et Delclós, 2006), comb.n. [Anaglyphites];
early Cretaceous, Barremian; Spain, Cuenca, 4
km NO from village Cierva, Las Hoyas.
Subgenus Anaglyphites Ponomarenko,
1964, stat.n.
The species of this subgenus have large cells
in the long longitudinal subparallel rows on the
elytra (sometimes rows are not clearly ex-
pressed), slightly expressed veins on the elytra
without fusion at apices (somewhat like those in
some Mesocupes), open and rather separated
antennal insertions, subflattened dorsal surface
of the head and subfiliform antennae with the
antennomere 3 very long while the species of
Mesocupes s.str. have the primary veins more
expressed and subfiliform antennae with com-
parably shorter antennomeres 2 and 3. Never-
theless Anaglyphites is very distinct due to the
comparatively shorter elytra which are usually
about 1.5 times as long as wide combined or not
infrequently slightly shorter while the elytra in
Mesocupes s.str. are generally longer. The type
specimen of Mesocupes (Anaglyphites) capita-
tus comb.n. shows a row of cells between elytral
Sc and lateral edge of the elytra while the elytra
of some specimens of M. (A.) admotus comb.n.
and M. (A.) clavatus comb.n. bear the sharply
and densely tuberculate or teethed (densely
crenellate at adge) stripes along the lateral edge
(similar to those in Priacma and Paracupes
s.str.). The pronotum of most specimens of this
subgenus demonstrate the finely crenelate sides
(Figs 95–102). Besides, the elytra of some spe-
cies of this subgenus have the rather steep ely-
tral sides with the sharp inflection [Mesocupes
(Anaglyphites) capitatus comb.n.] while the
paratype of M. (A.) mongolicus comb.n. has the
longitudinal rows of cells on the elytra which
give an impression of fusion of the traces of A1
and CuA (Fig. 100).
– Mesocupes (Anaglyphites) admotus (Pono-
marenko, 1964), comb.n. [Anaglyphites]; late
Jurassic, Oxfordian (or/and Kimmeridgian);
Kazakhstan Chimkent Region, Kara-Tau Range,
Karatau.
– Mesocupes (Anaglyphites) capitatus
(Ponomarenko, 1966), comb.n. [Anaglyphites];
early Cretaceous, Middle Neocomian; Russia,
Transbaikalia, Buryatya, Baissa.
– Mesocupes (Anaglyphites) clavatus (Pono-
marenko, 1964), comb.n. [Anaglyphites]; late
Jurassic, Oxfordian (or/and Kimmeridgian);
Kazakhstan, Chimkent Region, Kara-Tau Range,
Karatau.
– Mesocupes (Anaglyphites) minimus Pono-
marenko, 1997), comb.n. [Priacmopsis]; Early
Cretaceous, ? early Aptian, Bontsagan series;
Mongolia, Bayan-Khongor Ajmag, Bon Tsagan.
– Mesocupes (Anaglyphites) mongolicus
(Ponomarenko, 1997), comb.n. [Anaglyphites];
early Cretaceous, ? early Aptian, Bontsagan
series; Mongolia, Bayan-Khongor Ajmag, Bon
Tsagan.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
137
Figs 101–104. Fossil species of the subgenera Anaglyphites and Mesocupes s.str. (genus Mesocupes),
photographed under Leica MZ 9.0 with camera DFC290: 101–102 — holotype of Mesocupes (Anaglyphites)
clavatus (Ponomarenko, 1964), comb.n. (“PIN 2066/2275”): 101 — dry print; 102 — print covered by
alcohol; Upper Jurassic, Oxfordian (or/and Kimmeridgian); Kazakhstan, Karatau; length of body 6.0 mm;
103 — holotype of Mesocupes (Mesocupes) minor (Ponomarenko, 1968), comb.n. (“PIN 2239/812”);
Upper Jurassic, Oxfordian (or/and Kimmeridgian); Kazakhstan, Karatau; dry print, length of body 4.4 mm;
104 — additional specimen M. (M.) minor comb.n. (“PIN 2239/810”); Upper Jurassic, Oxfordian (or/and
Kimmeridgian); Kazakhstan, Karatau; dry print, length of body 4.0 mm.
Рис. 101–104. Ископаемые виды подродов Anaglyphites and Mesocupes s.str. (genus Mesocupes),
сфотографированные на микроскопе Leica MZ 9.0 с камерой DFC290: 101–102 — голотип Mesocupes
(Anaglyphites) clavatus (Ponomarenko, 1964), comb.n. (“PIN 2066/2275”): 101 — сухой отпечаток; 102 —
отпечаток, покрытый спиртом; верхняя юра, оксфордский (или/и кимериджский) яруса; Казахстан,
Каратау; длина тела 6,0 мм; 103 — голотип Mesocupes (Mesocupes) minor (Ponomarenko, 1968),
comb.n. (“PIN 2239/812”); верхняя юра, оксфордский (или/и кимериджский) яруса; Казахстан,
Каратау; сухой отпечаток, длина тела 4,4 мм; 104 — дополнительный экземпляр M. (M.) minor
comb.n. (“PIN 2239/810”); верхняя юра, оксфордский (или/и кимериджский) яруса; Казахстан,
Каратау; сухой отпечаток, длина тела 4,0 мм.
138
A.G. Kirejtshuk et al.
Figs 105–108. Fossil species of the subgenus Mesocupes s.str. (genus Mesocupes) and genus Cupidium,
photographed under Leica MZ 9.0 with camera DFC290: 105–106 — holotype of Mesocupes (Mesocupes)
paulus (Ponomarenko, 1964), comb.n. (“PIN 1789/91”): 105 — dry print; 106 — print covered by alcohol;
Upper Jurassic, Oxfordian (or/and Kimmeridgian); Kazakhstan, Karatau; length of body 3.8 mm; 107–108 —
holotype of Cupidium abavum Ponomarenko, 1968 (“PIN 2384/398”): 107 — dry print; 108 — counterprint
covered by alcohol; Upper Jurassic, Oxfordian (or/and Kimmeridgian); Kazakhstan, Karatau; length of body
9.6 mm.
Рис. 105–108. Ископаепмые виды подрода Mesocupes s.str. (рода Mesocupes) и рода Cupidium,
сфотографированные на микроскопе Leica MZ 9.0 с камерой DFC290: 105–106 — голотип Mesocupes
(Mesocupes) paulus (Ponomarenko, 1964), comb.n. (“PIN 1789/91”): 105 — сухой отпечаток; 106 —
отпечаток, покрытый спиртом; верхняя юра, оксфордский (или/и кимериджский) яруса; Казахстан,
Каратау; длина тела 3,8 мм; 107–108 — голотип Cupidium abavum Ponomarenko, 1968 (“PIN 2384/
398”): 107 — сухой отпечаток; 108 — противоотпечаток, покрытый спиртом; верхняя юра, оксфор-
дский (или/и кимериджский) яруса; Казахстан, Каратау; длина тела 9,6 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
139
Cainomerga subgen.n.
(see above Diagnosis of this subgenus)
– Mesocupes (Cainomerga) brevicornis
sp.n.; Paleogene, Paleocene; France, Puy-de-
Dome, Menat.
– Mesocupes (Cainomerga) fraternus sp.n.;
Paleogene, Paleocene; France, Puy-de-Dome,
Menat.
– Mesocupes (Mesocupes) immaculatus (Pi-
ton, 1940), comb.n. [Zonabris]; Paleogene,
Paleocene; France, Puy-de-Dome, Menat.
– Mesocupes (Cainomerga) palaeocenicus
sp.n.; Paleogene, Paleocene; France, Puy-de-
Dome, Menat.
– Mesocupes (Cainomerga) ponti sp.n.;
Paleogene, Paleocene; France, Puy-de-Dome,
Menat.
Genus Miocupes Ponomarenko, 1973
This genus is quite distinct and character-
ized by the medium body-size (body length 12.5
mm), the subparallel primary veins on the elytra
without fusion at apices, nine long longitudinal
rows of cells on the elytra, the very large and
wide head with the comparatively small eyes,
the head sides with distance from the temples to
the eyes distinctly greater than that from the
eyes to the anterior edge of the frons (i.e. the
very long temples markedly longer than frons
before the eyes), the antennal insertions rather
widely separated and the insertions somewhat
covered by the tubercles (convexities), the rath-
er widely separated and subparallel gular su-
tures; and the transverse pronotum.
– Miocupes rihai Ponomarenko, 1973; Neo-
gene, Middle Miocene; Czech Republic, Pochlo-
vice near Kynsperk.
Genus Priacmopsis Ponomarenko, 1966
The large body (ca. 30 mm) is the most
characteristic feature of this genus (after type
species — Priacmopsis adumbrata). Other char-
acters mentioned in the original description are
scarcely applicable for a reliable diagnostics,
except, probably, the small subquadrate cells in
the long longitudinal rows on the elytra of its
type species and the pronotum which is about
twice as wide as long and somewhat narrowing
anteriad, and also the short antennomere 2
(pedicel). The re-examination of the holotype
recovered also a distinct triangular depression
on the frons which is because of its large size
somewhat similar to the Y-shaped depression in
some other Mesozoic cupedines. On the print of
the holotype there are almost traced 10 long
longitudinal rows of cells on the elytra, but on
the counterprint a row of strongly transverse
cells between Sc and lateral (true anterior) edge
of the elytra is clearly visible. Besides, in con-
trast to other cupedines, the metacoxae of Pri-
acmopsis adumbrata are very short, with the
nearly subparallel anterior and posterior edges
and very slightly extended in their mesal part
forming a gentle curve along their posterior
edge. This feature was erroneously drawn in the
original description, as well as the shape of the
metaventrite (Figs 120–121).
The second species formerly included in this
taxon (Priacmopsis minimus Ponomarenko,
1997) is much smaller (body 9 mm), with the
pronotum much wider and apparently with larg-
er cells. Thus this second species can be scarce-
ly joined together with P. adumbrata in the
same genus and it was temporaly placed into the
subgenus Anaglyphites stat.n. of the genus Me-
socupes (see above).
– Priacmopsis adumbrata Ponomarenko,
1966; early Cretaceous, Middle Neocomian;
Russia, Transbaikalia, Baissa.
? Genus Pulchicupes Ren, 1995
See above Latocupes Tan et Ren, 2006.
Taxopsis gen.n.
(see above Diagnosis of this genus)
– Taxopsis motschulskyi (Kirejtshuk, 2005),
comb.n. [Cupes]; Paleogene, late Eocene; Bal-
tic See coast, Russia, near Kaliningrad (former-
ly Königsberg), Jantarny, Baltic amber.
8.2. List of fossil cupedine genera incertae
sedis (with indefinite position)
Genus Asimma Ponomarenko, 1966
This genus is represented by one species that
seems to be very similar to those of the genus
140
A.G. Kirejtshuk et al.
Figs 109–116. Fossil species of the genus Kirghizocupes, photographed under Leica MZ 9.0 with camera
DFC290: 109–111 — holotype of K. proporeius (Ponomarenko, 1969), comb.n. (“PIN 2240/3”): 109 — dry
print; 110 — print covered by alcohol, 111 — dry counterprint; ? Middle–Upper Triassic, Ladinian-Karnian;
Kyrgyzstan, Osh Region, Djailaucho (Dzhayloucho), Madygen; length of body 5.8 mm; 112 — specimen
of Kirghizocupes sp. (“PIN 3064/6424”, print covered by alcohol); ? Middle–Upper Triassic, Ladinian-
Karnian; Kyrgyzstan, Osh Region, Djailaucho (Dzhayloucho), Madygen; length of beetle remains 4.5 mm;
113–115 — holotype of Kirghizocupes indistinctus (Ponomarenko, 1969), comb.n. (“PIN 2240/72”): 113 —
dry print; 114 — print covered by alcohol, 115 — dry counterprint; ? Middle–Upper Triassic, Ladinian-
Karnian; Kyrgyzstan, Osh Region, Djailaucho (Dzhayloucho), Madygen; length of body 7.5 mm; 116 —
specimen of Kirghizocupes sp. (“PIN 3064/6424”, dry counterprint), length of beetle remains 5.9 mm.
Рис. 109–116. Ископаемые виды рода Kirghizocupes, сфотографированные на микроскопе Leica MZ
9.0 с камерой DFC290: 109–111 — голотип K. proporeius (Ponomarenko, 1969), comb.n. (“PIN 2240/
3”): 109 — сухой отпечаток; 110 — отпечаток, покрытый спиртом, 111 — сухой противоотпечаток;
? средний-верхний триас, Ладинский-Карнийский яруса; Киргизстан, Ошская обл., Джайляучо,
Мадыген; длина тела 5,8 мм; 112 — экземпляр Kirghizocupes sp. (“PIN 3064/6424”, отпечаток,
Taxonomy of the reticulate beetles of the subfamily Cupedinae
141
Figs 117–119. Fossil species of the genus Latocupes, photographed under Leica MZ 12.5 with camera Nikon
Digital Camera DXM1200C: 117 — holotype of L.s bellus Tan et Ren, 2006 (“CNU-C-LB2005013”, dry
print); Lower Сretaceous, Yixian Formation; China, Liaoning, Beipiao; length of body 12.7 mm; 118 —
holotype of . fortis Tan et Ren, 2006 (“CNU-C-LB2005012”, dry print); Lower Сretaceous, Yixian
Formation; China, Liaoning, Beipiao; length of body 27.5 mm; 119 — ? holotype of L.s collaris (Tan, Huang
et Ren, 2006), comb.n. (? “NIGPAS 142147”, found in collection of CNU without number); Middle Jurassic,
Jiulongshan Formation; Inner Mongolia; Chifeng City, Daohugou; dry print; length of body 13.3 mm.
Рис. 117–119. Ископаемые виды рода Latocupes, сфотографированные на микроскопе Leica MZ 12.5
с камерой Nikon Digital Camera DXM1200C: 117 — голотип L. bellus Tan et Ren, 2006 (“CNU-C-
LB2005013”, сухой отпечаток); нижний мел, формация исянь; Китай, Ляонинь, Бейпяо; длина тела 12,7
мм; 118 — голотип L. fortis Tan et Ren, 2006 (“CNU-C-LB2005012”, сухой отпечаток); нижний мел,
формация исянь; Китай, Ляонинь, Бейпяо; длина тела 27,5 мм; 119 — ? голотип L. collaris (Tan, Huang et
Ren, 2006), comb.n. (? “NIGPAS 142147”, найден в коллекции CNU без номера); средняя юра, формация
Джулиньшан; Внутренняя Монголия; Чифен, Даохугоу; сухой отпечаток; длина тела 13,3 мм.
покрытый спиртом); ? средний-верхний триас, Ладинский-Карнийский яруса; Киргизстан, Ошская
обл., Джайляучо, Мадыген; длина остатков жука 4,5 мм; 113–115 — голотип Kirghizocupes indistinctus
(Ponomarenko, 1969), comb.n. (“PIN 2240/72”): 113 — сухой отпечаток; 114 — отпечаток, покрытый
спиртом, 115 — сухой противоотпечаток; ? средний-верхний триас, Ладинский-Карнийский яруса;
Киргизстан, Ошская обл., Джайляучо, Мадыген; длина тела 7,5 мм; 116 — экземпляр Kirghizocupes
sp. (“PIN 3064/6424”, сухой противоотпечаток), длина остатков жука 5,9 мм.
142
A.G. Kirejtshuk et al.
Figs 120–123. Fossil species of the subfamily Cupedinae, photographed under Leica MZ 9.0 with camera
DFC290: 120 — holotype of Priacmopsis adumbrata Ponomarenko, 1966 (“PIN 1989/3000”); Lower
Cretaceous, Middle Neocomian; Russia, Transbaikalia, Baissa; dry print, length of specimen 30 mm; 121 —
additional specimen of P. adumbrata (“PIN 1989/3032”), print under alcohol; 122 — holotype of Taxopsis
motschulskyi Kirejtshuk, 2005, comb.n. (“PIN 363/130”); Paleogene, Upper Eocene; Russia, Jantarny,
Baltic amber; body, dorsal; 123 — idem, elytral apices, length of body 16.1 mm.
Рис. 120–123. Ископаемые виды подсем. Cupedinae, сфотографированные на микроскопе Leica MZ
9.1 с камерой DFC290: 120 — голотип Priacmopsis adumbrata Ponomarenko, 1966 (“PIN 1989/3000”);
нижний мел, средний неоком; Россия, Забайкалье, Бурятия, Байса; сухой отпечаток; длина экземп-
ляра 30 мм; 121 — дополнительный экземпляр P. adumbrata (“PIN 1989/3032”), отпечаток, покрытый
спиртом; 122 — голотип Taxopsis motschulskyi Kirejtshuk, 2005, comb.n. (“PIN 363/130”); палеоген,
верхний эоцен; Россия, Янтарный, балтийский янтарь; тело, сверху; 123 — то же, вершины
надкрылий, длина тела 16,1 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
143
Kirghizocupes by many characters including
the structure of pterothorax, the elytra probably
with 10 long longitudinal rows of cells and
without fusion of veins at apices. However, it is
differing mostly in the somewhat more slender
body and the pronotal sides less widely explan-
ate. Unfortunately the head of the remains of the
type species of this genus is missing and struc-
ture of underside of prothoracic segment is not
clear. To clarify the position of this genus it is
necessary to redescribe the holotype of this type
species.
– Asimma rara Ponomarenko, 1966; ? Mid-
dle–late Triassic, Ladinian-Karnian; Kyr-
gyzstan, Osh Region, Djailaucho (Dzhaylou-
cho), Madygen.
Genus Ensicupes Hong, 1976
This genus seems to be a member of Cupe-
didae, although it was described as Taldycupe-
didae, because the print with the holotype of its
type species (Ensicupes guyanensis: elytron
13.2 mm long) shows the characteristic features
in the polygonal cells arranged into not clear
long longitudinal rows and weakly raised pri-
mary veins with the fusion of A1 and CuA
characteristic of Cupedinae. However it has not
any more definite character for definion on its
position in this subfamily.
– Ensicupes guyanensis Hong, 1976; early
Cretaceous, Aptian, Guyang Formation; China,
Guyang County, Houmachi;
– Ensicupes obtusus Zhang, 1997; early
Cretaceous, Dalazi Formation; China, Jilin Prov-
ince, Zhixin Basin.
Genus Pterocupes Ponomarenko, 1966
This genus is represented by two species.
The type species of it (P. antennatus) seems to
be very similar to those in Asimma and Kir-
ghizocupes with the elytra having steeply slop-
ing and sharply inflected sides, however its
head, in contrast to that of the latter, is markedly
shorter. Another species of the genus (P. lepto-
cerus) is represented by the remains of a speci-
men that apparently was much more robust than
those of the type species, with the strongly
transverse head and the wider elytra with not
sharp (obliterated) inflection of elytral sides
outwards from the seventh row of cells. There-
fore, if the holotype of P. leptocerus was not
deformed by fossilization, it should be consid-
ered as member of another genus. To clarify the
position of this genus it is necessary to re-
examine the holotypes of both species.
– Pterocupes antennatus Ponomarenko,
1966; ? Middle–late Triassic, Ladinian-Kar-
nian; Kyrgyzstan, Osh Region, Djailaucho
(Dzhayloucho), Madygen.
– (?) Pterocupes leptocerus Ponomarenko,
1966; ? Middle–late Triassic, Ladinian-Kar-
nian; Kyrgyzstan, Osh Region, Djailaucho
(Dzhayloucho), Madygen.
8.3. List of fossil cupedine species without
generic attribution
– ‘Priacma’ sanzii Soriano et Delclós 2006;
early Cretaceous, Barremian; Spain, Cuenca, 4
km NE from village Cierva, Las Hoyas.
The remains of the holotype show the very
different characters from those of both Cupes
and Priacma (Kirejtshuk et al., 2010), particu-
larly concerning the shape of the pronotum with
the truncate anterior edge and the rounded sides,
the sculpture of elytra, and the very short anten-
nae. Therefore, this species should be regarded
as an “incertae sedis” among the Cupedinae till
a further re-examination of the type specimen.
– ‘?Anaglyphites’ pluricavus Soriano et
Delclós, 2006; early Cretaceous, early Barremi-
an; Spain, Ilerda, La Cabrua, Montsec Range,
Montsec.
In the description of this species there is no
clear diagnostic character usable for generic
attribution. However, at least the narrowly sep-
arated and open posteriorly procoxal cavities
can indicate that this species could be regarded
as a member of Cupedinae.
– ‘?Platycupes’ sogdianus Ponomarenko,
1966; ? Middle–late Triassic, Ladinian-Kar-
nian; Kyrgyzstan, Osh Region, Djailaucho
(Dzhayloucho), Madygen.
This species is rather different from the type
species of the genus Platycupes due to its rather
slender body. It is also characterized by the
144
A.G. Kirejtshuk et al.
comparatively narrowly separated procoxae and
10 long longitudinal rows of large subquadran-
gular cells (without expressed primary veins) on
elytra. To clarify the position of this species, it
is necessary to re-examine the holotype of it.
8.4. List of fossil cupedid genera incertae
sedis (uncertain or corrected subfamily), not
Cupedinae or doubtful members of Archo-
stemata
Genus Anthocoleus Hong, 1983
The type species of this genus is represented
by a narrow elytron (12 mm long). It has more or
less clear 10 long longitudinal rows of irregular
large cells and not raised primary veins. This
taxon was proposed as a member of Taldycupe-
didae, however it could belong to a cupedid
group, although its subfamily attribution is im-
possible to define with certainty.
– Anthocoleus hebeiensis Hong, 1983 (Tal-
dycupedidae); Middle Jurassic, Callovian/Ox-
fordian, Jiulongshan Formation; China, Lu-
oping County, Zhouyingzi village.
Genus Argentinocupes Martins-Neto et
Gallego in Martins-Neto et al., 2006
This genus was proposed for the elytra with
outlines of 2.5–9.3 mm long initially put in the
family Cupedidae (Martins-Neto et al., 2006a,
b). They are characterized by nine long longitu-
dinal rows of very large cells and weakly ex-
pressed primary veins. Ponomarenko regards
that these elytra could belong to Permocupe-
didae (Kirejtshuk, Ponomarenko, 2016), but to
clarify the position of this genus it would be
necessary to re-examine the accessible materials.
– Argentinocupes pulcher Martins-Neto et
Gallego in Martins-Neto et al., 2006; late Trias-
sic, Los Rastros Formation; Argentina, La Rioja
Province, Río Gualo;
– Argentinocupes abdalai Martins-Neto et
Gallego in Martins-Neto et al., 2006; late Trias-
sic, Los Rastros Formation; Argentina, La Rioja
Province, Río Gualo.
Genus Bothynophora Heer, 1865
This genus is a member of Cupedidae (un-
less Taldycupedidae), as the print with the holo-
type (elytron) of its type species shows the
characteristic features in cells arranged into the
long longitudinal rows. It is also characterized
by the unraised primary veins, without clear
fusion of them at apex. However the subfamilial
position of this genus is uncertain at the mo-
ment.
– Bothynophora elegans Heer, 1865 (Oede-
meridae); Middle Eocene; Germany, near Halle,
Geiseltal.
Genus Celocoleus Hong, 1983
The type species of this genus is represented
by one narrow elytron (11 mm long) and has
more than 10 unclear long longitudinal rows of
the irregular large cells and not raised primary
veins. It was proposed as a member of Taldycu-
pedidae, however it could belong to a cupedid
group, although the subfamily attribution of it is
scarcely possible to define.
– Celocoleus densus Hong, 1983 (Taldycu-
pedidae); Middle/late Jurassic, Callovian/Ox-
fordian, Jiulongshan Formation; China, Luop-
ing County, Zhouyingzi village.
Genus Chengdecupes Hong, 1983
This genus was proposed for the almost
complete specimen (22.0 mm long) initially put
in the family Cupedidae. The type species
(Chengdecupes jurassicus) has the subtriangu-
lar metacoxae and the large cells on elytra
arranged in 9 or 10 long longitudinal rows on the
elytra which are very characteristic of many
cupedines. The rather short metaventrite, ap-
parently the (sub) contiguous procoxae (if they
were correctly drawn by the descriptor), the
steeply sloping elytral sides and the weakly
raised primary veins with the fusion of A1 and
CuA before apex make it possible to suppose
the attribution of this genus to the Ommatinae.
Probably three species described as members of
this genus belong to different generic groups. In
particular, Chengdecupes kezuoense is repre-
sented by the elytral apex with the clear large
cells and the well expressed primary veins which
have a pattern apparently characteristic of the
species of Notocupes (Ommatinae) rather than
that of other cupedids.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
145
– Chengdecupes jurassicus Hong, 1983;
Middle/late Jurassic, Callovian/Oxfordian Jiu-
longshan Formation; China, Hebei, Chengde,
Xiaofanzhangzi (? Tongchuan Formation; Chi-
na, Shanxi, Shiluo);
– Chengdecupes kezuoense Hong, 1987;
early Cretaceous, Shahai Formation; China,
Liaoning, Xiwan village, Meileyingzi, Kezuo;
– Chengdecupes shiluoense Hong, 1984;
Middle Triassic, Tongchuan Formation; China,
Shilou County, Mengjiachang.
Genus Clathrocupes Hong, 1980
The type species of this genus was initially
attributed to Taldycupedidae. However the char-
acters in the original description do not make it
possible to conclude whether it belongs to Cu-
pedidae or to any other Mesozoic archostem-
atan family. Although the large cells on the
elytra arranged in the long longitudinal rows
without clear primary veins and without fusion
at apices as well as with the comparatively wide
elytral “epipleura” have some similarity with
those in Triadocupedinae, six abdominal ven-
trites drawn in the original description do not
give a reason to include it in Cupedidae.
– Clathrocupes anthrilegnotos Hong, 1980;
Middle Triassic, Tongchuan Formation; China,
Shanxi Province, Schiluo, Mengjiachang.
Genus Cupedites Ponomarenko, 1985
This “formal” genus in the family Cupe-
didae was proposed for the small elytra (1.8–2.2
mm long) characterized by the well raised and
(sub) parallel primary veins reaching their later-
al edge without fusion and elongately oblong
cells between them. The subfamily attribution
of this species (the type species, ! by monotypy)
is questionable, however, the available charac-
ters are enough to compare this supraspecific
taxon with other cupedid genera and subgenera.
– Cupedites minutissimus Ponomarenko,
1985; early Jurassic, Makarovskaya Formation;
Siberia, Krasnoyarsky Kray, Krasnoyarsk (Taty-
shev Island).
Genus Doggeriopsis Handlirsch, 1906
This genus seems to be a member of Cupe-
didae or Taldycupedidae, as the print with the
holotype of its type species (comparatively wide
elytron with length 19 mm) shows the character-
istic features in the cells arranged into clear long
longitudinal rows [it was drawn by Buckland
(1837) as Buprestis sp.]. It seems to be also
characterized by the absence of difference in
development in the primary and secondary veins
and the absence of clear fusion of them at apex.
However this genus is scarcely can be linked
with any subfamily without re-examination of
the type specimen.
– Doggeriopsis stonefieldiana Handlirsch,
1906; Middle Jurassic, Dogger, Taunton Lime-
stone Formation; England, Oxfordshire, Stones-
field Slate.
Genus Eumecoleus Haupt, 1950
This genus was proposed for the elytra with
the outlines of 12 mm long placed in Oedemer-
idae and compared with the sculpture Cupe-
didae. Hörnschemeyer et al. (1995) considered
them as probable archostematan, although, ex-
cept two striae along the middle of the distal half
that somewhat resemble the primary veins A1
and Cu in the cupedines, these prints have not
any character which can be used for definition
of their family or suborder attribution, although
its most probable placement could be among
cupedids.
– Eumecoleus tenuis Haupt, 1950 (Oede-
meridae); Middle Eocene; Germany, near Halle,
Geiseltal.
Genus Euteticoleus Hong, 1983
(= Eutecticoleus Hong, 1983; Eutecticupes Hong, 1983)
The type species of this genus is represented
by one comparatively large and narrow elytron
(8.5 mm long) and has more or less clear 10 long
longitudinal rows of the irregular large cells and
not raised primary veins. It was initially pro-
posed as a member of Taldycupedidae, however
it could belong to a cupedid group, but a sub-
family attribution of it is scarcely possible to
define.
– Euteticoleus radiatus Hong, 1983 (Taldy-
cupedidae); Middle Jurassic, Callovian/Oxfor-
dian, Jiulongshan Formation; China, Chengde
City, Xiaofanzhangzi.
146
A.G. Kirejtshuk et al.
Genus Gansucupes Hong, 1982
The type species is represented by one nar-
row elytron 10.0 mm long and 2.3 mm wide as
a member of Cupedidae. The photograph of this
elytron shows four primary veins (but the orig-
inal drawing is with five veins) with the charac-
teristic cells arranged in 10 long longitudinal
rows, although A1 and CuA are not distinct and
the primary veins at elytral apex are becoming
unclear (without visible fusion). This elytron
seems to be somewhat similar to that of Cu-
pedites minutissimus and could belong to Cupe-
dinae or Ommatinae due to comparatively large
oval to polygonal cells.
– Gansucupes attenuatus Hong, 1982; early
Cretaceous, Barremian, Chijinqiao Formation;
China, Gansu Province, Yumen, Chijinqiao (Ji-
uquan).
Genus Hebeicoleus Hong, 1992
This genus can be a member of Cupedidae or
Taldycupedidae, as the print with the holotype
(elytron of 8.5 mm long) of its type species
shows the characteristic features in the cells
arranged into the long longitudinal rows. It is
also characterized by the unraised primary veins,
the absence of clear fusion at apex and particu-
larly by the strongly transverse cells on the
elytral disk (as those in Latocupes and Taxopsis
gen.n.). However the subfamily attribution of
this genus remains uncertain without re-exami-
nation of the type specimen.
– Hebeicoleus sertulatus Hong, 1992; Mid-
dle Jurassic, Jiulongshan Formation; China,
Hebei, Chengde, Donggou village.
Genus Hebeicupes Zhang, 1986
This genus seems to be a member of Cupe-
didae, as the the holotype of its type species
(complete robust body of 9.3 mm long, with the
rather wide pronotum and the short elytra) shows
the characteristic features in the oval cells ar-
ranged into not clear long longitudinal rows and
the absence of clearly raised veins. To prove its
family attribution a further re-examination of
the type specimen is required.
– Hebeicupes formidabilis Zhang, 1986;
Middle/late Jurassic, Callovian/Oxfordian, Jiu-
longshan Formation; China, Luoping County,
Zhouyingzi village.
Genus Helopides Roemer, 1876
The type species of this genus was described
after the study of one separate elytron as similar
to that in Helops Fabricius, 1775 (Tenebrio-
nidae Latreille, 1802) and showing the charac-
teristic cupedid venation and cellulation with
somewhat expressed primary veins, the fusion
of A1 and CuA and the characteristic prescutel-
lar concentration of cells isolated by A2. At this
moment it is impossible to make any conclusion
about the subfamily attribution of this genus.
– Helopides hildsiensis Roemer, 1876; late
Triassic; Germany, Niedersachsen (Kraelaeh
near Hildesheim.
Genus Ironicus Handlirsch, 1906
(= Harpalidium Westwood, 1854)
The name Harpalidium nothus was pro-
posed for a species represented by one elytron
(10 mm long) which seems to have sculpture
somewhat looking like that in Cupes due to the
long longitudinal rows of large “cells”, although
as it can be seen in the original drawing (West-
wood, 1854: pl. 17, Fig. 3) the interspaces
between the “cells” are obliterated (somewhat
like that in Chalepocarabus). Handlirsch (1906–
1908) proposed for this genus a new name
Ironicus, probably because “hat nichts mit Har-
palus gemein” [Handlirsch, 1906 (1906–1908):
559] and also because the generic name Harpa-
lidium was used for another very different spe-
cies in the same publication (Westwood, 1854:
Harpalidium anactus: 393 and pl. 14, Fig.20)
without designation of type species for the pro-
posed taxon. Later the junior homonym was
proposed (Harpalidium Kolbe, 1883), which,
in turn, is regarded as a junior synonym of
Siopelus Murray, 1859 (Carabidae). Indeed the
possible affiliation of this fossil genus to Cupe-
didae is also still unclear. This confusing situa-
tion could be solved after a further re-examina-
tion of the type specimen.
– Ironicus nothus (Westwood, 1854) [Har-
palidium]; early Cretaceous, Berriasian, Pur-
beck Limestone Formation; England, Dorset,
Durlston Bay near Swanedge.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
147
Genus Katapontisus Handlirsch, 1906
The holotype of the type species (fragment
of apical part of elytron of 2 mm long) was
mentioned and drew by Brodie (1845: pl. 6, Fig.
9) as Limnius (?) sp. Later Giebel (1856) pro-
posed for it the name Elmis brodiei, and, finally,
Handlirsch (1906–1908) erected for this spe-
cies a separate genus with an unclear family
attribution. The fragment of the elytron demon-
strates the large cells and some traces of veins,
including the fusion of A1 and CuA character-
istic of Cupedidae, however is scarcely possible
to define a subfamily attribution of this species.
– Katapontisus brodiei (Giebel, 1856) [El-
mis]; early Cretaceous, Berriasian, Lulworth
Formation; England, Wiltshire, Dinton, Vale of
Wardour.
Genus Kelidus Handlirsch, 1906
(=Buprestium Westwood, 1854)
The name Buprestium bolbus was proposed
for a species represented by one elytron (5 mm
long) which seems to have sculpture somewhat
looking like that in Cupes due to the long longi-
tudinal rows of large “cells”. As it can be seen in
the original drawing (Westwood, 1854: pl. 17,
Fig. 5) and in Handlirsch (1906–1908: pl. 45,
Fig. 46) the interspaces between the “cells” are
without clear primary veins, although the fusion
of A1 and CuA is traceable (at least in the
disposition of cells). Handlirsch (1906–1908)
proposed for this genus a new name Kelidus,
probably because of “ist nichts sicher als Bu-
prestide zu erkennen” [Handlirsch, 1906 (1906–
1908): 557] and also because the generic name
Buprestium was used for some different species
in the same publication (Westwood, 1854: Bu-
prestium dardanus, B. stygnus, B. taleas, B.
valgus and B. woodlei) without designation of
type species for this used taxon. Unfortunately
the subfamily attribution of this fossil species is
scarcely possible even after a re-examination of
the type specimen, although some probability of
its link to cupedids can be supposed.
– Kelidus bolbus (Westwood, 1854) [Bu-
prestium]; early Cretaceous, Berriasian, Lul-
worth Formation; England, Dorset, Swanage/
Purbeck.
Genus Lecanopteryx Haupt, 1950
This genus was proposed for the elytra with
outlines of 10 mm long having neither any trace
of venation and cells nor another character which
can be used for definition of their family attribu-
tion. Hörnschemeyer et al. (1995) considered
them as probable archostematan, although, ex-
cept general outline looking like somewhat sim-
ilar to that in cupedines, these prints have not
any character which can be used for definition
of their family attribution.
– Lecanopteryx miranda Haupt, 1950 (Cu-
pedidae); Middle Eocene; Germany, near Halle,
Geiseltal.
Genus Liassocupes Zeuner, 1962
This genus with two species is re-interpreted
in the composition of Ommatinae after re-exam-
ination of the type species of the genus (Liasso-
cupes parvus Zeuner, 1962) (Ponomarenko,
2006). Besides, Liassocupes gigantea Whalley,
1985 in the latter publication was transferred
into the genus Mimema Handlirsch, 1906
(Schizocoleidae) (Ponomarenko, 2006). How-
ever the position of two other species needs to
be clarified after a further re-examination their
type materials. Hubert et al. (2003) identified
one elytron from New Jersey (U.S.A.) as “Lias-
socupes parvus”, but it would be better to regard
it without subfamily and generic attribution (see
below Chapter 8.5).
– Liassocupes maculatus Whalley, 1985;
early Jurassic, Sinemurian; England, Dorset,
Stonebarow, Charmouth;
– Liassocupes parvus Zeuner, 1962; early
Jurassic, Sinemurian; England, Dorset, Stone-
barow, Charmouth.
Genus Longaevicupes Ren, 1995
This taxon is traditionally regarded as a
junior synonym of Tetraphalerus (Kirejtshuk,
Ponomarenko, 2016), although it was proposed
as a member of the Taldycupedidae Rohden-
dorf, 1861. However, the remains of the holo-
type of the type species of this genus are 8 mm
long and represented by the elytra with nine long
longitudinal rows of cells, the pterothorax and
the abdomen similar to those of the slender
148
A.G. Kirejtshuk et al.
cupedines rather than to slender ommatines. It
particularly concerns the outline of the elytra.
Its very important features are expressed in the
weak primary veins combining with the fusion
of A1 and CuA at elytral apex, which are more
characteristic of various cupedines, but not Tet-
raphalerus. The very wide metepisterna of this
species are rather unique among members of the
family Cupedidae at all.
– Longaevicupes macilentus Ren, 1995; early
Cretaceous, Aptian, Lushangfen Formation;
China, Western Beijing Province, Jingxi Basin,
Lushangfen village.
Genus Lupicupes Ren, 1995
According to the original description the
remains of this species (holotype of 16 mm long
and exposing the elytra, meso- and metatibiae,
and abdomen) demonstrate 12 (or not less than
11) long longitudinal rows of strongly trans-
verse cells on the elytra. The primary elytral
veins of this specimen are more or less distinct
and showing the fusion of A1 and M while CuA
is diappearing between these veins. This gener-
ic name was recently synonymized with Noto-
cupes without argumentation (Ponomarenko,
2006). However it is difficult to be sure whether
Lupicupes trachylenus Ren, 1995 belongs to
Cupedinae or Ommatinae, although its elytra
with the strongly transverse cells seem to be
similar to those in Latocupes, Mesothoris and
Pulchicupes (see above) rather than any other
cupedids.
– Lupicupes trachylenus Ren, 1995; early
Cretaceous, Aptian, Lushangfen Formation;
China, Western Beijing Province, Jingxi Basin,
Lushangfen village.
Genus Mesothoris Tillyard, 1916
This genus was proposed for the elytron
named Mesothoris clathrata Tillyard, 1916 (type
species, monotypy), which Tillyard regarded as
resembling “Thoris” Pascoe, 1867 (Ceramby-
cidae Latreille, 1802) and compared it also with
Omma (Tillyard, Dunstan, 1916). This elytron
demonstrates 10 long longitudinal rows of the
subquadrangular cells and unclear primary veins
characteristic of Cupedidae. Later Dunstan
(1923) added to this genus three additional
species (M. grandis Dunstan, 1923, M. quadri-
partite Dunstan, 1923 and M. tenuiclathrata
Dunstan, 1923). The remains of the first species
include the part of elytron with 10 long longitu-
dinal rows of the strongly transverse cells and
unclear primary veins, pterothorax and abdo-
men. This species (M. grandis) could be regard-
ed as a member of the same genus. However,
Mesothoris tenuiclathrata and M. quadripar-
tite have much greater number of long longitu-
dinal rows and could be transfered to the genus
Mesolobites Carpenter, 1985 (= Lobites Dun-
stan, 1923, non Mojsisovics, 1873) of the fam-
ily Schizocoleidae Rohdendorf, 1961 (Kirej-
tshuk, Ponomarenko, 2016). Another species
was described after a separate elytron as a
members of the genus Elaterium Westwood,
1854 (E. punctomarginum Dunstan, 1923) with
11 long longitudinal rows of cells, including one
row placed along explanate stripe of the elytral
side. Riek (1968) described a further species in
this genus, Mesothoris westraliensis Riek, 1968,
and put it in the family Tenebrionidae. This
species together with M. clathrata and M. gran-
dis instead of their differences in structure of
elytra are here preliminarily regarded as cupe-
dids with unclear position (Cupedinae or Om-
matinae). These attributions are confirmed by
the pictures of the type specimens published by
Jell (2004).
– Mesothoris clathrata Tillyard, 1916; late
Triassic, Carnian; Australia, Queensland, Den-
mark Hill.
– Mesothoris grandis Dunstan, 1923; late
Triassic, Carnian; Australia, Queensland, Den-
mark Hill.
– Mesothoris punctomarginum (Dunstan,
1923), comb.n. [Elaterium]; late Triassic, Car-
nian; Australia, Queensland, Denmark Hill.
– Mesothoris westraliensis Riek, 1968; late
Triassic; Australia, West Australia, Hill River.
Genus Mesotricupes Hong, 1982
Two species of this genus are represented by
the medium-sized elytra about 5 mm long and 2
mm wide described as members of Cupedidae.
The photograph of the elytron of the type spe-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
149
cies (Mesotricupes lineatus) shows three appar-
ently convex narrow stripes looking like prima-
ry veins not fused but converging at the apex and
with probable sculpture between them appar-
ently consisting of small diffuse cells. The char-
acter of the “venation” of these elytra seems to
be somewhat similar to that in some members of
Tricoleidae Ponomarenko, 1969, although their
sculpture between “primary veins” is complete-
ly different, and, therefore, it would be reason-
able to temporarily consider them without sub-
order attribution.
– Mesotricupes lineatus Hong, 1982; early
Cretaceous, Aptian, Chijinpu Formation; China,
Gansu Province, Yumen, Chenjiwan, Changma;
– Mesotricupes reticulatus Hong, 1982; early
Cretaceous, Aptian, Xiagou Formation; China,
Gansu Province, Yumen, Chenjiwan, Chang-
ma.
Genus Metacupes Gardiner, 1961
The type species of this genus is represented
by six fragments (five pieces of elytra and one
piece of metaventrite) having no clear diagnos-
tic character to identify their family and genus.
The large punctures or cells on elytra arranged
in the long longitudinal rows could make it
possible to suppose a link of this species with
any group of Archostemata, however the these
punctures or cells have circular outlines in the
drawing giving rise to doubt that they could be
really archostematan cells.
– Metacupes harrisi Gardiner, 1961; late
Triassic – early Jurassic, Rhaetian; England,
near Bridgend (Cnap Twt quarry, St. Bride’s
Island).
Genus Nannocurculionites Handlirsch,
1906
The type species of this genus was described
in the composition of the genus Curculionites
Heer, 1847, nec Kolbe, 1888 (Curculionidae
Latreille, 1802). It is represented by the part of
one elytron (1.7 mm long) showing characteris-
tic cellulation and venation with the expressed
primary veins with the fusion of A1 and CuA.
This elytron could belong to a member of Cupe-
didae (Cupedinae or Ommatinae).
– Nannocurculionites carlsoni (Heer, 1878)
[Curculionites]; late Triassic; Sweden, Scania,
Rhaetian Bjuf.
Genus Parabuprestites Handlirsch, 1906
The type species of this genus was described
in the composition of the genus Buprestites
(Buprestidae Leach, 1815). It is represented by
the part of one elytron (3.0 mm long) showing
characteristic cellulation and venation with the
expressed primary veins and the fusion of A1
and CuA which are somewhat similar to that in
Nannocurculionites carlsoni (see above). This
elytron could belong to a member of Cupedidae
(Cupedinae or Ommatinae).
– Parabuprestites rugulosus (Heer, 1878)
[Buprestites]; late Triassic; Sweden, Scania,
Rhaetian Bjuf.
Genus Paracurculionites Handlirsch, 1906
The type species of this genus was initially
described in the composition of the genus Cur-
culionites (Curculionidae). It is represented by
the part of one elytron (3.0 mm long) showing
the characteristic cellulation and venation with
somewhat expressed primary veins and the fu-
sion of A1 and CuA. This elytron could belong
to a member of Cupedidae (Cupedinae or Om-
matinae).
– Paracurculionites parvulus (Heer, 1878)
[Curculionites]; late Triassic; Sweden, Scania,
North of Sofiero.
Genus Paradoggeria Handlirsch, 1906
This genus is a member of Cupedidae or
Taldycupedidae, as the holotype (moderately
narrow elytron of 16 mm long) of its type
species shows the characteristic features in the
cells arranged into the clear long longitudinal
rows [it was drawn by Buckland (1837) as
Buprestis sp.]. It seems to be also characterized
by the absence of difference in development in
the primary and secondary veins and the absence
of clear fusion of them at apex. However this
genus is scarcely can be linked with any subfam-
ily without re-examination of the type specimen.
– Paradoggeria acuminata Handlirsch,
1906; Middle Jurassic, Dogger, Taunton Lime-
stone Formation; England, Oxfordshire, Stones-
field Slate.
150
A.G. Kirejtshuk et al.
Genus Petalocupes Hong, 1982
The type species of this genus described as
a member of Cupedidae is represented by a
small and narrow elytron about 4 mm long and
1 mm wide. The photographs of this elytron are
not clear and the drawings show four elongate
convex narrow stripes looking somewhat like
primary veins and not fused but convergent at
apex; and the sculpture between them consists
of small and irregular (frequently transverse)
diffuse cells. To clarify its family and suborder
attribution a further re-examination of the type
specimen is necessary.
– Petalocupes arcus Hong, 1982; early Cre-
taceous, Aptian, Xiagou Formation; China,
Gansu Province, Yumen, Xiagou City, Qing-
quan.
Genus Pseudosilphites Zeuner, 1930
The type species of this genus is represented
by the elytra (8.2 mm long) with strongly raised
primary veins and characteristic fusion of A1
and CuA before apices, which suggests an attri-
bution to the Cupedinae or Ommatinae (in the
original description, it was compared with Sil-
pha Linneus, 1758, Silphidae Latreille, 1807).
– Pseudosilphites triassicus Zeuner, 1930
(Silphidae); Middle Triassic; Germany, Nor-
drhein-Westfallen, “Strasse Vlotto-Bonneberg”.
– Pseudosilphites natalensis Zeuner, 1960
(Silphidae); Middle Triassic; Molteno beds,
South Africa.
Genus Stigmenamma Handlirsch, 1906
The type species of this genus is represented
by the elytron about 9.5 mm long described as a
member of Harpalus by Giebel (1956) and
figured before by Brodie (1945), who also men-
tioned that this elytron has “pale elytra with dark
brown markings”. The elytron bears 9–10 long
longitudinal subparallel rows of large cells and
no clear vein. It very probably belongs to Cupe-
didae. However, it is impossible to clarify sub-
family attribution without re-examination of the
type specimen.
– Stigmenamma heeri (Giebel, 1856) [Har-
palus]; late Triassic, Rhaetian (formerly “Early
Liassic”); England, Gloucestershire, Hasfiel.
Genus Synodus Hong, 1982 (Coleoptera),
not Scopoli, 1777 (Alepisauriformes)
The type species of this genus is represented
by the small and narrow elytron about 4 mm
long and 1.5 mm wide described as a member of
Cupedidae. The photographs of this elytron are
not clear and the drawings show nine elongate
convex stripes some of which look somewhat
like primary veins becoming, however, not clear
at apex and the sculpture between them appar-
ently consisting of irregular elongate cells. To
clarify family attribution a further re-examina-
tion of the type specimen is necessary, although
the belonging of it to Archostemata remains
very probable.
– Synodus changmaensis Hong, 1982; early
Cretaceous, Aptian, Chijinpu Formation; Chi-
na, Gansu Province, Yumen, Chenjiwan, Chang-
ma.
Genus Tentyridium Westwood, 1854
This genus was proposed for a species (Ten-
tyridium peleus) represented by one lateral part
and apex of elytron (7 mm long) which has the
rather large and dense cells arranged in the long
longitudinal rows and no trace of veins. Hand-
lirsch (1906–1908) proposed another species
name in this genus for another elytron (8.5 mm
long) illustrated by Westwood in the same pub-
lication which before remained without name
(Westwood, 1894: pl. 16, Fig. 17). The latter
elytron is rather different from that of Tentyrid-
ium peleus and probably belongs to a species
from another genus. However the subfamily
attribution of both is scarcely possible to define
even after a re-examination of these specimens.
– Tentyridium peleus Westwood, 1854; ear-
ly Cretaceous, Berriasian, Lulworth Formation;
England, Dorset, Swanage/Purbeck.
– Tentyridium dilatatum Handlirsch, 1906;
early Cretaceous, Berriasian, Lulworth Forma-
tion; England, Dorset, Swanage/Purbeck.
Genus Tetrocupes Hong, 1983
This genus was proposed for one elytron
12.0 mm long initially put in the family Cupe-
didae. It show four veins with 4–5 “rows of the
pore” (?cells) between them. Besides, in the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
151
description four long parallel veins are men-
tioned, although the picture and drawing of this
elytron show that probable Cu and M are run-
ning to be fused at apex. In order to clarify the
subfamily attribution and position of this genus
an additional re-examination of type specimens
is needed.
– Tetrocupes cavernasus Hong, 1983; Mid-
dle Jurassic, Jiulongshan Formation; China,
Hebei, Chengde, Xiaofanzhangzi.
8.5. List of fossil cupedid species without
generic attribution, incertae sedis (with in-
definite positions)
– “Liassocupes parvus” Hubert et al., 2003
(not Zeuner, 1962); early Jurassic, Towaco For-
mation; U.S.A., New Jersey, Roseland, Rose-
land Querry.
Hubert et al. (2003) identified an ellytron
collected in USA as “Liassocupes parvus” de-
scribed from U.K. and they also published a
picture of it which looks like characteristic of
cupedids with fused A1 and CuA and weakly
raised primary veins. Nevertheless it is impossi-
ble to be sure in such identification and this
specimens would be better to regard as a cupe-
did without subfamily and generic attribution.
– ‘Priacma’ corrupta Ponomarenko, 1986;
early Cretaceous, early Neocomian; West Mon-
golia Hovd Ajmag, Myangad (Myangat).
The remains of its holotype represent the
ventral sclerites of the pterothorax, and also the
sclerites of legs and the basal parts of the hind-
wings. They share the characters of some cupe-
dine genera, but, in contrast to Priacma serrata,
the metaventrite of this species is transverse and
its abdominal ventrite 1 is much longer than
each of the ventrites 2–4. The proposed original
attribution of this fossil specimen to any sub-
family also gives rise to doubt.
– ‘Priacma’ longicapitis Ponomarenko,
1997; early Cretaceous, ? early Aptian; Mongo-
lia, Bayan-Khongor Aymag, Gobi-Altay, Shar-
Tologoy.
These remains correspond to the head (ho-
lotype) and to a separated elytron (additional
specimen) somehow fitted to the holotype.
This taxon shares the characters of some cupe-
dine genera with the slender and more or less
subparallel-sided body showing the rather large
cells on the elytra. The holotype has the head
somewhat similar to that of Priacma serrata
but without any characteristic feature that
could provide an evidence for such attribu-
tion. The elytron seems to be similar to that of
P.
serrata and to species of some other cupe-
dine genera with the narrow elytra wearing a
weak primary venation. However, in contrast
to members of Priacma and the similar gen-
era, the mentioned elytron (additional speci-
men) has nine long longitudinal rows of cells.
Therefore, the proposed original attribution
of this fossil specimen to any subfamily also
gives rise to doubt.
– ‘Priacma’ oculata Ponomarenko, 1997;
early Cretaceous, ? Early Aptian; Mongolia,
Bayan-Khongor Aymag, Gobi-Altay, Shar-Tol-
ogoy.
The remains of its holotype have the char-
acters of Priacma as some other cupedine
genera with narrow body (more or less similar
head, part of thorax, elytron with slightly ex-
pressed primary veins and large cells), but
there is no character that could increase the
probability of its attribution to Priacma. Be-
sides, the proposed original attribution of this
fossil specimen to any subfamily is also prob-
lematical.
– ‘Priacma’ striata Ponomarenko, 2000;
early Cretaceous, Argun’ Formation; Trans-
baikalia, Chitiskaya Region, Elizavetinskaya
depression, Semen.
The holotype is represented by the metatho-
rax with elytra (with rather feeble primary veins)
and abdomen (15 mm long). It differs from
Priacma serrata by the long abdominal ventrite
1 and comparatively long hypopygidium. Pono-
marenko (2000) mentioned that this species
“has been allocated to the genus Priacma tenta-
tively based on the long metathorax and the last
abdominal sternite, and the elytra having char-
acteristic colour pattern with longitudinal
flecks”. However, the proposed original attri-
bution of this fossil specimen to a certain sub-
family also gives rise to doubt.
152
A.G. Kirejtshuk et al.
9. Position of the genus Priacma LeConte,
1874
Genus Priacma LeConte, 1874
Type species Cupes serratus LeConte, 1861,
recent, by monotypy
History of studies. For a long time this
genus was considered as perhaps the most inter-
esting to study because it was regarded as an
“archaic” member of the subfamily or family,
even though more usually it was Omma and/or
Tetraphalerus that were interpreted as more
“primitive”, “archaic” or “basal” for the “cupe-
dine clade” (Atkins, 1958a, b, 1963; Hörn-
schemeyer, 2009; etc.). However, Priacma ser-
rata had only been known from male adults until
Ross and Pothecary (1970) obtained a female
with eggs and even some freshly hatched 1st
instar larvae. Crowson (1962), albeit without a
proper diagnosis, had proposed the tribe Priac-
mini to solely encompass the genus Priacma,
but later a number of fossil genera were added to
Priacmini as well (Ponomarenko, 1969, 1997;
Tan, Ren, 2006, 2009; Tan et al., 2006; etc.).
Hörnschemeyer (2009) proposed different phy-
logenetic hypotheses in which Priacma, togeth-
er with the genus Paracupes, formed a separate
clade in every case. Hünefeld et al. (2011) agree
that Priacma and Paracupes represent a sepa-
rate clade: “They likely represent the first two
branches (or the first branch as a clade) within
the family” (Hünefeld et al., 2011: 157). How-
ever, Kirejtshuk et al. (2010a) showed that
taking into consideration also available fossils
Priacma is distinct from the other cupedines
only in structure of the male genitalia (but not in
its external characters) and proposed to aban-
don this tribal taxon as inapplicable. In contrast,
Bouchard et al. (2011) elevated its rank to the
subfamily Priacminae, yet without any explana-
tion or any note on its composition. Because
only the male genitalia are a unique character of
Priacma to distinguish the tribe or subfamily,
only two members of this suprageneric taxon,
Priacma and Gracilicupes, would be recogniz-
able while many fossil genera with still un-
known male genitalia are to be regarded as taxa
of unclear positions. The authors of the present
paper consider such taxonomic treatments as
irrational and only confusing the classification,
because most of the generic categories in the
family Cupedidae still remain unassigned to a
tribe or subfamily. Some paleocoleopterists have
attributed a few fossils to the genus Priacma,
but in all such cases this has been done without
due evidence (see above “Notes on the taxono-
my and systematics of the subfamily” and Chap-
ter 8). Some of these forms are placed here in
genera quite distinct from Priacma (see the
genus Apriacma gen.n.).
Diagnosis. Body elongate of medium size
(9.5–16.5 mm). Integument with moderately
coarse sculpture and punctation; large subqua-
drangular (transverse) to polygonal cells of elytra
arranged into 10 long longitudinal rows inter-
spaced by weak secondary and more raised
primary veins; A1 straight, almost reaching ely-
tral apex and fusing with CuA before apex.
Head usually subtriangular, with moderately
narrow neck (about 3/4 as wide as head at
temples), with a pair of long paramedial con-
vexities along eye edge to temples and covering
base of antennal insertions and with very slight-
ly expressed Y- or V-shaped depression, well
projecting temples; antennal insertions moder-
ately widely separated. Mandibles rather strong
and arcuately curved at outer edge before stout
bidentate apex. Mentum rather short and wide
(more than twice as wide as long). Gular sutures
widely separated, subgradually and slightly con-
vergent posteriorly. Antennae submoniliform
and moderately long; antennomere 1 (scape)
subequal in length with flagellomere 3 and 2.5–
3.0 times as long as antennomere 2 (pedicel) and
more or less thicker; all flagellomeres subconi-
cal and progressively becoming shorter distad,
although ultimamate and subultimate antenno-
meres subcylindrical and longer. Pronotum sub-
trapezoid, very slightly and rectilinearly wid-
ened anteriad, with subexplanate sides, project-
ing and subacute anterior angles, feebly tuber-
culate surface and weak median elevated stripe.
Part of prosternum before procoxae markedly
longer than procoxae. Elytra about three times
as long as wide combined, with broadly arcuate
lateral edges, subtruncate to subarcuate (or sub-
Taxonomy of the reticulate beetles of the subfamily Cupedinae
153
Figs 124–128. Priacma serrata LeConte, 1861 (ZIN “Montana, Hathead Co, Glacier Nat. Park, West
Glacier, USA, 17 May 1993, Ivie”); recent; photographed under Leica MZ 12.0 with camera DFC290: 124 —
head and pronotum, dorsal; 125 — elytral apex, dorsal view under impinging light; 126 — idem, under
transmitted light; 127 — elytral apices, ventral; 128 — posterior part of body, lateral. Length of body 16.4
mm. Abbreviation: fef — fixing epipleural fold.
Рис. 124–128. Priacma serrata LeConte, 1861 (ZIN “Montana, Hathead Co, Glacier Nat. Park, West
Glacier, USA, 17 May 1993, Ivie”); современный; сфотографированный на микроскопе Leica MZ 12.0
с камерой DFC290: 124 — голова и переднеспинка, сверху; 125 — вершина надкрылья, сверху при
падающем свете; 126 — то же, при проходящем свете; 127 — вершины надкрылий, снизу; 128 —
задняя часть тела, сбоку. Длина тела 16,4 мм. Сокращение: fef — замыкающая эпиплевральная
складка.
excised) apices and comparatively gently slop-
ing sides, lateral edge thickened and with stripe
bearing small, dense and sharp tubercles, epi-
pleura with fixing fold. Metaventrite about as
wide as long. Abdominal ventrite 1 somewhat
shorter to subequal to each of ventrites 2–4 and
hypopygidium markedly longest and rounded at
apex. Legs moderately thin, moderately long;
with protibia straight and tarsomeres 1–4 slight-
ly narrower to subequally wide lobed. Aedea-
gus of Priacma-type, with rather long, subcylin-
drical and strongly curved parameres, volsellar
plate, “digitus volsellaris”, “cuspis volsellaris”,
penis ended with a laterally compressed dolab-
riform apex (“uncus”).
On the structure of the male genitalia.
Priacma is highly distinct among all modern
beetles in an extraordinary structure of its male
genitalia. Edwards (1953a, b) published very
detailed studies of the genital capsule and aede-
agus of Priacma serrata, emphasizing their
considerable similarity to those in some Hy-
menoptera. Regrettably, the results of those
important investigations remained almost fully
154
A.G. Kirejtshuk et al.
ignored. Only Ponomarenko (1969) repeated
the striking peculiarities which Edwards had
observed in the male genitalia of Priacma ser-
rata while Iablokoff-Khnzorian (1980), noting
the Priacma-type aedeagus, questioned Ed-
wards’ interpretations of its structures. Lawrence
(1999) wrote that “the aedeagus of all modern
Archostemata (including Micromalthidae and
Crowsoniellidae) is of a distinctive type de-
scribed by Sharp & Muir (1912) as having
peculiarities which are “indications of an old
relationship with Insects of another Order (per-
haps something that preceded the existing Sial-
idae)”. Later Martynov (1931) developed a hy-
pothesis on the closer relation of Coleoptera and
Megaloptera which became generally acknowl-
edged (see below Discussion). However, after
the Edwards’ description of the male genitalia
of Priacma the facts on the structure of the
coleopteran male genitalia needed to be recon-
sidered. During some decades coleopterists did
not pay attention on this situation. By the way,
the structure of the coleopteran ovipositor can
be consideredas a possible “plesiomorphicform
of the Neuropteroidian ovipositor” (Mickoleit,
1973: 38). Hörnschemeyer (2009) regarded that
“clade XXVII (Priacma, Paracupes)”, includ-
ing “Paracupes” ascius (= Paracupoides
subgen.n. here), was more basal to the other
cupedines, treating them in a separate family,
Cupedidae, but he did not pay attention to con-
tradictions between the similarities in structure
of the male genitalia and elytra. Eventually, the
male genitalia in Priacma and Paracupes s.lato
are very different (see Figs 129–134 and Vulca-
no & Pereira, 1975: 51, Figs 27–35), but their
elytra are even more strikingly different from
those of Paracupoides subgen.n. (see above
Chapter 6). Besides this, Hörnschemeyer (2009)
found only three differences in structure of the
male genitalia of Priacma and Paracupes s.lato:
“dorsal plate” lying on the penis, “bifurcate
process” between sternite 9 and the aedeagus,
and the “position of the basal opening of the
aedoaegus”. However, even in the case of cor-
rectly coding all formalized characters of this
organ (characters 58–84) in the matrix, of all
110 characters listed by that author, the result-
ing cladogram would look essentially different.
Many neontologists like Hörnschemeyer (2009)
attach special importance to the “ventromargin-
al spines” and to the apical hooks of the
parameres as characteristic features of the Ar-
chostemata, even though these are sometimes
membranous, except for Priacma serrata, on
the one hand, while also membranous rudiments
of these structures are present in the parameres
of certain groups of Polyphaga, on the other
hand (e.g. very clear in the Recent members of
Cerophytidae Latreille, 1834 — see Fig. 138–
139; illustrations in Costa et al., 2003: 381, Fig.
8; and particularly Costa et al., 2014: Figs 11–
19; etc.; Dascillidae Guérin-Méneville, 1843 —
see illustrations in Jin et al., 2013: 620, Fig. 18
P–R; Pleocomidae LeConte, 1861 — see illus-
trations in Lawrence et al., 2011: 136, Fig. 55 F;
etc.). The species of Potergus Bonvouloir, 1871
(Throscidae Laporte, 1840) show articulated
appendages located at base of parameres or
places at joining parameres and phallobase (for
example, Cobos, 1961: 2, Fig. 5).
The term “ventro-marginal spines” (Neboiss,
1984) for the processes along the parameres in
archostematans, which is considered as a fea-
ture only characteristic of Archostemata as a
whole (e.g. Hörnschemeyer, 2009), needs spe-
cial comment as well, because the above inter-
pretation leads to a serious confusion. Hörn-
schemeyer (2009) denoted the processes locat-
ed along basal half of the parameres as “spines”,
but those located at the apices of the parameres
as “hooks”. Indeed, the aedeagus of Priacma
has two pairs of short curved sclerites (each a
“digitus volsellaris”, or a “cuspis volsellaris”)
which, as in hymenopterans, are separated from
both parameres (“lateral lobes”) and penis trunk
(“median lobe of aedeagus”) and which are
regarded as derivatives of gonocoxites 9 (Sn-
odgrass, 1941). In other groups, these sclerites
may have become fused to the base of the
phallobase (one of the pairs) and parameres
(another pair), this being observable in many
groups of archostematans and polyphagans
(mostly in elateriformians, particularly in the
superfamily Byrrhoidea, very clearly in Che-
lonariidae, but in other groups with various
degrees of expression: see members of Cneo-
glossa Guérin-Méneville, 1843 (Cneoglossidae
Taxonomy of the reticulate beetles of the subfamily Cupedinae
155
Champion, 1897) in Costa et al., 1999: 270,
Figs 45–48; numerous Eucnemidae Eschscholtz,
1829, with “secondary lateral lobes”, in Muona,
1993: 94, Figs 111–114 etc.; some Eulichas
Jacobson, 1913 (Eulichadidae Crowson, 1973),
in Hajek, 2007: 15, Figs 31–32). Articulation
between phallobase and parameres in the poly-
phagan aedeagus could be formed on base of a
ring of volsellar plates and parameral plates in
Priacma-type aedeagus.
The male genitalia of Priacma serrata are in
need of further detailed investigations and com-
parisons with different groups of holometab-
olans and other orders [the male genitalia exam-
ined by the authors (Figs 129–134) look some-
what different from those illustrated by Ed-
wards (1953a: 4–5, Figs 1–2; 1953b: 19, Fig.
1)]. Its genital capsule is also different from that
of other archostematans, and of the beetles in
general which are known to have this structure.
The ventral part of it in Priacma serrata (deriv-
ative of abdominal sternite 9) including a pair of
sclerites gradually widening posteriad and fused
together by their proximal ends (somewhat sim-
ilar to those in Paracupes: Vulcano, Pereira,
1975: 51, Fig. 35) while the dorsal part (deriva-
tive of abdominal tergite IX) with two sharp
paramedian pairs of narrow acuminate process-
es at the apex which are oriented mesally is
extremely distinct. Nevertheless, this segment
is more similar to that in the other beetles rather
than in hymenopterans and its peculiarities are
usually regarded as an apomorphy of the order
Coleoptera (Hünefeld et al., 2011). In contrast,
the aedeagus of Priacma serrata is more similar
to that in the hymenopterans (Boulange, 1924;
Snodgrass, 1941) rather than in the other beetles
and even in other orders of Neuropterida (As-
pöck, 2000), although Sharp and Muir (1912)
had found some “indications” of a relation of
male genitalia in Coleoptera (but not Priacma
ones) to those in Megaloptera (Sialidae). How-
ever the similarity of Priacma and hymenopter-
ans concerns the volsellar plate, “digitus volsel-
laris”, or “cuspis volsellaris”, which are charac-
teristic of many hymenopteran groups (Kimsey,
Wasbauer, 2013; Pesenko, 1983; Smith, Shino-
hara, 2011; etc.) which could, in turn, be homol-
ogous with basal appendages of both parameres
and penis trunk. As in hymenopterans each
“cuspis voscellaris” of the Priacma male aede-
agus has a strong muscle provides producing an
approaching of digiti and cuspes each other to,
when the muscles of both cuspes are contracted
(Edwards, 1953b). However, the aedeagus of
Priacma serrata also shares certain similarities
to those of elateriformians (see above) which, in
contrast to the aedeagus of archostematans,
show some conformations reminiscent of the
volsellar structures of hymenopterans. At the
same time, in addition to the notes of Edwards
(1953a) on the similarity of the male genitalia of
P. serrata to those of Hymenoptera, there are
some similarities to those of the other Holome-
tabola. The “uncus” of the penis trunk of P.
serrata is more similar to that in certain groups
of Lepidoptera (e.g. some Sphingidae Latreille,
1802: see in Simonsen & Kitching, 2014) rather
than to the probable homologue in some Co-
leoptera (as in the cupedine Rhipsideigma: see
Neboiss (1984: 466, Figs 59–66), as well as in
the elateriformians. S.Yu. Sinev considers that
the structure denoted as “uncus” could have had
independent origins in different lepidopteran
groups and frequently its shape is functionally
conditioned (personal communication). Some
similarities are also traced in the male genital
structures of Priacma and some dipterans (e.g.
Sinclair, 2000). However, probable homologi-
zations of structural details need a more careful
study. For instance, the parameres in the flies
seem to be in the same position as the cuspis
volsellaris of Priacma while the parameres in
Priacma seem to be in the same position as the
gonostyli of the flies (cf. Fig. 129 to Sinclair,
2000: fig. 2.1).
On some external structures. The genus
under consideration is also characterized by the
distinct subtruncate elytral apices, as well as by
the particular rows of sharp teeth along the
lateral elytral edge and the “fixing” epipleural
fold in Priacma serrata (Fig. 127). In most
cupedines, however, the elytral apices are con-
jointly subacute, forming a joint angle, but fos-
sil representatives of Anaglyphites stat.n., Cu-
pidium, Kirghizocupes, and Priacma, as well as
the modern species of Paracupes s.str. and
156
A.G. Kirejtshuk et al.
Figs 129–134. Male genitalia of recent Priacma serrata LeConte, 1861 (ZIN “Montana, Hathead Co,
Glacier Nat. Park, West Glacier, USA, 17 May 1993, Ivie”): 129 — aedeagus extracted from body without
preparation, dorsal, photographed under Olympus SCX9 with camera Olympus, length 1.7 mm; 130 —
idem, ventral; 131 — idem, lateral; 132 — aedeagus after maceration by KOH, dorsal, photographed under
Leica MZ 12.0 with camera DFC290; 133 — idem, ventral; 134 — male genital capsule without ventral
“bifurcate process” (“abdominal segment IX”), dorsal, length 2.0 mm. Abbreviations: ape — apex of penis
trunk (“uncus”), cus — “cuspis volsellaris”, dig — “digitus volsellaris”, par — parameres.
Рис. 129–134. Гениталии самца современного Priacma serrata LeConte, 1861 (ZIN “Montana, Hathead
Co, Glacier Nat. Park, West Glacier, USA, 17 May 1993, Ivie”): 129 — эдеагус, извлеченный из тела, без
обработки, сверху, сфотографированный на микроскопе Olympus SCX9 с камерой Olympus, длина
1,7 мм; 130 — то же, снизу; 131 — то же, сбоку; 132 — эдеагус после обработки в едкой щелочи
(KOH), сверху, сфотографированный на микроскопе Leica MZ 12.0 с камерой DFC290; 133 — то же,
снизу; 134 — генитальная капсула самца без нижних выростов “bifurcate process” (“брюшного
сегмента IX”), сверху, длина 2,0 мм. Сокращения: ape — вершина ствола пениса (“uncus”), cus —
“cuspis volsellaris”, dig — “digitus volsellaris”, par — парамеры.
Paracupoides subgen.n., demonstrate the ely-
tral apices separately rounded or truncate (Figs
84, 87, 125, 126). Observing this character in
fossils is associated with some potential obsta-
cles because of possible deformations of the
elytral apices specified by the positions of the
remains during, and by the circumstances of
fossilization. The important characters give the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
157
Figs 135–138. Male genitalia of recent representatives of Cupedidae and Cerophytidae, photographed under
Leica MZ 12.0 with camera DFC290: 135 — male genital capsule (“abdominal segment IX”) of specimen
of Cupes mucidus Chevrolat, 1829 (ZIN “Okr. Chernigovki, Prim. obl., Emel’yanov, 3 VII 914 “ (in
Cyrillics)), ventral; 136 — aedeagus of this specimen, ventral; 137 — male genital capsule (“abdominal
segment IX”) of specimen of Cerophytum japonicum Sasaji, 1999 (ZIN “Vinogradovka, Ussur. kr., 1.vi.929,
D’yakonov Philip” (in Cyrillics)), ventral; 138 — aedeagus of this specimen, ventral. Abbreviations: ah —
apical hook of parameres, ph — parameran hook, vms — ventro-marginal spine of base of parameres.
Рис. 135–138. Гениталии самцов современных представителей Cupedidae и Cerophytidae, сфотогра-
фированные на микроскопе Leica MZ 12.0 с камерой DFC290: 135 — генитальная капсула (“брюшной
сегмент IX”) экземпляра Cupes mucidus Chevrolat, 1829 (ZIN “Окр. Черниговки, Прим. обл., Емель-
янов, 3 VII 914 ”), снизу; 136 — эдеагус этого же экземпляра, снизу; 137 — генитальная капсула
(“брюшной сегмент IX”) экземпляра Cerophytum japonicum Sasaji, 1999 (ZIN “Виноградовка, Уссур.
кр., 1.vi.929, Дьяконов Филип”), снизу; 138 — эдеагус этого экземпляра, снизу. Сокращения: ah —
вершинный крючок парамеры, ph — крюк парамеры, vms — вентромаргинальный отросток основа-
ния парамеры.
elytral sides sometimes bearing the longitudinal
stripe with sharp and dense tubercles (teeth or
crenulation at adge) along epipleuran carina are
well expressed in Priacma, Paracupes s.str.
(Figs 125–128, 155, 157), while in Paracu-
poides subgen.n., there are only traces of such
stripes (Fig. 156) and other recent cupedines
such stripes are completely reduced (Figs 71,
72, 158). Unfortunately, these characters of
Priacma remain without an adequate compari-
son among the members of Cupedinae, because
it is scarcely traceable among fossils. Neverthe-
less, the holotype of Mesocupes (Anaglyphites)
admotus certainly has serrate edges of the elytra
showing teeth arranged in two parallel rows (as
in Priacma and Paracupes s.str.). At the same
time, this might not have been a random coinci-
dence between the type of sloping of the elytral
sides and the development of rows of teeth
along the lateral sides, even though it is difficult
to suggest any functional or evident morphoge-
netic link between these characters. The fixing
epipleural fold (Fig. 127) at the elytral apices is
unique among the modern cupedids, but like-
wise it seems to be almost impossible to trace
this feature in fossils. The first idea for the usage
of this structure that comes to mind is to promote
a reliable fixation of the abdomen. The cu-
158
A.G. Kirejtshuk et al.
pedines have a rather strong fixation of the
abdomen and elytra due to the subvertical lat-
erosternites and steeply sloped elytral sides
forming tightly congruent surfaces of these or-
gans. The fixing epipleural folds could serve as
an additional mechanism providing a more reli-
able fixation of abdomen and elytra. If so, their
presence in Priacma could provisionally be
interpreted as an adaptation to living in (semi)
aquatic or other very wet conditions, by analogy
with the structures along elytral sides consid-
ered by Crowson (1981), Kirejtshuk et al. (2013)
and others.
The long paramedian convexities on the
head of Priacma serrata (at least in a certain
position in respect of the light) show an impres-
sion of the “Y-depression” (Fig. 124) while
such a formation is not so well expressed in
other modern cupedines. Instead it is somewhat
similar to those in many Mesozoic cupedids. In
some cases it grants a good diagnostic character
(see above “Key to genera and subgenera”), but
its phylogenetic interpretation is frequently rather
restricted because of various reasons, including
its bad preservation in some fossils.
The comparatively short abdominal ven-
trite 1 of Priacma serrata is similar to that in
representatives of Kirghizocupes, although
these genera demonstrate enough of other clear
characters to be easily distinguished (see be-
low). At the same time, this similarity can be
combined with some similarity in the elytral
apices of both groups which are clearly not
conjointly subacute, but rather separately
rounded or subtruncate (see Figs 109–116 and
125). Nevertheless, Priacma and Kirghizo-
cupes seem to be hardly closely related groups
among cupedines because, in contrast to Pri-
acma serrata, members of Kirghizocupes have
the elytra with very weak veins which are not
fused at the elytral apices. Finding a specimen
of Kirghizocupes with exposed male genitalia
would be essential to clarify the relationship
between these groups.
Comparison. This genus has the Priacma-
type male aedeagus similar to that in Gracili-
cupes but differs from the latter in the longer
prosternum before the procoxae, the somewhat
longer the head with smaller eyes, the convexi-
ties covering antennal insertions, the much wid-
er and subquadrangular pronotum (wider than
head), the thicker antennae with the longer and
thicker scape and very short antennomere 2, the
shorter abdominal ventrite 1. Taking into con-
sideration the expression of raised elytral M and
isolation of lateral elytral stripes in the species
of Gracilicupes, the steepness of the slopes of
elytral sides in the latter is markedly greater than
that in Priacma. Also, in contrast to Priacma,
the elytral lateral edges of Gracilicupes are
simple and have no trace of serrate stripes.
Priacma, in addition to the distinct male
genitalia, differs from other cupedine genera
with 10 long longitudinal rows of cells on elytra
in the following characters:
– Anaglyphites stat.n. in the more slender
body, the clear convexities on the frons cover-
ing base of the antennal insertions, the prono-
tum with the rectilinear sides and strongly pro-
jecting anterior angles, the much narrower and
longer elytra with the clear primary veins and
fusion of A1 and CuA, the shorter abdomimal
ventrite 1;
– Cainomerga subgen.n. in the not so slen-
der body with more rectilinear elytral sides, the
more covered antennal insertions, the some-
what shorter and thicker legs and the much
shorter abdominal ventrite 1;
– Chalepocarabus in the more slender body
with elytral sides not arcuate, the shorter and
transverse pronotum with the rectilinear sides
and projecting anterior angles, the raised prima-
ry veins, the longer and thicker scape and very
short antennomere 2, the shorter abdominal
ventrite 1;
– Kirghizocupes in the shorter (not elon-
gate) head, the clear convexities on the frons
covering the base of antennal insertions, the
narrower prothoracic segment with the narrow-
ly subexplanate pronotal sides and the some-
what less quadrangular cells on elytra;
– Menatops gen.n. in the much longer head
with smaller eyes, the clear convexities on the
frons covering the base of antennal insertions,
the narrower and longer elytra, and the shorter
abdomimal ventrite 1;
Taxonomy of the reticulate beetles of the subfamily Cupedinae
159
– Mesocupes s.str. in the more marked pri-
mary veins, the clear convexities on the frons
covering the base of antennal insertions, the
thicker antennae with the longer and thicker
scape and very short antennomere 2, the shorter
abdominal ventrite 1;
– Paracupes sensu str. in the frons with two
pair of tubercles, the widely separated gular
sutures, the longer ventrite 1 and the not curved
protibiae;
– Paracupoides subgen.n. in the frons with
two pair of tubercles, the elytra with the less
expressed primary veins, the gently sloping
sides and the serrate lateral edge, the widely
separated gular sutures, the longer ventrite 1
and the not curved protibiae.
In addition to another type of aedeagus,
Priacma differs from modern cupedine genera
after the keys proposed by Atkins (1948) and
Crowson (1962) and from the fossil genera with
nine long longitudinal rows of cells on the elytra
in the number of these rows of cells and also in
the following characters:
– Apriacma gen.n. in the absence of Y-
shaped depression behind the antennal inser-
tions, the subtruncate elytral apices, the shorter
abdominal ventrite 1;
– Cretomerga gen.n. in the absence of Y-
shaped depression behind the antennal inser-
tions, the wider neck behind temples, the longer
prosternum and the narrowly separated proco-
xae;
– Cupes in the gently sloping elytral sides,
the subtruncate elytra apices, the absence of
tarsal gooves on prosternum;
– Cupopsis gen.n. in the head with the small
eyes and not concave along the anterior part of
their edges;
– Furcicupes in the subtruncate elytral api-
ces, the absence of Y-shaped depression behind
the antennal insertions and the wider neck be-
hind temples;
– Latocupes (? =Pulchicupes) in the more
slender body, the longer pronotum, the not trans-
verse cells on the elytral disk, and the absence of
Y-shaped depression on the frons;
– Miocupes in the more slender body, the
narrow and not transverse head with the not
strongly projecting temples, the longer prono-
tum, the shorter prosternum before the procox-
ae, and also in the not parallel primary veins and
more distinct cells on the disk of elytra;
– Taxopsis gen.n. in the smaller body-size,
the not transverse cells on the elytral disk, the
narrow prosternal process.
Finally, Priacma differs from the genera
with unclear number of long longitudinal rows
of cells, namely from Priacmopsis in the small-
er and much more slender body, and the shorter
prosternum before the procoxae, and from Cu-
pidium in the markedly more slender body, the
shorter antennomere 2, the longer pronotum
with the rectilinear sides and the projecting
anterior angles, the truncate (but not separately
rounded or conjointly subacute) elytral apices.
Note. If the male genitalia of Priacma are
similar to those of Gracilicupes, other external
characters of Priacma seem to share more sim-
ilarities with those of Furcicupes (see above).
On a phylogenetic interpretation. The tra-
ditional interpretation of a relationship of Pri-
acma and other cupedines as admitted by Ed-
wards (1953a, b), Atkins (1958a) and many
later authors cited in the below list of reference
is supported by new data analyzed in the present
paper. The numerous plesiomorphic characters
observed in structure of the male genitalia can
be seen as the trace of a very old split between
cupedines, and also ommatines and probably
polyphagan beetles. An independent structural
instauration of “ancestral” structure of the male
genitalia in both Priacma and Gracilicupes (as
changes in the sequence of a normal ontogenetic
development as described by Tichomirova
(1981) in the genital structures of some sta-
phylinids) can scarcely be admitted. Therefore,
the attribution of Priacma together with Graci-
licupes to a very old lineage (clade) of Co-
leoptera can be a more reasonable assumption
expressed in a joint structure of the male genita-
lia that may have been formed at the very early
stage of diversification of the order, when some
its subgroups maintained the comparatively ple-
siotypic male genitalia while others could have
already had a derived (apotypic) structure of
this organ which might have given rise at least to
160
A.G. Kirejtshuk et al.
Figs 139–141. Mesocupes (Cainomerga) fraternus sp.n. (holotype “MNHN.F.A52773”); Paleogene,
Paleocene; France, Menat: 139 — dry print, photographed under Olympus SCX9 with camera Olympus;
140 — body, ventral, with broken line outlined anterior edge of pronotum; 141 — integument of pterothorax.
Length of specimen without apical parts of abdomen and elytra 14.6 mm.
Рис. 139–141. Mesocupes (Cainomerga) fraternus sp.n. (голотип “MNHN.F.A52773”); палеоген, палео-
цен; Франция, Мена: 139 — сухой отпечаток, сфотографированный на микроскопе Olympus SCX9 с
камерой Olympus; 140 — тело, снизу, прерывистой линией очерчен передний край переднеспинки;
141 — поверхность птероторакса. Длина экземпляра без вершинных частей брюшка и надкрылий
14,6 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
161
Figs 142–144. Genus incertus sp.1 (MNHN “R 63906”) (subfamily Cupedinae); Paleogene, Paleocene;
France, Menat: 142 — dry print, photographed under Olympus SCX9 with camera Olympus; 143 — body,
dorsal, with broken line outlined anterior edge of prosternum; 144 — integument of anterior part of body.
Length of specimen 12.6 mm.
Рис. 142–144. Genus incertus sp.1 (MNHN “R 63906”) (подсем. Cupedinae); палеоген, палеоцен;
Франция, Мена: 142 — сухой отпечаток, сфотографированный на микроскопе Olympus SCX9 с
камерой Olympus; 143 — тело, сверху, прерывистой линией очерчен передний край переднегруди;
144 — поверность передней части тела. Длина экземпляра 12,6 мм.
162
A.G. Kirejtshuk et al.
Figs 145–150. Fossil species of the subfamily Cupedinae, photographed under Olympus SCX9 with camera
Olympus; Paleogene, Paleocene; France, Menat: 145 — apex of elytron of (?) Mesocupes (Cainomerga) sp.1
(“MNHN.F.A.52768”), dry print, length of apex of elytron 7.0 mm; 146 — Cupes aff. manifestus
(“MNHN.F.A.52771”), dry print, length of body 5.6 mm; 147 — Cupes aff. manifestus (“MNHN.F.A.52770”),
dry print, length of body 7.8 mm; 148 — Mesocupes (Cainomerga) sp., “MNHN.F.A.52769”, dry print,
length of elytron 12.6 mm; 149 — Genus incertus sp. 2, “MNHN.F.A.52794”, dry print, length of body 14.4
mm; 150 — idem, dry counterprint.
Рис. 145–150. Ископаемые виды подсем. Cupedinae, сфотографированные на микроскопе Olympus
SCX9 с камерой Olympus; палеоген, палеоцен; Франция, Мена: 145 — вершина надкрылья (?)
Taxonomy of the reticulate beetles of the subfamily Cupedinae
163
Figs 151–152. Genus incertus sp.3 (“MNHN.F.A.52772”) (subfamily Cupedinae), photographed under
Olympus SCX9 with camera Olympus; Paleogene, Paleocene; France, Menat: 151 — anterior part of dry
print, length of body 13.2 mm; 152 — anterior part of dry counterprint.
Рис. 151–152. Genus incertus sp.3 (“MNHN.F.A.52772”) (подсем. Cupedinae), сфотографированный на
микроскопе Olympus SCX9 с камерой Olympus; палеоген, палеоцен; Франция, Мена: 151 — передняя
часть сухого отпечатка, длина тела 13,2 мм; 152 — передняя часть сухого противоотпечатка.
Cupedinae, Ommatinae, Micromalthidae, Crow-
soniellidae, and Polyphaga, if not to all other
modern groups of this order. If so, 10 long
longitudinal rows of cells on the elytra of Priac-
ma and Gracilicupes could also represent an
initial state of the character to that with less
number of rows of cells. Both types of sloping of
the elytral sides apparently associated with the
serration of the elytral edges and the tubercula-
tion of the frons could have been homoplastic in
the phylogeny of cupedines. The recurrence of
the different conditions of these structures dur-
ing the historical development of the subfamily
Cupedinae might have been related to some
morphocyclic evolutionary processes (Emel’-
yanov, 2000) providing the parallel similar
changes in different structures and causing a
regular convergence (see “Discussion” below).
An example of such a recurrence in elytral
structure is shown here in the pair Paracupes
s.str. — Paracupoides subgen.n. Priacma to-
gether with Gracilicupes having the “archaic”
Mesocupes (Cainomerga) sp.1 (“MNHN.F.A.52768”), сухой отпечаток, длина вершины надкрылья 7,0
мм; 146 — Cupes aff. manifestus (“MNHN.F.A.52771”), сухой отпечаток, длина тела 5,6 мм; 147 —
Cupes aff. manifestus (“MNHN.F.A.52770”), сухой отпечаток, длина тела 7,8 мм; 148 — Mesocupes
(Cainomerga) sp., “MNHN.F.A.52769”, сухой отпечаток, длина надкрылья 12,6 мм; 149 — Genus
incertus sp.2, “MNHN.F.A.52794”, сухой отпечаток, длина тела 14,4 мм; 150 — то же, сухой
противоотпечаток.
164
A.G. Kirejtshuk et al.
Figs 153–158. Recent species of the subfamily Cupedinae, photographed under Olympus SCX9 with camera
Olympus: 153 — Paracupes (Paracupes) brasiliensis Kolbe, 1908 (MNHN, “Brasil, (?) Mur” “ex Museo
Mniczech” “coll. R. Oberthur”), elytron, lateral; length of body 16 mm; 154 — P. (Paracupoides) ascius
Neboiss, 1989, holotype (MNHN, “Equateur Or., de Baсos а Canelos, M. de Mathan, IX-X.1894”), elytron,
lateral; length of body 10.4 mm; 155 — P. (Paracupes) brasiliensis (same specimen — see above), elytra,
posterior; width 4.9 mm; 156 — P. (Paracupoides) ascius (same specimen — see above), elytra, posterior;
width 2.9 mm; 157 — Priacma serrata LeConte, 1861 (ZIN “Montana, Hathead Co, Glacier Nat. Park, West
Glacier, USA, 17 May 1993, Ivie”), elytra, posterior; width 4.5 mm; 158 — Adinolepis (Ascioplaga)
mimeta (Neboiss, 1984), comb.n., holotype, male (MNHN, “Mt Koghi, 15.iii. 1978 (Delobel)”), elytra,
posterior; width 2.6 mm.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
165
aedeagus type could thus be suggested to be the
sister to Permocupes unless to any tschekar-
docoleid group (see “Discussion” below).
10. Taxonomic rank of Adinolepis Neboiss,
1984 and Ascioplaga Neboiss, 1984
Neboiss (1984) initially proposed both Ad-
inolepis and Ascioplaga as two separate genera,
the former is endemic to Australia and the latter
to New Caledonia. The main difference of them
consists in the character of distribution of scales
on the elytra and some features in the structure
of male genitalia (Neboiss, 1984: 459: “differ-
ent elytra vestitute and male genitalia character-
istics”). Both groups under consideration have a
clear current isolation in space. In contrast to the
synonyms between Cupes, Tenomerga, and
Distocupes, the pair Adinolepis and Ascioplaga
seems to be somehow analogous to the pair
Paracupes s.str. and Paracupoides subgen.n.,
although the latter pair of two taxa is certainly
more distinguishable than the former pair. To
get somehow a balanced classification with com-
parable hiati between taxa of the same rank is
the reason to lower the rank of Ascioplaga
stat.n. and consider it in the genus Adinolepis
s.lato.
Discussion
Historical overview of studies on Ar-
chostemata
The Archostemata, Cupedidae and Micro-
malthidae have been placed in the Polyphaga by
most of the earliest authors. Ganglbauer (1903)
assigned the Cupedidae in the Adephaga and
Lameere (1903) proposed the Cupediformia as
the most “primitive” group of the Adephaga.
Kolbe (1908) coined the name Archostemata
and included that group in his division Symphy-
togastra of the suborder Heterophaga. Sharp &
Muir (1912) placed the Cupedidae in the Byr-
rhoidea (currently the infraorder Elateriformia)
on the basis that the male genital tube of Cupes
is of the “trilobe” type (herein, the Cupes-type),
the most “generalised” (“primitive”) existing in
the living Coleoptera. Finally, Forbes (1926)
erected the suborder Archostemata (including
Cupedidae and Micromalthidae) to separate
them from the Adephaga, based on the differ-
ence in the spiral rolling of the forewing apex.
Böving and Craighead (1930) recognised the
subordinal level, considering, however, that the
larval structures allowed for archostematans to
be regarded as representatives of a distinct an-
cient group. Since that time the Archostemata is
considered as the oldest group in the order, but
its relationships with the other suborders are
still under discussion. Crowson (1981) tried to
divide the Archostemata into three suborders,
adding the usage of the Protocoleoptera Tillyard,
1924 and Archecoleoptera Crowson, 1981.
However, the name Protocoleoptera had been
proposed only for the family Protocoleidae
(Tillyard, 1924), which is now considered a
component of the order Protelytroptera (Forbes,
1928; Carpenter, 1992; etc.). The Archeco-
leoptera was introduced by Crowson for the
Permian beetle groups except Tshekardoco-
leidae, but the usage of this name is scarcely
warranted because it is probably a paraphyletic
group and includes the late Permian taxa only
sharing some structural similarities which could
appear as convergence from an Early Permian
Рис. 153–158. Современные виды подсем. Cupedinae, сфотографированные на микроскопе Olympus
SCX9 с камерой Olympus: 153 — Paracupes (Paracupes) brasiliensis Kolbe, 1908 (MNHN, “Brasil, (?)
Mur” “ex Museo Mniczech” “coll. R. Oberthur”), надкрылье, сбоку; длина тела 16 мм; 154 — P.
(Paracupoides) ascius Neboiss, 1989, голотип (MNHN, “Equateur Or., de Baсos а Canelos, M. de Mathan,
IX-X.1894”), надкрылье, сбоку; длина тела 10,4 мм; 155 — P. (Paracupes) brasiliensis (тот же
экземпляр, см. выше), надкрылья, сзади; ширина 4,9 мм; 156 — P. (Paracupoides) ascius (тот же
экземпляр, см. выше), надкрылья, сзади; ширина 2,9 мм; 157 — Priacma serrata LeConte, 1861 (ZIN
“Montana, Hathead Co, Glacier Nat. Park, West Glacier, USA, 17 May 1993, Ivie”), надкрылья, сзади;
ширина 4,5 мм; 158 — Adinolepis (Ascioplaga) mimeta (Neboiss, 1984), comb.n., голотип, самец
(MNHN, “Mt Koghi, 15.iii. 1978 (Delobel)”), надкрылья, сзади; ширина 2,6 мм.
166
A.G. Kirejtshuk et al.
tshekardocoleid ancestor (Kirejtshuk et al.,
2013).
The most important apomorphies of the
order Coleoptera are manifested in the sclero-
tized forewings (elytra) and their venation (Sc
simple; RA convex and simple; RP simple,
more concave than RA; M less convex than R
and CuA, simple, and basally fused with, but
distally separated from CuA; M+CuA com-
pletely separated from radial stem; and CuA
nearly reaching forewing apex), thorax (mono-
lithic prothorax), legs (short trochanter and
non-projecting coxae) (Kirejtshuk, Nel, 2013;
Nel et al., 2013). All these characters are well
expressed in the suborder Archostemata. The
morphological characters shared by Coleoptera
in body structure are very clear in the late
Permian archostematans, viz. a dorsoventral
compression of a compact body; thorax (pro-
hypomeres); forewing (elytral) venation (CuA
and CuP separating at forewing base or very
close to it; CuP relatively short or reduced,
rather concave and simple; first anal vein con-
vex and more or less parallel to posterior forew-
ing edge; second anal vein of comparable con-
vexity, shorter, but also more or less parallel to
posterior edge; a third anal vein present, also
closely parallel to posterior forewing edge in
its distal part and fused with the second and
first anal veins to constitute an ‘adsutural line’);
hindwing venation (automatic folding device
and some autapomorphic features in venation
patterns with the radial system diminished and
anal one enlarged, see Kukalová-Peck &
Lawrence, 1993; Fedorenko, 2009); metacox-
ae extending on abdominal base and corre-
sponding fusion of derivates of two basal ab-
dominal sternites; modification of the abdomen
(laterosternites); and the coordinated structural
transformations of the organs of the pterothorax
and abdomen to constitute an isolated subelytral
space. Later, the subgroups of this suborder
maintained many above-mentioned features in
the condition close to the initial situation (Kire-
jtshuk et al., 2013). Characters of the Archo-
stemata probably include the restricted number
of antennomeres (usually 11–13), meso- and
metathoraces (pterothorax) fused together im-
movably; metathoracic flight motor; tegula lost;
subapical abdominal segments (beyond 8th one),
invaginated genitalia and abdominal segments 8
and 9 of both sexes. The tegular or overlapping
articulation of abdominal ventrites characteris-
tic of both Tshekardocoleidae (Moravocoleus
permianus Kukalová, 1969) and Cupedidae
(many groups) could also be provisionally rec-
ognized as an apotypic feature for Archostema-
ta. The order Coleoptera is usually treated in
composition of the superorder Coleopterida
which could include also Strepsiptera and Skle-
roptera (Kirejtshuk, Nel, 2013) and which is
traditionally linked to even joined with Neurop-
terida (frequently as the sister group) (Mar-
tynov, 1931; Hennig, 1981; Kristensen, 1999;
Rasnitzyn, 2002; Kukalová-Peck, Lawrence,
2004; Labandeira, 2011; Peters et al., 2014;
etc.). This concept has found some support from
current molecular studies (Wiegmann, Kim,
2009; McKenna, Farrell, 2010; etc.). However,
the early phyletic differentiations of holometab-
olans still remain rather unclear. Rohdendorf
considered that the supraorder “Coleopteroi-
dea” could be “descendants of the oldest car-
boniferous orthopteroids” (Rohdendorf, 1961:
294). In some phylogenetic reconstructions, the
lineage Coleopterida deviates next after the
lineage of Hymenopterida from general branch
after splitting Holometabola and Caloneuridea
(Rasnitsyn, 1980). Representatives of these su-
perorders appear as the first holometabolans in
the fossil record (Nel et al., 2013) and the data
on the structure of the male genitalia (see below)
are in accordance with this interpretation rather
than others, because the Priacma-type aedea-
gus maintain more common features with the
hymenopteran structure of this organ than those
of other holometabolan groups (see above Chap-
ter 9). It is worth noting that Mickoleit (1973)
concluded that the secondary telescopic ovipos-
itor, “vaginal palps in the Coleoptera may pos-
sibly represent a plesiomorphic form of the
Neuropteroidian ovipositor” (Mickoleit, 1973:
38). This paleontological and morphological
interpretation obtained also support in the re-
sults of molecular comparisons (up to sister
Taxonomy of the reticulate beetles of the subfamily Cupedinae
167
Figs 159–164. Fossil species of subgenenus Cainomerga subgen.n. (genus Mesocupes), photographed
under Olympus SCX9 with camera Olympus; Paleogene, Paleocene; France, Menat: 159 — M. (C.) sp.1
(“MNHN.F.A. 57518”), dry print, length of body 12.0 mm; 160 — M. (C.) sp. (“MNHN.F.A. 57519”), dry
print, length of elytron 14.6 mm; 161 — M. (C.) sp. (“MNHN.F.A. 57520”), dry print, length of posterior
part of elytron 4.5 mm; 162 — M. (C.) sp. (“MNHN.F.A. 57521”), dry print, length of exposed part of elytron
4.1 mm; 163 — M. (C.) sp. (“MNHN.F.A. 57523”), dry print, length of elytron 4.6 mm; 164 — (?) M. (C.)
sp.1 (“MNHN.F.A. 57522”), dry print, length of body, lateral 12.1 mm.
Рис. 159–164. Ископаемые виды подрода Cainomerga subgen.n. (рода Mesocupes), сфотографированные
на микроскопе Olympus SCX9 с камерой Olympus; палеоген, палеоцен; Франция, Мена: 159 — M. (C.)
sp.1 (“MNHN.F.A. 57518”), сухой отпечаток, длина тела 12,0 мм; 160 — M. (C.) sp. (“MNHN.F.A.
57519”), сухой отпечаток, длина надкрылья 14,6 мм; 161 — M. (C.) sp. (“MNHN.F.A. 57520”), сухой
отпечаток, длина задней части надкрылья 4,5 мм; 162 — M. (C.) sp. (“MNHN.F.A. 57521”), сухой
отпечаток, длина выступающей части надкрылья 4,1 мм; 163 — M. (C.) sp. (“MNHN.F.A. 57523”),
сухой отпечаток, длина надкрылья 4,6 мм; 164 — (?) M. (C.) sp.1 (“MNHN.F.A. 57522”), сухой
отпечаток, длина тела сбоку 12,1 мм.
168
A.G. Kirejtshuk et al.
Figs 165–168. Menatops bartenevi gen. et sp.n. (“MNHN F.A. 57524”, print and counterprint), photographed
under Olympus SCX9 with camera Olympus); Paleogene, Paleocene; France, Menat: 165 — dry print; 166 —
body, dorsal, with broken lines for outlines of below inner eye edges, sclerites of legs and abdomen; 167 —
head and antennae, dorsal; 168 — dry counterprint. Length of specimen 7.2 mm.
Рис. 165–168. Menatops bartenevi gen. et sp.n. (“MNHN F.A. 57524”, отпечаток и противоотпечаток),
сфотографированные на микроскопе Olympus SCX9 с камерой Olympus); палеоген, палеоцен;
Франция, Мена: 165 — сухой отпечаток; 166 — тело сверху, сверху, прерывистой линией очерчены
нижние внутренние края глаз, склериты ног и брюшка; 167 — голова и усики, сверху; 168 — сухой
противоотпечаток. Длина тела 7,2 мм.
group relations: Savard et al., 2006; Castro,
Dowton, 2007; Simon et al., 2012; etc.).
The studies of the Upper Carboniferous ho-
lometabolous larvae of the Moscovian age [Nel
et al., 2013: Metabolarva bella Kirejtshuk,
Prokin, Nel et Wappler, 2013 and Haug et al.,
2015: Srokalarva bertei J. Haug, Labandeira,
Santiago-Blay, C. Haug et Brown, 2015 (=Sro-
kalarva bertei Kukalova-Peck, 1997, nomen
nudum)] make it possible to suppose that larvae
looking like beetle ones came into the fossil
record much earlier than the first true beetles,
but at the age comparable with the findings of
the oldest representatives of the probable stem
Taxonomy of the reticulate beetles of the subfamily Cupedinae
169
lineages of hymenopterans and coleopterans
(Nel et al., 2013). These larvae have some
correspondence to many early Permian larvae
from Tshekarda, among which the larvae of
tshekardocoleids can be expected (Ponomaren-
ko, 1969; Prokin et al., 2015). The structure of
some of these larvae corresponds well to the
opinion that larvae of oldest beetles were asso-
ciated with tree trunks inhabiting inside sub-
strate, having their structures similar to those of
eruciform larvae of the Recent beetles (Pono-
marenko, 1969, 1988; Crowson, 1981; Kirejt-
shuk, et al., 2013; Nel et al., 2013; etc.). How-
ever, Haug et al. (2015) considered that the
Carboniferous larvae are members of the clade
Neuropterida and they were “herbivorous as
external foliage feeders” (Haug et al., 2015: 8).
This opinion was explained by these authors, on
the one hand, by the hypothetic reconstruction
of mouthparts of these larvae, which, however,
is rather uncertain and does not give a clear
reason to suppose the larval diet, and, on other
hand, by the findings of the Palaeozoic plants
damaged by some arthropods by chewing. Nev-
ertheless, the Recent members of the orders of
Neuropterida are mostly predaceous and only
the freshwater sisyrid larvae (Sisyridae Zalessky,
1932) feed on sponges (Porifera: Spongillidae
Gray, 1867) and probably on bryozoans (Phy-
lactolaemata). Labandeira (2011) regarded that
Srokalarva could belong to the Antliophora and
admited its feeding inside substrate. The inter-
pretation of surface inhabitance of Srokalarva
also contradicts to the concept by Lameere
(1899) on origin of the Holometabola as result
of certain transformations during individual
development connected with splitting of places
of larval inhabitance (concealed) and of place of
adult ihabitance (open). The important circum-
stance is the probable traces of metapneustic
respiratory system with an orifice at abdominal
apex observed in Metabolarva bella which could
testified its inhabitance in wet or even freshwa-
ter invironment (Nel et al., 2013). Among the
modern holometabolans this respiratory type
occurs in larvae of dipterans, coleopterans and
hymenopterans. The coleopterous larvae are
more probable pretenders to be linked with the
Carboniferous larvae by analogous or homolo-
gous structures, although all modern coleopter-
ous larvae with metapneustic respiratory system
are aquatic and very diverse in structure, but not
eruciform (Nel et al., 2013). Ponomarenko
(1988) following Lameere (1899) supposed that
the ancestors of holometabolans were eruci-
form and had habitats different from those of
their adults. The structure of Metabolarva bella
with probable metapneustic respiratory system
allowing living inside wet substrate is in accor-
dance with such interpretation. If the ancestors
of beetles were really associated with helophil-
ous forest, their larvae could have metapneustic
respiratory system allowing to be in wet inter-
stices under bark and in wood.
Phylogeny and historical develop-
ment of the suborder Archostemata
The oldest beetles known from the early
Permian are currently included in the suborder
Archostemata (Kirejtshuk et al., 2013). They
have the forewings with more or less expressed
remains of venation, which is frequently regard-
ed as key for tracing the phyletic diversifica-
tions because of great conservatism of this struc-
ture. Ponomarenko (1969) divided the suborder
Archostemata in the two phyletic cupedoid and
schizophoroid lineages (later called the in-
fraorders Cupedomorpha Ponomarenko, 2002,
and Schizophoromorpha Ponomarenko, 2002,
respectively). The former group is more similar
to members of the oldest known beetles (Tshe-
kardocoleidae) by structure of the elytra and
general appearance (Ademosynidae Ponomar-
enko, 1968, Cupedidae, Permocupedidae, Taldy-
cupedidae). This could have been somehow
associated with structural transformations lead-
ing to a syndrome of polyphagan morphology
(mostly on the basis of hindwing venation: see
Ponomarenko, 1973b). This archostematan
group, including both Tshekardocoleidae and
Cupedidae, maintains more plesiomorphic char-
acters than other beetles in the forewings, even
in Recent forms. The other group of archostem-
atan families (Schizophoromorpha: Asiocolei-
dae Rohdendorf, 1961, Catiniidae Ponomaren-
170
A.G. Kirejtshuk et al.
ko, 1968, Rhombocoleidae Rohdendorf, 1961;
Schizophoridae Rohdendorf, 1961, Tricoleidae)
could have been better associated with Adeph-
aga and Myxophaga and, therefore, according
to Ponomarenko (1973b), they could be linked
by phyletic relations characterized by the loss of
“traces of an archedictyon” (presence of cells as
transparent “windows” between veins arranged
in a less modified pattern) and multiplications
of long longitudinal rows of cells on the elytra at
the early stages of diversification (during the
Permian and Triassic). Later, the family Schizo-
coleidae Rohdendorf, 1961, has been added to
the latter lineage (infraorder) (Kirejtshuk, Pono-
marenko, 2016). Besides this, Ponomarenko
(1969) regarded Micromalthidae as showing an
unclear position within Archostemata. Howev-
er, the archedictyon in the strict sense seems to
have disappeared in the Holometabola before
the emergence of an ancestor of the cohorte
Coleopterida, since all other major holometab-
olan clades (Hymenopterida, Mecopterida, Neu-
ropterida) show no true archedictyon at all.
Kukalová-Peck & Lawrence (1993) proposed
the following phylogenetic hypothesis for the
beetles {Polyphaga + [Archostemata + (Myx-
ophaga + Adephaga)]}, which could be in some
agreement with the fossil record. Taking into
account the transformational rows in aedeagal
structure and the pattern of wing venation, it
could be provisionally suggested that the Cupe-
didae, the Micromalthidae, and the Crowsoniel-
lidae with five abdominal ventrites, probably
also the Jurodidae Ponomarenko, 1985 with six
abdominal ventrites, supposedly belong in the
cupedoid lineage of Archostemata together with
the probable ancestor of Polyphaga, while
Adephaga and Myxophaga could have origi-
nated within the schizophoroid lineage of this
suborder.
All lineages of the archostematans seem to
have emerged during the end of the Palaeozoic,
although many characteristic groups appeared
in the fossil record in the second half of the
Triassic (Ponomarenko, 1969, 1995, 2004;
Kirejtshuk, Ponomarenko, 2016). Therefore,
the main diversifications of the order can be
supposed to have happened during a compara-
tively short time (second half of the Permian to
the earliest Triassic). Various structural types
adapted to different ecological niches had al-
ready been represented among the late Permian
members of these lineages, in addition to the
oldest xylomycetophagans from Tshekar-
docoidae (Kirejtshuk, 1991; Ponomarenko,
1995; Kirejtshuk et al., 2013). Most findings of
late Permian beetles originated from Laurasia
and only few are known from South African
deposits (Pinto, 1987; Geertsema, van den Heev-
er, 1996; Ponomarenko, Mostovski, 2005). The
Mesozoic was the era of archostematan boom.
The greatest diversity of families and genera of
this suborder is observed during that era, al-
though some Palaeozoic groups (viz. Tshekar-
docoleidae, Asiocoleidae, Permocupedidae and
Rhombocoleidae) had disappeared earlier or
near the Palaeozoic to Mesozoic boundary
(Ponomarenko, 1969; Kirejtshuk, Ponomaren-
ko, 2016). Most Mesozoic findings of archo-
stematans also originated from Europeand Asia,
although their total range is much wider than
that of the Palaeozoic ones and includes many
forms from Australia (Dustan, 1923; Jell, 2004),
few taxa from Brazilian Crato from the Early
Cretaceous (Wolf-Schwenninger, Schawaller,
2007; Kirejtshuk, Nel, in preparation), Argen-
tinian Río Gualo from the early Triassic (Mar-
tins-Neto et al., 2006a, b) and few from the
Cretaceous deposits of North America (Pono-
marenko 1969b; Lubkin, 2003; Kirejtshuk,
Ponomarenko, 2016). The highest diversity
(number of archostematan genera and species)
is found in the Jurassic deposits, to progressive-
ly decline thereafter. To the end of the early
Cretaceous, the suborder Archostemata con-
sisted of families represented in the extant fauna
by a smaller number of genera and species.
The family Cupedidae could be regarded as
one of the groups “closely” related to the family
Tshekardocoleidae. Ponomarenko (1969) men-
tioned that the former mostly differed from the
latter by the reduction of CuP in the elytra, by
two rows of cells between all primary veins of
the elytra, and by open procoxal cavities. He
regarded Permocupedidae as a close relative of
Tshekardocoleidae. The elytra of both latter
Taxonomy of the reticulate beetles of the subfamily Cupedinae
171
families show remains of a CuP, less regular
(sometimes with a multiplied number and var-
ied sizes of the cells) and not always strictly
double rows of cells on the interspaces between
the primary veins, without fusion of A1 and
CuA at the apices; although the species of Per-
mocupes Martynov, 1933 (Permocupedidae)
have both A1 and CuA fused, while not all
Triadocupedinae, Cupedinae, and some Om-
matinae show this fusion very clearly (see above
“Notes on the taxonomy and systematics of
Cupedinae”). Besides this, Permocupes species
demonstrate almost regular double rows of cells
on each interspace at least between A3, A1,
CuA, M and R (as those in many Cupedidae).
According to Ponomarenko (1969), representa-
tives of the Tshekardocoleidae and Permocupe-
didae, presumably also Taldycupedidae, had
the procoxae closed posteriorly. This is the
reason to suggest an “ancestor” of Cupedidae
closer to the Permocupedidae rather than to any
other archostematans (Ponomarenko, 1969).
However, the structure of the prothorax of the
Permian beetles is generally poorly known, while
the presence of the posteriorly closed procoxae
among all the Palaeozoic groups remains more
or less hypothetic because of scarcity of com-
plete fossils studied. Rohdendorf (1961) pro-
posed to unite Cupedidae and Permocupedidae,
but Ponomarenko (1969) later restored the fam-
ily rank for both. However, the relations be-
tween these families still need revision after
additional studies on known and new materials
accessible in different collections. Ponomaren-
ko (1969) considered the genus Permocupes
with the elytral CuA and A1 fused at the apices
(as in most cupedines) as “progressive” among
permocupedid genera. However, if we admit
this character as synapomorphic for both Per-
mocupedidae and Cupedidae while separate ends
of these veins as having appeared secondarily, a
hypothesized sister-group relation of these fam-
ilies from a single tshekardocupedid lineage
may be recognized as more probable. Ponomar-
enko (1969) regarded that the first cupedids
should have had wide epipleura, also character-
istic of the Triadocupedinae, but not so frequent
in Cupedinae and Ommatinae. This feature is
usually associated with a stripe of more or less
large cells between Sc and the lateral (true
anterior) elytral edge and present in many tshek-
ardocoleids demonstrating explanate sides of
the elytra. One row of cells also occurs in many
triadocupedines and some members of both
cupedines and ommatines. Thus, these submar-
ginal cells could have been initial while their
lack in many cupedines and ommatines with
narrow sides (as in modern species) is a second-
ary homoplastic trait.
Unfortunately, the aedeagi of tshekar-
docoleids, permocupedids and taldycupedids
remain unknown. Moreover, many Permian
groups are only known from elytra which dem-
onstrate frequent parallel transformations and
cannot be used for reliable conclusions on their
phylogenetic relations. Therefore, it is impossi-
ble to determine whether these families had
male genitalia similar to those in Priacma or
Cupes. This point is important to define a diver-
gence in the lineage maintaining the probably
plesiomorphic Priacma-type aedeagus and the
lineage (clade) with the apomorphic Cupes-
type aedeagus which corresponds to the roots of
all other coleopteran groups. It is difficult to
admit that the appearance of the Priacma-type
aedeagus is a result of instauration of the ances-
tral condition of cupedines which could have
appeared from the clade uniting at least the
ommatines and cupedines, i.e. it is difficult to
consider the Priacma-type aedeagus as a de-
rived form the “plesiomorphic” (if it can be
admitted) cupedine type. On the other hand, as
the ontogenetic differentiations proceed the same
way in both sexes (Tichomirova, 1981), any
crucial change in this process’ sequence in one
sex should cause correlated changes in the op-
posite sex. The existence of a secondary tele-
scopic ovipositor in Tshekardocoleidae (Kirej-
tshuk et al., 2013) suggests that the male genita-
lia of this group were also already invaginated.
The ovipositor in Priacma remains still unde-
scribed.
The Cupedinae, including the forms with
both Priacma-type and Cupes-type aedeagi can
also be regarded as plesiomorphic in relation to
the male genitalia of Ommatinae apparently
172
A.G. Kirejtshuk et al.
Figs 169–174. Fossil species of subfamily Cupedinae, photographed under Olympus SCX9 with camera
Olympus; Paleogene, Paleocene; France, Menat: 169–172 — Cupes manifestus Kirejtshuk, Nel et Colomb,
2010: 169 — “MNHN.F.A. 57525”, head and antennae on dry print; 170 — same specimen, dry print, length
of body 8.9 mm; 171 — “MNHN.F.A. 57526”, dry print, length of body 7.8 mm; 172 — “MNHN.F.A.
57527”, dry print, length of body 8.9 mm; 173–174 —Genus incertus sp.4: 173 — “MNHN.F.A. 57529”,
dry print, length of elytron 5.9 mm; 174 — “MNHN.F.A. 57530”, dry print, length of elytron 6.2 mm.
Рис. 169–174. Ископаемые виды подсем. Cupedinae, сфотографированные на микроскопе Olympus
SCX9 с камерой Olympus; палеоген, палеоцен; Франция, Мена: 169–172 — Cupes manifestus Kirejtshuk,
Nel et Colomb, 2010: 169 — “MNHN.F.A. 57525”, голова и усики на сухом отпечатке; 170 — тот же
экземпляр, сухой отпечаток, длина тела 8,9 мм; 171 — “MNHN.F.A. 57526”, сухой отпечаток, длина
тела 7,8 мм; 172 — “MNHN.F.A. 57527”, сухой отпечаток, длина тела 8,9 мм; 173–174 — Genus
incertus sp.4: 173 — “MNHN.F.A. 57529”, сухой отпечаток, длина накрылья 5,9 мм; 174 — “MNHN.F.A.
57530”, сухой отпечаток, длина надкрылья 6,2 мм.
Taxonomy of the reticulate beetles of the subfamily Cupedinae
173
lacking traces of a volsellar plate, “digitus volsel-
laris”, “cuspis volsellaris” (see above Chapter
9). Edwards (1953a), following the Snodgrass’
terminology of the hymenopteran aedeagal struc-
tures (Snodgrass, 1941), considered these three
structures together with the parameran plate and
a broad sclerite at the base of the parameres (see
Fig. 131) as having participated in the formation
of a phallobase (basal piece), like it may have
arisen in the Hymenoptera, as we have shown
above. On the other hand, some researchers
regard the membranous appendages of the
parameres in Micromalthidae and Crowsoniel-
lidae as homologous with the derivative aedeagal
structures of the cupedids (Crowson, 1976;
Hörnschemeyer, 2009). Hünefeld et al. (2011)
regards that the “potential autapomorphy of
Archostemata is the median division of the
sclerotization of sternum IX, which is conspic-
uous in Tetraphalerus. In Cupedidae, two alter-
native conditions are found: an unpaired sternal
plate is present in most species of this family,
but paired ventro-lateral sclerites are found in
the small genera Priacma (1 sp.) and Paracupes
(2 spp.).” (Hünefeld et al., 2011). This hypoth-
esis is not evident even in the case of ignorance
that the embryonic base of abdominal sterna
have a paired nature (Tichomirova, 1991) and
that many genital structures of both sexes deri-
vated from abdominal sclerites have paired di-
visions (Tanner, 1927; Matsuda, 1976; Crow-
son, 1981; etc.) because of paired embryonic
base and genetics (Lewis et al., 2000). As some
polyphagans show remains of these structures
as well, we may suppose that the aedeagus of the
modern ommatines might have originated in-
dependently from that in polyphagans, some
groups of which also maintain such remains (see
above Chapter 9). Sharp and Muir (1912) pos-
tulated that the adephagan type of aedeagus
without a phallobase (op. cit. 1912: 622: “the
basal-piece is greatly reduced, or entirely mem-
branous”) has “no approximation” to that in the
cupedids and regarded it as rather specialized.
Crowson (1981) considered that the polyph-
agans may have obtained the phallobase through
fusion of the gonocoxites as an apomorphy of
this suborder. Although Lawrence (1999: 384)
supposed that “the absence of a phallobase may
be a synapomorphy of Archostemata + Adeph-
aga + Myxophaga”, some homologies in differ-
ent structures showing the formation of a phal-
lobase can be traced in the aedeagi of the Priac-
ma-type, the Cupes-type and some polyphagans.
This suggests that the volsellar complex and
phallobase could present in some archostem-
atans and polyphagans a symplesiomorphy while
it could have been reduced (as a volsellar plate,
“digitus volsellaris”, “cuspis volsellaris” or their
derivates) in the adephagans and myxophagans.
Fedorenko & Ponomarenko (2012) found a
schizophoroid hindwing from Isady (late Per-
mian, Tatarian, Poldarsa Formation), together
with a hindwind of a cupedomorph beetle. Tak-
ing this into account, it is possible that the
ancestors of both Adephaga and Myxophaga
could have been related to the Schizophorimor-
pha, while the ancestor of Polyphaga could have
been related to the lineage of the Cupedidae,
probably during the late Permian when beetles
had already become rather diverse (Volkov,
2012; Ponomarenko, 2015; etc.). The differenc-
es in their aedeagal structures could be a trace of
the sequence in the old divergence of this lin-
eage into a sublineage (leading to Recent Priac-
ma) and a different one which includes the most
well-known cupedines and a probable polyph-
agan “ancestor” with a Cupes-like aedeagus
(see above Chapter 9). Thus, the basal diver-
gences of the order can be defined as follows:
Tshekardocoleidea + {Schizophorimorpha +
[Cupedinae + (Ommatinae + other families of
Cupedomorpha)]}. The traditional phyletic link
of Schizophorimorpha to Adephaga and “Myx-
ophaga” and, accordingly, that of Cupedomor-
pha to Polyphaga can be admitted perhaps as the
most probable hypothesis (Ponomarenko,
1973b, 2002; Kirejtshuk, 1991; etc.).
On the phylogeny and historical de-
velopment of the subfamily Cupedinae
The hypothetic generalized pattern (ground-
plan, or archetype) of elytral venation seems to
be most stable among members of the subfamily
Cupedinae compared to any other archostem-
174
A.G. Kirejtshuk et al.
atan subfamily with more than ten described
members. It seems to include a weak or usually
reduced Sc (if raised), a long and weak R run-
ning along the lateral curve of the elytra and
ending at the very apex (or fusing to A1 just at
apex), a long M subparallel to R and fusing with
the latter at the apex, a long CuA subparallel to
the suture and fusing to A1 at the apex, a long A1
subparallel to the suture and reaching to the
latter’s edge, but somewhat deviating externally
at the scutellum, a short A2 subparallel to the
scutellum and slightly surpassing its apex along
the suture. The most usual place of change in the
elytral disc plane to sloping sides proceeds
along M. Sc and R are frequently very weak or
even scarcely traceable. The space between Sc
and the lateral (initially anterior) edge of the
elytra seems to be occupied by a row of cells
located on an explanate part of the elytral sides
(observed in Kirghizocupes, Priacmopsis, some
Anaglyphites, etc.) while the interspaces be-
tween the long primary veins (A3, A1, CuA, M,
R and Sc) bore a double row of cells. There are
some modes of transformation of this pattern.
One of these modes includes changes when A1
does not to reach the lateral edge of the elytra
and ends up in the distal part of the elytral suture
(in many Triadocupedinae and Ommatinae with
known venations, but only in the genus Prolixo-
cupes among Recent Cupedinae). Such differ-
ences in the ending of the primary veins can also
be observed among members of Permocupe-
didae, Taldycupedidae, and Ademosynidae. So
it can scarcely be regarded as a feature of great
phylogenetic meaning. Another mode consists
of an independent ending of all primary veins
along the apical part of the lateral (true anterior)
edge of the elytra, which is not infrequentamong
ommatines, but also occurs in the cupedine
genus Miocupes with parallel primary veins,
some cupedine groups with gradually approach-
ing primary veins but without fusion at the
apices (particularly members of the genera
Chalepocarabus, Kirghizocupes and Latocupes,
but also the subgenera Anaglyphites stat.n. and
Mesocupes s.str., etc.). Such a trend in elytral
venation is also observed in some other archo-
stematan families (Taldycupedidae, Permocu-
pedidae, Tricoleidae, Rhombocoeidae, etc.).
They likewise have the primary veins ending
independently along the lateral edge of the elytra.
This trend could supposedly be reversible in
some cases of an incomplete reduction of veins.
Another very frequent transformation is the
homoplastic reduction and disappearance of a
short A2 and a correspondingly short row of
cells at the scutellum. Finally, in some cases the
vein traces become completely obliterated (the
cupedine genera Mesocupes and Chalepocara-
bus) and make the elytra look similar to those in
some Permian groups. Such obliterated elytral
veins, coupled with the disappearance of clear
rows of cells, have happened in many archo-
stematan families and is to be recognized as a
sequence of homoplastic transformations which
without additional evidence, can hardly be re-
garded as a ground for phylogenetic conclu-
sions.
A rather extraordinary transformation of ely-
tral venation is found in the Cenozoic Caino-
cups aixensis gen. et sp.n. This species had four
long primary veins on a flat elytral disc, with an
unusual pattern of their fusion, and also 12 long
longitudinal rows of cells, instead 9 or 10 as
typical of Cupedinae. This species of an unclear
position between Cupedinae and Ommatinae
shows inflections to the steeply sloping lateral
sides along R (but not M) and the external four
rows of cells on a very wide sloping elytral side
(see above the description of this new genus and
species in Chapter 7). Nevertheless, the absence
of such a type of elytra in the previous fossil
record can suggest that this type is scarcely old
in origin.
The overlapping abdominal ventrites (char-
acteristic also of tshekardocoleids) can be rec-
ognized as plesiomophic for cupedines, be-
cause it occurs in both other cupedid subfami-
lies (Triadocupedinae and Ommatinae) as well
as the abutting type. The subvertical abdominal
laterosternites known both in Ommatinae and
Cupedinae with rather convex elytra seem to
have appeared during the early period of diver-
sifications of the early cupedids and could be a
synapomorphy for the family in general. The
fossil specimens with steep elytral sides should
Taxonomy of the reticulate beetles of the subfamily Cupedinae
175
certainly have such subvertical laterosternites,
including the oldest cupedines (Kirghizocupes).
Comparative studies of structures of the Recent
Cupedinae and Ommatinae show that the mod-
ern Omma and Tethrapalerus maintain a greater
number of plesiomorphic characters (Hörn-
schemeyer et al., 2006) which, according to the
cladogram created on the basis of a matrix by
these authors, could form the sister group to all
other Recent archostematans (including Crow-
soniella Pace, 1976 and Micromalthus LeCon-
te, 1878, but not Sikhotealinia Lafer, 1996).
Futhermore, Beutel et al. (2008: 292) found that
the “Ommatidae” could be “placed as the sister
group of the remaining Archostemata, includ-
ing the extinct families Ademosynidae, Schizo-
phoridae and Catiniidae. The monophyly of this
clade is supported by one apomorphy of adults,
the absence of the tentorial bridge, and by seven
apomorphies of larvae. The larval features
strongly support a clade comprising Microma-
lthidae and Cupedidae, but not Ommatidae”.
Unfortunately, because tentorial bridges of adults
are not available in most fossils, also because
only larvae of Micromalthidae are known among
fossil archostematans (no fossil cupedid larva
has been found so far), and because a compari-
son based only on a few Recent archostematan
larvae is certainly not enough for a reliable
reconstruction of the whole phylogeny of recent
and fossil taxa, the hypothesis proposed by
these authors can scarcely be tested in a proper
way without data on fossil larvae. Atkins (1958a),
among other things, noted that Priacma serrata
lacks membranous lobes on the tarsomeres, but
this species shows what seem to be reduced or
rudimentary empodia. At the same time, extant
members of Omma have distinct bisetose empo-
dia. It is noteworthy that some recent compara-
tive studies conclude that “representatives of
Archostemata are characterized by many de-
rived features of the head, which are partly due
to a more exposed lifestyle and partly to special-
ized pollen-feeding habits” (Hörnschemeyer et
al., 2002: 312). However, the particularly stable
morphotype of many taxa in the subfamily Cu-
pedinae beginning at least from the Middle/late
Triassic gives reason to suggest that both “life-
style” and “feeding habits” of cupedine larvae
and adults may have maintained stable for a very
long time (at least from the Middle/late Trias-
sic) and they could have been basal for further
changes in both “lifestyle” and “feeding habits”
of the polyphagans which might have derived
from a cupedine-like ancestor. Crowson (1975)
regarded that the boring holes in trunks of Arau-
carioxylon Knowlton, 1888 (Araucariaceae) “in
petrified forest” of Arizona could have being
made by cupedine larvae. Such habits could be
associated with an appearance of resins in the
Mesozoic conifers caused by protection from
beetle attacks. Thus, similarity of mouthparts
and other structures in the Jurassic and modern
cupedines can be explained by similar mode of
life including regular visiting plant generative
organs. The archostematans used for compari-
sons by neontologists mostly represent the mod-
ern fauna, i.e. only few terminal phylogenetic
branches. In particular, the extant cupedines,
except for Priacma serrata, with similar head
tuberculation patterns and partly covered anten-
nal insertions, seem to be descendants of the
same phylogenetic lineage of the subfamily.
The difference in the male genital structures of
Priacma and Gracilicupes, on one hand, and
those of other genera with known male genita-
lia, on the other hand, seems to support the
viewpoint that these lineages which produced
the Recent (1) Priacma and (2) other genera
represented in the modern fauna must be rather
old, although some diversification events in
these genera could be quite recent, most proba-
bly during the Cenozoic. This was the reason for
an unresolved polytomy in the strict consensus
tree obtained by Beutel et al. (2008). This latter
paper, as well as some others published by
specialists on the modern fauna, contain no
studies on fossil beetles and consider the char-
acters of Recent species alone, the results being
directly and uncritically extrapolated on fossil
species, with some features of Recent represen-
tatives frequently being assigned to fossils.
However, the distribution of plesiotypic and
apotypic characters available in the extant mem-
bers of a group rarely corresponds to the se-
quence of diversifications, at least such situa-
176
A.G. Kirejtshuk et al.
tion is clearly observed in cupedines whose
fossil diversity is incommensurably greater than
recent diversity. After a rather broad compari-
son of many species of the modern fauna even
more paradoxical interpretation have been ob-
tained by Lawrence et al. (2011: 65): “Within
the Archostemata, neither Ommatidae nor Cu-
pedidae is monophyletic, Ommatinae (com-
ments: these authors only mean Recent species
of Omma) forming a clade with Cupedinae
(comments: these authors only mean all Recent
cupedines except Priacma serrata) and Tet-
raphalerinae (comments: these authors only mean
Recent species of the genus Tetraphalerus) with
Priacminae (comments: these authors only mean
Priacma serrata), while Micromalthidae is sis-
ter to the remaining taxa.” Not surprisingly,
such a hypothesis of relationships fails to corre-
spond to what we could observe here. Nor can it
be applied to the fossil material examined or re-
examined in the present paper.
The fossil specimens studied so far allow for
the structural cupedine type or cupedine mor-
photype to be admitted as having evolved at a
very early stage of the historical development of
the family, which also included a plesiotypic
pattern of elytral venation, a significant convex-
ity of the body, the elytra with steeply sloping
sides and reduced epipleura, and also the Priac-
ma-type male genitalia. The Priacma-type aede-
agus is to be recognized as probably plesiomor-
phic because it is hardly possible at the moment
to admit a reversed transformation of the de-
rived and less complex cupedine type back to
the plesiomorphic and more complex Priacma-
type. If the characters accompanying the plesio-
typic aedeagus could also be considered as
somewhat plesiotypic, the similarities of the
oldest cupedines to Priacma serrata may tenta-
tively be postulated as general features of cupe-
dine morphotype.
The Middle/late Triassic Kirghizocupes, one
of the oldest cupedine genera, shares with Pri-
acma serrata the rather slender medium-sized
body, the more or less steeply sloping sides of
the elytra (but apparently not as vertical as in
Cupes and its closest allies), the ground-plan
elytral venation pattern), 10 (or 11) long longi-
tudinal rows of cells on the elytra, the tubercu-
late head and (sub)filiform antennae (see above
“Notes to Kirghizocupes” and Chapter 9). How-
ever, Kirghizocupes differs from Priacma in
one additional row of cells visible in some prints
just between Sc and the lateral edge of the elytra,
the reduction or lack of fusion of primary veins
at the elytral apices, the rather long head with
paramedian convexities behind the antennal in-
sertions forming a more or less expressed Y-
shaped depression in the anterior part of the
frons, the open antennal insertions, the rather
wide pronotum and very widely (sub)explanate
sides. Therefore, the distinguishing characters
of Priacma can preliminarily be recognized as
apomorphic, albeit the lack of fusion of primary
veins at the elytral apices in some specimens of
Kirghizocupes is to be regarded as secondary. It
is noteworthy that the above distinguishing char-
acters of Kirghizocupes are characteristic not
only of members of Cupedinae, but also of
Asimma and Pterocupes, both of uncertain po-
sitions, partly also of some other groups initially
included in triadocupedines. Thus, these char-
acters could provisionally be consideredas sym-
plesiomorphic. The very wide explanate prono-
tal sides occur mostly in the Triassic cupedines
(Kirghizocupes and Asimma) and triadocu-
pedines (Cupesia Ponomarenko, 1966, Platy-
cupes and Triadocupes). Among other cu-
pedines, only Latocupes (including Pulchicu-
pes) has such a wide pronotum with so widely
explanate sides, which seems better to be inter-
preted as a secondarily developed similarity.
The comparatively long head occurs in the cu-
pedine Kirghizocupes and triadocupedine Pro-
cupes Ponomarenko, 1966, as well as many
Ommatinae. The comparatively deep Y-shaped
depression in the anterior part of the frons
associated with the peculiar convexities on the
head are observed in many Mesozoic cupedines.
Ponomarenko (1966) mentioned this character
only for Mesocupoides (a junior synonym of
Kirghizocupes) among Triassic cupedids, but
the elongate paramedian convexities behind open
antennal insertions can be traced back to numer-
ous Mesozoic cupedines, triadocupedines and
ommatines; in triadocupedines with a wide head,
Taxonomy of the reticulate beetles of the subfamily Cupedinae
177
the lateral branches of this depression barely fail
to form a transverse groove with a slight (very
blunt) angle in the middle. Probably this struc-
tural peculiarity cannot be unambiguously in-
terpreted in all cases; however its distribution
across many groups of the family allows to
suppose some probability of a symplesiomor-
phy to be admitted for most of these cases. The
open antennal insertions are somehow structur-
ally associated with the tubercles and depres-
sions on the frons (considered above). If the
convexities or tubercles are not raised or located
behind the antennal insertions, they leave the
antennal insertions open. If these convexities or
tubercles shift anteriorly, they cover the poste-
rior edge of the antennal insertions while the Y-
shaped depression on the frons becomes dis-
placed further forward, where the frons is trans-
formed into a small triangular depression before
the antennal insertions (as in all Recent cu-
pedines). So the formation of covered antennal
insertions seems to have frequently been ho-
moplastic, albeit in the Recent species of most
genera similar to Cupes they may have had a
phylogenetic succession because of close rela-
tionships expressed in a considerable similarity
of most Recent taxa (see above Chapter 4).
All three subfamilies of the family Cupe-
didae (Triadocupedinae, Ommatinae and Cupe-
dinae) appeared in the fossil record as clearly
defined forms only since the Middle/late Trias-
sic. The Triadocupedinae is only represented in
the Middle/late Triassic while the Ommatinae
and the Cupedinae have survived until now.
Besides this, beginning from the Middle Trias-
sic, elytra with characteristic venation patterns
and steep sloping of elytra have been found, but
regrettably they are impossible to accurately
assign to any of the subfamilies (see above
Chapter 8). Nevertheless, the lack of earlier
evidence of cupedines seems better to be ex-
plained by scarcity of material obtained from
the early Triassic. The possibly plesiomorphic
character of the aedeagus in Priacma might
have been inherited from the ancestor of Ho-
lometabola. Therefore, the cupedines could have
emerged from a taxon with a similar aedeagal
type. Plausible candidates could be either an
early Permian tshekarcocoleid or any late Per-
mian related group. The male genitalia of such
groups are still unknown (see above) and no
Triassic cupedid with exposed male genitalia
has been recorded. It seems likely that a close
relative of Permocupes and the “ancestor” of
cupedids may have had a sister-group relation.
Besides this, two alternative evolutionary sce-
narios could be presumed. Either taxa directly
related to the Palaeozoic Tshekarcocoleidae or
any late Permian group reached the Middle/late
Triassic without representation in the fossil
record to give rise to the Cupedidae, or the
Mesozoic Cupedidae appeared during the times
when the Palaeozoic groups were more numer-
ous, but the cupedids had failed to enter fossil
deposits till the Middle/late Triassic. The latter
option seems to be more viable, taking into
account the significant similarities between the
elytra of Permocupes and many cupedids (see
above).
Changes of the number of long longitudinal
rows of cells on the elytra may have followed a
progressive decrease in number from 10 or 11 to
nine. This process is characteristic of different
groups and occurs independently, as it was
demonstrated in the modern groups by Neboiss
(1984). The opposite change, i.e. an increase of
the number of rows of cells could also be admit-
ted. In Cainocups aixensis sp.n., the number of
long longitudinal rows was increased to 12
because the primary vein R adjoined, through a
longitudinal inflection, the flat disk of the elytron
and the steep sloping of the elytral sides in this
species proceeded along R, but not along M (as
usual in cupedines). Nevertheless, it seems more
reasonable to provisionally join C. aixensis sp.n.
with Cenozoic archostematan groups rather than
search for another explanation for placing this
taxon in the classification of the suborder.
Many structural details easily accessible in
modern cupedines simply cannot be analyzed to
the same extent in fossil representatives. Atkins
(1958a) considered that the living Priacma ser-
rata have maintained the conditions that are
more archaic than those occurring in the mem-
bers of the genus Cupes. This author mentioned
that, in Cupes, abdominal ventrites 2 and 3 are
178
A.G. Kirejtshuk et al.
connate, without any apparent suture between
them. Unfortunately, this structural peculiarity
which could be important for tracing the phy-
logeny is not testable in fossils. Nevertheless,
this conclusion strongly contradicts the timing
of the distribution of the overlapping (tegular)
type of articulation in the abdominal ventrites
among the Permian and early Mesozoic archo-
stematans, which can be traced not only in most
fossil cupedids (including many fossil members
of Cupes), but also in Permian tshekardocoleids,
as well as the distribution of the abutting (flat)
type present in Priacma serrata, other cupedids
and archostematans. The venation of the hind-
wings in Priacma serrata is more complex than
in other cupedid hindwings previously described,
there being more cross-veins and six enclosed
cells (Atkins, 1958b). The persistence of the
more complex venation with numerous cross-
veins is usually interpreted as plesiotypic. Ac-
cording to Atkins (1958a: 533), “Priacma ser-
rata has a larger number of cross veins than
Cupes, which might indicate that Priacma is the
more primitive of these two genera”. Of course,
this agrees with differences in the type of aede-
agus (see above Chapter 9).
Another interesting aspect is the difference
in the pair of genera Paracupes s.lato and Pri-
acma, as well as in the pair of subgenera Para-
cupes s.str. and Paracupoides subgen.n. The
former pair shows many similarities in external
characters and a striking contrast in structure of
the male genital sclerites. If the divergence in
the genital structure of the common “ancestors”
of these groups happened long time ago, the
similarities in external structure should be sup-
ported by a certain ecological conditionality to
maintain such a great stability. Such condition-
ality could be great conservatism in habitat
preferences and the mode of life (lifestyle) at
least over two hundred million years (since the
Middle/late Triassic). Similar cases are repre-
sented not only in the ommatine Omma and
Tetraphalerus, but also in the genera Jurodes
Ponomarenko, 1985 (early–late Jurassic) and
Sikhotealinia (modern) from the archostematan
family Jurodidae. Thus, such a conservatism in
structure and probably also in ecology and bion-
omy is somehow more or less characteristic of
cupedomorphan Archostemata in general. At
the same time, this conservatism during such a
long time is combined with a comparatively
rapid transformation in the types of elytra dis-
covered in the studies of the pair of Paracupes
s.str. and Paracupoides subgen.n. (see above
Chapter 6). This reccurrence of the structural
type of not too closely related “groups” (Para-
cupoides ascius comb.n., on the one hand, and
modern species of the genera near Cupes, on the
other hand) could be accounted for by a mor-
phocyclic process of conservative ground-plans
causing some convergence (Emel’yanov, 2000),
because it seems to be completely impossible to
link genealogically one of these groups (Para-
cupes s.str. and Paracupoides subgen.n.) to
Priacma and the other to Cupes. In a formal
sense, this could be made using a matrix of the
characters traditionally used in a cladistic anal-
ysis of Cupedinae (e.g. Hörnschemeyer, 2009).
It is noteworthy in the context of the pairs
considered that the oldest of the known cu-
pedines had their elytra with steeply sloping
sides and 10 long longitudinal rows (Figs 109–
116) like those in Cupes and Paracupoides
subgen.n., rather than those in Priacma and
Paracupes sensu str. Thus, the elytra with steeply
sloping sides similar to those in Recent Cupes
could have appeared many times and scarcely
manifest a reliable evidence of common ances-
try without additional, more reliable support.
The next type of similar differences can be
observed in the elytra of the subgenus Anag-
lyphites stat.n. Mesocupes (Anaglyphites) cap-
itatus comb.n. shows a row of cells between
elytral Sc and the lateral edge (Figs 95–98),
whereas the elytra of M. (A.) admotus comb.n.
and M. (A.) clavatus comb.n. (Figs 101–102)
bear densely tuberculate (crenelate) stripes along
the lateral edge, similar to those in Priacma and
Paracupes s.lato. These types of difference
demonstrate a mosaic distribution of many char-
acters among Recent and fossil cupedines.
The subfamily Cupedinae, as cupedids in
general, certainly had a much greater signifi-
cance in the Mesozoic faunas than in the Ceno-
zoic ones. Having appeared in deposits since the
Taxonomy of the reticulate beetles of the subfamily Cupedinae
179
Middle Triassic, the cupedids in general repre-
sent up to one-third of the coleopterous remains
in the Jurassic deposits, the other greatest pro-
portions being comprised by schisophoroids
from Archostemata and various polyphagans
(mostly elateroids) (Ponomarenko, 1995, 2002,
2003; etc.). Cupedidae as a major and the best-
known Mesozoic group occurred only in com-
paratively hot climatic zones (Ponomarenko,
2016). The Recent representatives of that fam-
ily, however, do not clearly show such prepon-
derance. All extant cupedids with known bion-
omy exist owing to the wood of angiosperms, a
plant group absent in the Jurassic and earlier.
Changes in food plants seem to have not become
effected in structural features of the family.
Therefore, the ecology and bionomy of its fossil
and Recent members, despite the above-men-
tioned differences, were and probably still are
more dependant on the structure of wood tis-
sues, also on the fungal and microbial composi-
tions associated with wood rather than the sys-
tematic position of food trees.
The triadocupedines disappeared in the fos-
sil record at the end of the Triassic (Kirejtshuk,
Ponomarenko, 2016) while both other subfam-
ilies have reached the present times. They are
now represented by a small number of species
regarded as relict groups scattered in a mosaic
pattern across the areas with temperate and
subtropical climates. At the same time, the cu-
pedines (31 species) are more diverse in the
modern fauna than ommatines (only six spe-
cies). Data on the Cenozoic faunas of cupedines
are still scant and many species already collect-
ed and deposited in scientific collections are
still awaiting a study. At least, in addition to the
published cupedine species (Gersdorf, 1976;
Tröster, 1993; Kirejtshuk, 2005), some unde-
scribed species from the Eocene deposits which
could mostly belong to Cupes, as well as some
others undescribed genera are known to the
authors from Baltic amber (Museum of Amber,
Kaliningrad, ZIN and several private collec-
tions), Messel (SFNFM) and Eckfeld (LNNR).
In addition, separate apices of cupedine elytra
have been revealed in younger outcrops, i.e, the
boundary between the Eocene and Oligocene of
Bembridge Marls (Ponomarenko, Kirejtshuk,
in press) and the latest Oligocene of Aix (Cain-
ocups aixensis sp.n.); the youngest cupedine
remains are represented by Cupes praeglacia-
lis from the late Pliocene of Willershausen,
although its generic assignment needs a further
revision. Thus, during the Neogene the subfam-
ily Cupedinae can be supposed to have de-
clined, probably in accordance with global cli-
matic changes with reinforced seasonality and
zonality. In Europe, the Cupedinae could have
disappeared during the Pleistocene due to glaci-
ations. At least in Europe, this group survived
the Pliocene global cooling.
The Paleocene fauna of Menat seems to
have had a considerable representation of Cupe-
dinae, consisting of some genera as remains of
the Mesozoic fauna [at least Mesocupes (Cain-
omerga subgen.n.)], although at the same time
some close relatives of the modern fauna seem
to have also been represented by Cupes species.
This representation should be associated with
forest ecosystems around the place of their
deposition. The general composition of beetles
in the outcrops of Menat includes a significant
proportion of weevils (more than half of beetle
remains), also many specimens of Ceramby-
cidae Latreille, 1802, Elateridae Leach, 1815,
Nitidulidae Latreille, 1802, Buprestidae Leach,
1815, eumolpine-like leaf beetles (Chrysome-
lidae Latreille, 1802) and some other arboreal
groups which certainly were dominant at that
locality at the time of deposition (Nabozhenko,
Kirejtshuk, 2014).
Conclusions
1. Taxonomic and phylogenetic aspects:
– Cupedinae represents one of the oldest
groups of the order and some of its forms main-
tain traces of “pre-coleopteran” features (Priac-
ma-type aedeagus in Recent Priacma and fossil
Gracilicupes inherited from probable ancestor
of the Holometabola) and therefore early diver-
gence of the order Coleoptera can preliminary
defined as Tshekardocoleidea + {Schizopho-
rimorpha + [Cupedinae + (Ommatinae + other
families of Cupedomorpha)]};
180
A.G. Kirejtshuk et al.
– The differences between subfamilies of
the Cupedidae are not so great and some genera
proposed in composition of Triadocupedinae
should be considered in composition of Cupedi-
nae [Kirghizocupes (=Mesocupoides syn.n.), ?
Asimma and ? Pterocupes];
– Cupedinae is regarded as a separate sub-
family which is not possible to divide into the
tribes and, therefore, the tribes formerly pro-
posed (Priacmini and Mesocupedini) are re-
garded as not reasonable taxa;
– The cupedines demonstrate a great stabil-
ity in structure and sculpture of elytra and body
appendages (antennae, mouth parts and legs),
but some variability is exposed mostly in the
proportions of body sclerites, structure of tho-
rax and epicranium;
– Most fossil taxa described before this
publication could not be put in Cupedinae or
Ommatinae, frequently even cannot be recog-
nized among other generic taxa because of not
relable available characters or partly confusing
original descriptions;
– Recent cupedines represent remains of a
small part of ancient diversity (Mesozoic and
Paleogene) and seem to consist of two group of
the closely related (one of them is Priacma and
second is all other taxa);
– In order to reach more or less equal hiati
between Recent taxa and make them somehow
comparable with extinct taxa, the taxa Adinole-
pis and Ascioplaga are regarded as subgenera of
the same genus (Adinolepis) and for Paracupes
ascius was proposed a new subgenus (Paracu-
poides subgen. n.).
2. Distributional aspects:
– Cupedinae have been recorded in deposits
of the Middle/late Triassic from Middle Asia:
[Kirghizocupes (=Mesocupoides syn.n.), Asim-
ma and Pterocupes), Europe (Helopides, Meta-
cupes, Nannocurculionites, Parabuprestites,
Paracurculionites and Pseudosilphites] and
Australia (Mesothoris), although position of
some Triassic forms needs to be confirmed by
further studies;
– Portions of cupedines in Mesozoic (begin-
ning from Middle/late Triassic) and Paleogene
(not later than Eocene) outcrops are decreasing
in later deposits; in the Neogene outcrops cu-
pedines became rare and only 31 species have
been found in the modern fauna diffusely spread
in different continents, although in the Recent
European and Antarctic faunas the cupedines
are not known;
– Cenozoic extinct faunas in Europe repre-
sent considerable species and generic diversity
of Cupedinae during the Paleogene (Menat,
Messel, Eckfeld, Baltic amber), however later
they became rather rare in deposits and reached
only the Pliocene (Cupes praeglacialis).
3. Ecological aspects:
– Maintenance of many structural characters
in all Recent species in the state not changed
from the Middle/late Triassic ones testifies an
extremely stable preference of habitats and ex-
traordinal conservatism mode of life (probably
since of time of formation of this subfamily);
– such stability can be maintained only un-
der immutable conditions of life of the cu-
pedines during the whole time of existence of
this subfamily. These conditions could be pro-
vided by the larval development in interstices of
tree trunks and comparatively short imaginal life;
– Diet of cupedines seems to have depen-
dence on the structure of wood tissues, also on
the fungal and microbial compositions associat-
ed with wood but not on systematic position of
plants producing wood (gymposperms or ang-
isperms).
Acknowledgements
The authors have a pleasure to receive generous
assistance from many colleagues. Many thanks to
Clotilde Berger-Pompili and Leygnac Mathilde, di-
rectors of the EHPAD du Pays de Menat, for their
kind help and authorisations to collect fossil insects
in a small but rich outcrop near the village of Menat
(Puy-de-Dôme, France). Very special thanks to Jean-
Marc Pouillon who helped in the field to collect
material. Particular thanks are expressed to A.G.
Ponomarenko (PIN), who provided the authors with
many consultations and advices in study the speci-
mens examined. S. Wedmann (SFNFM), H. Lutz
(LNNR), C. Gröhn (GPIM), W. Weitschat (GPIM),
Chr. and H.W. Hoffeins (Hamburg), G. Herrling
(Engter), T. Hörnschemeyer (GZUG), and F. Kerneg-
ger (Hamburg) essentially promoted the studies by
assistance in acquisition of specimens examined or
Taxonomy of the reticulate beetles of the subfamily Cupedinae
181
photographs of them. The authors also sincerely
thank S. Pont (Laboratoire de Minéralogie et de
Cosmochimie, MNHN) who helped the authors in
the study of the fossils by electronic microscopy. T.
Deuve, O. Montreuil, and A. Mantilleri (UMR 7205,
Entomologie, MNHN) helped in management of the
specimens from the collection of the modern beetles
in MNHN and M.V.L. Barclay in BMNH. An essen-
tial help in preparation of the manuscript was ob-
tained from S.I. Golovatch (Institute for Problems of
Ecology and Evolution, Moscow) who checked En-
glish in its most important part and provided the
authors with many valuable linguistic consultations.
Bo Wang (Institute of Geology and Palaeontology of
Chinese Academy of Sciences, Nanjing) and Mingyi
Tian (South China Agricultural University, Guang-
dong) helped in translation and transliteration of
Chinese papers. A.A. Prokin (Papanin Institute for
Biology of Inland Waters of Russian Academy of
Sciences, Borok, Yaroslavl Oblast) and Huali Chang
(CNU) assisted the authors to prepare photographs
of the specimens examined from the collection of
CNU. D.Ye. Shcherbakov (PIN) made some photo-
graphs and assisted in studies of the specimens from
the collection of PIN. A.V. Kovalev (ZIN) helped in
preparation of photographs of the specimens from
the collection of ZIN and discussed with authors
different aspects of studies and preparation of this
paper. R.E. Nelson (Department of Geology, Colby
College, Waterville, Maine, USA) facilitated to find
contacts with S. Lubkin who allowed the authors to
use her pictures from Lubkin (2003). S.A. Beloko-
bylskij (ZIN), S.Yu. Sinev (ZIN) and I.V. Shamshev
(ZIN) provided the authors with valuable consulta-
tions on the structure of male genitalia of different
groups of Holometabola. The studies of the first
author were partly carried out under the framework
of the Russian state research project no.
01201351189, programme ‘Research in Paris’ of the
City of Paris (Mairie de Paris), programme of the
Presidium of the Russian Academy of Sciences “Prob-
lems of the origin of life and formation of the
biosphere”, Sorbonne Universités (Programme
d’Accueil de Chercheurs de Haut niveau) and the
Russian Foundation of Basic Research (grant 15-04-
02971-a). The authors greatly appreciate useful com-
ments on an earlier version of the manuscript re-
ceived from anonymous reviewers.
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Responsible editor K.G. Mikhailov
Appendix 1. Material studied
Fossil species
Subfamily Cupedinae
– Apriacma clavata (Tan, Ren et Shih, 2006), comb.n. [Priacma] — holotype (“CNU-C-
LB2006001”).
– (?) Apriacma latidentata (Tan, Ren et Shih, 2006), comb.n. [Priacma] — holotype (“CNU-
C-LB2005010”).
– Apriacma renaria (Tan, Ren et Shih, 2006), comb.n. [Priacma] — holotype (“CNU-C-
LB2006002”).
– Apriacma tuberculosa (Tan, Ren et Shih, 2006), comb.n. [Priacma] — holotype (“CNU-C-
LB2005011”).
– Cupes distinctissimus sp.n. — holotype (“MNHN.F.A.51118”).
– Cupes eckfeldensis (Troester, 1993), comb.n. (Tenomerga) — holotype (LNNR “1992/461”).
– Cupes groehni Kirejtshuk, 2005 — holotype (“4420”, “4071 ex. Coll. C. Gröhn”, GPIM) and
4 paratypes (“372”, DEIM (coll. Chr. and H.W. Hoffeins); “4421”, “4318 ex Coll. C. Gröhn”,
GPIM; “790”, from coll. Herrling; “792”, from coll. Herrling).
– Cupes hoffeinsorum Kirejtshuk, 2005 — holotype (“1036-2”, DEIM (coll. Chr. and H.W.
Hoffeins)).
– Cupes kerneggeri Kirejtshuk, 2005 — holotype (“1036-2”, ZIN (coll. H. Kernegger)).
– Cupes komissari Kirejtshuk, 2005 — holotype (“III, B233”, “Cupes tesselatus (Motschulsky,
1856), Yu. Popov det. 1990”, GZUG).
– Cupes manifestus Kirejtshuk, Nel et Colomb, 2010 — holotype (“MNT-06-902”, MTM,
print and counterprint) and 5 additional specimens (“MNHN.F.A.52770”, print and counterprint;
“MNHN.F.A.52771”, print; “MNHN F.A. 57525”, print; “MNHN F.A. 57526”, print: “MNHN
F.A. 57527”, print).
– Cupes messelensis (Troester, 1993), comb.n. (Tenomerga) — holotype (SFNFM “MeI
2806”).
188
A.G. Kirejtshuk et al.
– Cupes ponomarenkoi Kirejtshuk, Nel et Colomb, 2010 — holotype (“PA 6457”, MNHN).
– Cupes rohdendorfi Iablokoff-Khnzorian, 1960 — holotype (“PIN 364/105”) and six
additional specimens (“3/1999”, ZIN (coll. H. Kernegger); “360”, GPIM (coll. C. Gröhn); (“999-
2”, DEIM (coll. Chr. and H.W. Hoffeins) (“999-2”, DEIM (coll. Chr. and H.W. Hoffeins); “GPIM-
22”; “792”, from coll. Herrling).
– Cupes simillimus sp.n. — holotype (“MNHN.F.A5 1120”).
– Cupes tesselatus (Motschoulsky, 1856) [Cupoides] — neotype (“¹ 4422”, GPIM).
– Cupes weitschati Kirejtshuk, 2005 — holotype (“PA 6457”, MNHN).
– Cupidium abavum Ponomarenko, 1968 — holotype (“PIN 2384/398”).
– Forcicupes raucus Tan et Ren, 2006 — holotype (“CNU-C-LB2006004”, print and
counterprint).
– Gracilicupes crassicruralis Tan, Ren et Shin, 2006 — holotype (“CNU-C-LB2005001”).
– Gracilicupes tenuicruralis Tan, Ren et Shin, 2006 (=tenuocruralis: Tan & Ren, 2009) —
holotype (“CNU-C-LB2005002”).
– Kirghizocupes indistinctus (Ponomarenko, 1969), comb.n. — holotype (“PIN 2240/72”,
print and counterprint).
– Kirghizocupes proporeius (Ponomarenko, 1969), comb.n. — holotype (“PIN 2240/3”, print
and counterprint) and three specimens (“PIN 2240/82”, “PIN 2240/104”, “PIN 2069/1034”).
– Kirghizocupes sp. — one specimen (“PIN 3064/6424”, print and counterprint).
– Latocupes bellus Tan et Ren, 2006 — holotype (“CNU-C-LB2005013”).
– Latocupes collaris (Tan, Huang et Ren, 2006), comb.n. — ? holotype (? “NIGPAS 142147”,
found in CNU without number).
– Latocupes fortis Tan et Ren, 2006 — holotype (“CNU-C-LB2005012”).
– Menatops orbiculatus (Kirejtshuk, Nel et Colomb, 2010), comb.n. — holotype (“MNT-05-
198”( MTM), print and counterprint).
– Menatops bartenevi sp.n. — holotype (“MNHN F.A. 57524” , print and counterprint).
– Mesocupes (Anaglyphites) admotus (Ponomarenko, 1964), comb.n. — holotype (“PIN 2066/
2293”) and 11 possible additional specimens (M. (A.) pr. admotus comb.n.: “PIN 2066/2280”,
“PIN 2066/2279”, “PIN 2066/2271”, print and counterprint), “PIN 2384/412”, “PIN 2465/976”,
“PIN 2554/438”, “PIN 2465/968”, “PIN 2384/405”, “PIN 2239/1505”, “PIN 2066/2277”, “PIN
2239/804”).
– Mesocupes (Anaglyphites) capitatus (Ponomarenko, 1966), comb.n. — holotype (“PIN
1989/2993”) and 1 additional specimen — (!) counterprint of holotype (“PIN 1989/1869”).
– Mesocupes (Anaglyphites) clavatus (Ponomarenko, 1964), comb.n. — holotype (“PIN 2066/
2275”, print and counterprint) and 9 additional specimens (M. (A.) pr. clavatus comb.n.: “PIN
2384/402”, “PIN 2384/403”, “PIN 2384/406”, “PIN 2384/408”, “PIN 161/167”, “PIN 2232/807”,
“PIN 2232/808”, “PIN 2232/809”, “PIN 2066/2276”, “PIN 2066/2285”).
– Mesocupes (Anaglyphites) minimus (Ponomarenko, 1997), comb.n. — holotype (“PIN 3589/
6052”, print and counterprint ).
– Mesocupes (Anaglyphites) mongolicus (Ponomarenko, 1997), comb.n. — holotype (“PIN
3559/1608”) and two paratypes (“PIN 3559/6055”, “PIN 3790/277”).
– Mesocupes (Cainomerga) brevicornis sp.n. — holotype (“MNHN.F.A51119” , print and
counterprint).
– Mesocupes (Cainomerga) fraternus sp.n. — holotype (“MNHN.F.A52773”).
– Mesocupes (Cainomerga) immaculatus (Piton, 1940), comb.n. — holotype (“MNHN
R07012”).
– Mesocupes (Cainomerga) palaeocenicus sp.n. — holotype (“MNHN.F.A51117”).
– Mesocupes (Cainomerga) ponti sp.n. — holotype (“MNHN.F.A51116”, print and counter-
print).
Taxonomy of the reticulate beetles of the subfamily Cupedinae
189
– Mesocupes (Cainomerga) sp.1 — 1 specimen (“MNHN.F.A.57518”, print)
– Mesocupes (Cainomerga) spp. — 4 specimens (“MNHN.F.A.52769”; “MNHN.F.A.57519”,
print and counterprint; “MNHN.F.A.57520”; “MNHN.F.A.57521”, print).
– (?) Mesocupes (Cainomerga) sp.1 — 1 specimen (“MNHN.F.A.52768”).
– (?) Mesocupes (Cainomerga) sp.2 — 1 specimen (“MNHN.F.A. 57522”).
– (?) Mesocupes (Cainomerga) sp.3 — 1 specimen (“MNHN.F.A. 57523”).
– Mesocupes (Mesocupes) primitivus Martynov, 1926 — neotype (“PIN 2452/682”) and three
additional specimens (“PIN 2452/196”, “PIN 2066/2272”, “PIN 2066/2283”).
– Mesocupes (Mesocupes) minor Ponomarenko, 1968 — holotype (“PIN 2239/812”) and one
additional specimen (“PIN 2239/810”, print and counterprint).
– Mesocupes (Mesocupes) paulus (Ponomarenko, 1964), comb.n. — holotype (“PIN 1789/
91”) and two additional specimens (M. (T.) pr. paulus comb.n.: “PIN 2384/411”, “2384/405”).
– Mesocupes (Mesocupes) bidens Ponomarenko, 1964 — holotype (“PIN 2465/980”).
– Mesocupes (Mesocupes) spinosus Ponomarenko, 1964 — holotype (“PIN 2066/2294”, print
and counterprint).
– Priacmopsis adumbrata Ponomarenko, 1966 — holotype (“PIN 1989/3000”, print and
counterprint) and 1 additional specimen (“PIN 1989/3032”, print and counterprint).
– Taxopsis motschulskyi (Kirejtshuk, 2005), comb.n. — holotype (“PIN 363/130”) and one
paratype (“3750”, GZUG).
– Genus incertus sp.1 — 1 specimen (MNHN “R 63906”).
– Genus incertus sp.2 — 1 specimen (“MNHN.F.A.52794”).
– Genus incertus sp.3 — 1 specimen (“MNHN.F.A.52772”).
– Genus incertus sp.4 — 2 specimens (“MNHN.F.A.57529”, “MNHN.F.A.57530”).
Subfamily Ommatinae
– Brochocoleus angustus Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006009”).
– Brochocoleus sulcatus Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006011.
– Cionocoleus cervicalis Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006007”).
– Cionocoleus planiusculus Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006008”).
– Lithocupes punctatus Ponomarenko, 1969 — holotype (“PIN 2240/326”).
– Notocupes alienus Tan et Ren, 2006 — holotype (“CNU-C-LB2005001”).
– Notocupes eumerus (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005007”).
– Notocupes minusculus (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005005”).
– Notocupes porrectus (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005006”).
– Notocupes protensus (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005008”).
– Notocupes psilatus (Tan, Ren et Liu, 2005) — holotype (“CNU-C-LB2004001”).
– Notocupes rudis (Tan, Ren et Liu, 2005) — holotype (“CNU-C-LB2004004”).
– Notocupes sibiricus (Ponomarenko, 2000) — 1 additional specimen (“PIN 2385/192”).
– Notocupes tylodes (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005003”).
– Notocupoides triassicus Ponomarenko, 1966 — holotype (“PIN 2070/617”, print and
counterprint).
– Tetraphalerus curvinervis Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006006”).
– Tetraphalerus latus Tan, Ren et Shih, 2007 — holotype (“CNU-C-LB2006005”).
– Tetraphalerus trachylaenus (Tan, Ren et Shih, 2006) — holotype (“CNU-C-LB2005009”).
Subfamily Triadocupedidae
– Cupesia serricornia Ponomarenko, 1969 — holotype (“PIN 2240/85”).
– Triadocupes ferghanensis Ponomarenko, 1966 — holotype (“PIN 2240/85”).
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Family Cupedidae (subfamilia incerta)
– Cupedites minutissimus Ponomarenko, 1985 — holotype (“PIN 1256/33”) and three
additional specimens (C. pr. minutissimus: “PIN 1256/39”, “PIN 1256/75”, “PIN 1256/76”).
– ‘Priacma’ corrupta Ponomarenko, 1986 — holotype (“PIN 3152/1302”).
– ‘Priacma’ oculata Ponomarenko, 1997 — holotype (“PIN 4271/182”, print and counter-
print).
– ‘Priacma’ longicapitis Ponomarenko, 1997 — holotype (“PIN 4271/182”, print and
counterprint).
Recent species
– Adinolepis (Adinolepis) mathesonae (Neboiss, 1960) — 1 specimen (MNHN, “Vic., Pascoe
Vale, Melbourne, 25 Dec. 1960, H. Hellgerien”).
– Adinolepis (Adinolepis) youanga (Neboiss, 1960) — 1 specimen (ZIN, “27.30S 152.58E,
Austr., Indooroopilly, QLD, 17-21.12.1990, A. Kirejtshuk”).
– Adinolepis (Ascioplaga) mimeta (Neboiss, 1984), comb.n. — holotype, male (MNHN, “Mt
Koghi, 15.iii. 1978 (Delobel)”); 1 paratype, male (“For & De La Thy, i.1960 (Chazeau & Saile)”).
– Cupes anguliscutis Kolbe, 1886 — 19 specimens (MNHN, ZIN), Southern China, Annam,
Tonkin.
– Cupes concolor Westwood, 1835 — 13 specimens (MNHN, ZIN), from different localities
of USA.
– Cupes capitatus Fabricius, 1801 — 25 specimens (BMNH, MNHN, ZIN), different localities
of USA.
– Cupes leucophaeus Newman, 1839 — 3 specimens (BMNH, MNHN), South Africa.
– Cupes mucidus Chevrolat, 1829 — type of C. mucidus (MNHN, “Cupes mucidus Chevr,
Yc.R.a. Porte P.58, des. Philippines”), lectotype and one paralectotype of C. clathratus (ZIN,
“Vladivostok, 13–15.vii.1868, Cupes cancellatus mihi”) and more than 120 specimens (MNHN,
ZIN), Russian Far East (Primorsky Kray), Japan, Philippines.
– Cupes varians Lea, 1902 [Distocupes] — 1 specimen, “Austral.”.
– Paracupes (Paracupes) brasiliensis Kolbe, 1898 — 1 specimen (MNHN), “Brasil” and one
specimen (MNHN) “Bahia”.
– Priacma serrata LeConte, 1861 — about 100 specimens (males) (MNHN, ZIN), different
localities of western states of USA.
– Prolixocupes lathriellei (Solier, 1849) — holotype (MNHN, “Illapel”, “Museum Paris, Chili,
Cl. Gay, 1845”, “TYPE”, “Cupes Latreillei, Santjago et Illapel”) and more than 30 specimens
(MNHN) — Chili.
– Prolixocupes lobiceps (LeConte, 1874) — 4 specimens (BMNH, MNHN), California.
– Rhipsideigma adjuncta Neboiss, 1984 — holotype (MNHN, “Rhipsideigma adjuncta sp. n.,
det, Neboiss, 1983, Holotype, #”, “Andringitra Est, Anjavidilava, 2000 m, 18. xii .– 15 .i. 1971”,
“l/FDSMP/” “Museum Paris, Madagascar Centre Mission C. N . R.S .I RCP no 225”).
– Rhipsideigma anosibense Neboiss, 1984 — holotype (MNHN, “Rhipsideigma anosibense
sp. n., det, Neboiss, 1988, holotype, #”, “Anosibe, d-te 3-67, Collection Dujardin-Delacour ”,
“Madagascar Nord, 1500–2500 m”).
– Rhipsideigma cretaceotincta (Kolbe, 1887) — one specimen (BMNH, “Tanganyika,
31.iii.1955, ex Piphadenia sp., dead part living tree (Gardner).
– Rhipsideigma lugubris (Fairmaire, 1895) — 15 specimens (MNHN), Madagascar.
– Rhipsideigma raffrai (Fairmaire, 1884) — holotype (MNHN, $, “Cupes Raffrayi Fairmaire,
Madagascar/Museum Paris/Madagascar/Collection Leon Fairmaire 1906” and about 40 speci-
mens (BMNH, MNHN) — Madagascar.
