Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea

The paper is the first of a series, which aims to present a consistent interpretation of the suprageneric taxa of fossil beetles in the current century and their generic and species composition. Order Coleoptera is considered in composition of the superorder Coleopteroidea Handlirsch, 1903 (= Coleopterida sensu Boudreaux, 1979, nec Pearse, 1936) together with orders Skleroptera and Strepsiptera, and also with the family Umenocoleidae of unclear position. This paper includes the archostematan superfamilies Coleopseoidea and Cupedoidea of the infraorder Cupediformia, i.e., Coleopseidae (one genus and one species), Tshekardocoleidae (12 genera, 15 species), Labradorocoleidae (one genus, one species), Permocupedidae (together with Taldycupedinae, stat. nov., 24 genera and 54 species) and Cupedidae (three subfamilies, 49 genera, 253 species). The preliminary information on structure of the larva of Tshekardocoleidae from Tshekarda is done. There are also described the new taxa: genus Afrotaldycupes Kirejtshuk, gen. nov. with the type species: genus Taldycupes africanus Ponomarenko in Ponomarenko & Mostovski, 2005 [Afrotaldycupes africanus comb. nov.] and Afrotaldycupes lidgettoniensis (Ponomarenko in Ponomarenko & Mostovski, 2005), comb. nov. [Taldycupes]; genus Allophalerus Kirejtshuk, gen. nov. with the type species: Tetraphalerus aphaleratus Ponomarenko, 1969 [Allophalerus aphaleratus comb. nov.], and also with Allophalerus antiquus (Ponomarenko, 1964), comb. nov. [Tetraphalerus], Allophalerus bontsaganensis (Ponomarenko, 1997), comb. nov. [Tetraphalerus], Allophalerus incertus (Ponomarenko, 1969), comb. nov. [Tetraphalerus], Allophalerus latus (Tan, Ren et Shih, 2007), comb. nov. [Tetraphalerus], Allophalerus maximus (Ponomarenko, 1968), comb. nov. [Tetraphalerus], Allophalerus okhotensis (Ponomarenko, 1993), comb. nov. [Tetraphalerus], Allophalerus tenuipes (Ponomarenko, 1964), comb. nov. [Tetraphalerus], Allophalerus verrucosus (Ponomarenko, 1966), comb. nov. [Tetraphalerus]; genus Bukhkalius Kirejtshuk et Jarzembowski, gen. nov. with the type species: Tetraphalerus lindae Jarzembowski, Wang et Zheng, 2017 [Bukhkalius lindae comb. nov.]; genus Burmocoleus Kirejtshuk, gen. nov. with the type species: Burmocoleus prisnyi sp. nov. and Burmocoleus zhiyuani (Liu, Tan, Ślipiński, Jarzembowski, Wang, Ren et Pang, 2017), comb. nov. [Brochocoleus]; genus Cionocups Kirejtshuk, gen. nov. with the type species: Cionocups manukyani sp. nov.; genus Echinocups Kirejtshuk et Jarzembowski, gen. nov. with the type species: Notocupes neli Tihelka, Huang et Cai, 2020 [Echinocups neli comb. nov.], and also Echinocups ohmkuhnlei (Jarzembowski, Wang et Zheng, 2020), comb. nov. [Notocupes] and Echinocups denticollis (Jiang, Li, Song, Shi, Liu, Chen et Kong, 2020), comb. nov. [Notocupes]; genus Jarzembowskops Kirejtshuk, gen. nov. with the type species: Brochocoleus caseyi Jarzembowski, Wang et Zheng, 2016 [Jarzembowskops caseyi comb. nov.]; genus Lobanovia Kirejtshuk, gen. nov. with the type species: Simmondsia permiana Ponomarenko, 2013 [Lobanovia permiana comb. nov.]; genus Pintolla Kirejtshuk, gen. nov. with the type species: Kaltanicupes ponomarenkoi Pinto, 1987 [Pintolla ponomarenkoi comb. nov.]; genus Polyakius Kirejtshuk, gen. nov. with the type species: Polyakius alberti Kirejtshuk, sp. nov. and Polyakius pubescens Kirejtshuk, sp. nov.; Clessidromma zengi Kirejtshuk, sp. nov.; Cupes golovatchi Kirejtshuk, sp. nov.; Cupes legalovi Kirejtshuk, sp. nov.; Cupes lutzi Kirejtshuk, sp. nov.; Cupes nabozhenkoi Kirejtshuk, sp. nov.; Cupes wedmannae Kirejtshuk, sp. nov.; Mallecupes prokini Kirejtshuk, sp. nov. and Omma janetae Kirejtshuk, sp. nov. The new synonymy is established for the generic names Clessidromma Jarzembowski, Wang et Zheng, 2017 and Lepidomma Jarzembowski, Wang et Zheng, 2019, syn. nov. The rank of Cainomerga A. Kirejtshuk, Nel et P. Kirejtshuk, 2016 is elevated from subgeneric to generic. Also other new combinations are proposed: Cainomerga brevicornis (A. Kirejtshuk, Nel et P. Kirejtshuk, 2016), comb. nov. [Mesocupes], Cainomerga fraterna (A. Kirejtshuk, Nel et P. Kirejtshuk, 2016), comb. nov. [Mesocupes], Cainomerga immaculata (Piton, 1940: 194), comb. nov. [Zonabris, Mesocupes], Cainomerga palaeocenica (A. Kirejtshuk, Nel et P. Kirejtshuk, 2016), comb. nov. [Mesocupes], and Cainomerga ponti (A. Kirejtshuk, Nel et P. Kirejtshuk, 2016), comb. nov. [Mesocupes], Clessidromma tianae (Jarzembowski, Wang et Zheng, 2019), comb. nov. [Lepidomma], Diluticupes applanatus (Tan et Ren, 2009), comb. nov. [Brochocoleus], Diluticupes crowsonae (Jarzembowski, Yan, Wang et Zhang. 2013), comb. nov. [Brochocoleus], Diluticupes magnus (Tan et Ren, 2009), comb. nov. [Brochocoleus], Diluticupes minor (Ponomarenko, 2000), comb. nov. [Brochocoleus], Diluticupes validus (Tan et Ren, 2009), comb. nov. [Brochocoleus], Diluticupes yangshuwanziensis (Jarzembowski, Yan, Wang et Zhang. 2013), comb. nov. [Brochocoleus], Monticupes curtinervis (Tan, Ren et Shih, 2007), comb. nov. [Tetraphalerus], Monticupes decorosus (Tan, Wang, Ren et Yang, 2012), comb. nov. [Tetraphalerus], Odontomma sulcatum (Tan, Ren et Shih, 2007), comb. nov. [Brochocoleus], Omma ancistrodontum (Tan, Wang, Ren et Yang, 2012), comb. nov. [Pareuryomma], Omma grande (Ponomarenko, 1964), comb. nov. [Tetraphalerus], Omma longicolle (Ponomarenko, 1997), comb. nov. [Tetraphalerus], Pareuryomma angustum (Tan, Ren et Shich, 2007), comb. nov. [Brochocoleus], Pareuryomma magnum (Tan et Ren, 2009), comb. nov. [Brochocoleus], Zygadenia aliena (Tan et Ren, 2006), comb. nov. [Ovatocupes], Zygadenia baojiatunensis (Hong 1992), comb. nov. [Chengdecupes], Zygadenia brachycephala (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia caduca (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia caudata (Ponomarenko, 1966), comb. nov. [Notocupes], Zygadenia cellulosa (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia crassa (Ponomarenko, 1969), comb. nov., [Notocupes], Zygadenia cyclodontus (Tan, Ren, Shih et Ge, 2006), comb. nov. [Amblomma, Notocupes], Zygadenia dischdes (Zhang, 1986), comb. nov. [Notocupes], Notocupes dundulaensis (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia elegans (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia epicharis (Tan, Ren et Liu, 2005), comb. nov. [Amblomma, Notocupes], Zygadenia eumeura (Tan, Ren et Liu, 2005), comb. nov. [Amblomma, Notocupes], Zygadenia excellens (Ponomarenko, 1966), comb. nov. [Notocupes], Zygadenia exigua (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia foersteri (Ponomarenko, 1971), comb. nov. [Procarabus, Notocupes], Zygadenia homora (Lin, 1986), comb. nov. [Conexicoxa, Notocupes], Zygadenia issykkulensis (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia jurassica (Hong 1983), comb. nov. [Chengdecupes], Zygadenia kezuoensis (Hong 1987), comb. nov. [Chengdecupes], Zygadenia khasurtuiensis (Strelnikova, 2019), comb. nov. [Notocupes], Zygadenia khetanensis (Ponomarenko, 1993), comb. nov. [Notocupes], Zygadenia kirghizica (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia laeta (Lin, 1976), [Tetraphalerus], Zygadenia laiyangensis (Hong et Wang, 1990), comb. nov. [Forticupes, Notocupes], Zygadenia lapidaria (Ponomarenko, 1968), comb. nov. [Notocupes], Zygadenia laticella (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia lata (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia lenta (Ren, Lu, Guo et Ji, 1995), comb. nov. [Tetraphalerus], Zygadenia lini (Ponomarenko, Yan, Wang et Zhang, 2012), comb. nov. [Notocupes], Zygadenia longicollis (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia ludongensis (Wang et Liu, 1996), comb. nov. [Notocupes], Zygadenia minuscula (Tan, Ren, Shih et Ge, 2006), comb. nov. [Amblomma, Notocupes], Zygadenia mongolica (Ponomarenko, 1994), comb. nov. [Notocupes], Zygadenia nigrimonticola (Ponomarenko, 1968), comb. nov. [Notocupes], Zygadenia oxypyga (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia patula (Ponomarenko, 1985), comb. nov. [Notocupes], Zygadenia pingi (Ponomarenko et Ren, 2010), comb. nov. [Notocupes], Zygadenia porrecta (Tan, Ren, Shih et Ge, 2006), comb. nov. [Amblomma, Notocupes], Zygadenia protensa (Tan, Ren, Shih et Ge, 2006), comb. nov. [Amblomma, Notocupes], Zygodenia psilata (Tan, Ren et Liu, 2005), comb. nov. [Amblomma, Notocupes], , Zygadenia pulchra Ponomarenko, 1968, comb. nov. [Notocupes], Zygadenia reticulata (Oppenheim, 1888), comb. nov. [Procarabus, Notocupes], Notocupes rostrata (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia rudis (Tan, Ren et Liu, 2005), comb. nov. [Amblomma, Notocupes], Zygadenia shiluoensis (Hong 1984), comb. nov. [Chengdecupes], Zygadenia sogutensis (Ponomarenko, 1969), comb. nov., Zygadenia stabilis (Tan, Ren et Liu, 2005), comb. nov. [Amblomma, Notocupes], Zygadenia tenuis (Ponomarenko, 1969), comb. nov. [Notocupes], Zygadenia tripartita (Oppenheim, 1888), comb. nov. [Procarabus, Notocupes], Zygadenia tuanwangensis (Hong et Wang, 1990), comb. nov. [Picticupes, Notocupes], Zygadenia valida (Lin, 1976), comb. nov. [Sinocupes, Notocupes], Zygadenia vitimensis (Ponomarenko, 1966), comb. nov. [Notocupes].


Introduction
This publication is the first of a series, which aims to provide a consistent interpretation of the suprageneric taxa of fossil beetles in the current century and their generic and species composition. The preparation of such an attempt is quite complicated, because probably no recent palaeocoleopterist can be competent in all groups of the order. Another great problem is to find a reasonable compromise between alternative opinions on the placement of established taxa proposed by somebody. The author of this publication is aware of a lot of complexities in the attempt undertaken by him and possible criticism from Many other researchers who are more experienced in Many groups of the order studied by the present author. A serious particular problem is connected with correct interpretation of the previous descriptions and illustrations supplementing them, most of which contains rather ambiguous or inadequate information on the described forms. In most cases the formerly described species need to be retested and redescribed for a correct usage. Nevertheless, the general importance of such understanding is they can provide preliminary generalizations to put together a great quantity of facts obtained by the author and borrowed from literature in order. Many years of previous debates on this Matter among neontologists and palaeontologists clarified several complex problems but were not enough to produce a balanced result. However, even taking into consideration the deficiencies published in the previous taxonomic generalizations on fossil beetles, such as Handlirsch [1], Rohdendorf [2], Carpenter [3] and others, it is difficult to overestimate their significance in research and usage in different fields of knowledge. In recent decades, publication of numerous separate annotated lists of fossil beetles on separate groups or palaeofaunas provided Many very important contributions (for examples, reviews and catalogues [4][5][6][7][8][9][10][11][12][13][14][15][16][17][18][19][20][21][22][23], and Many others).
Ponomarenko and Zherikhin, palaeocoleopterists of the Paleontological Institute of the Russian Academy of Sciences, prepared the "catalogue" (a filing cabinet including Many carboard cards with used or proposed names for fossil beetles), later Ponomarenko put a considerable part of this information in some computer DOC-files. The next step of development of the basis for this publication was a preparation of the "Catalogue of fossil Coleoptera" presented on the site of "Beetles (Coleoptera) and coleopterists" of the Zoological Institute of the Russian Academy of Sciences (https://www.zin.ru/animalia/coleoptera/rus/index.htm). This preparation was Made in 2002 and 2003 with the very essential contribution of Lobanov who helped to find a way presenting this "catalogue" on the Internet and after that Lobanov devoted years in adding new data and correcting previous data in the web-pages. Participation of Ponomarenko in preparation of the web-version of this catalogue during few first years was rather essential, however, later he could not find time for contribution in the development of this catalogue and, as a result, Many important corrections and changes in it were Made without his attention and agreement. Nevertheless an initial impulse of collecting of taxonomic facts from palaeontological literature to put them in Internet appeared thanks to the Ponomarenko's desire to save the data of carboard cards for use by colleagues from over the world.
During the current century other resources of data on fossil beetles appeared and got a rather quick development. One of most important and a rather large source of different palaeontological  Genus Uralocupes Ponomarenko, 1969 [26]. Type species: Kaltanicupes Major Ponomarenko, 1963 [69], by monotypy. One species.
Diagnosis. This new genus is similar to Kaltanicupes, but differs from it in the Markedly more weakly curved veins and longitudinal rows of cells on its elytra (veins in the photographs of the holotype of Pintolla ponomarenkoi comb. nov. are scarcely observable, although in one of the photographs the traced veins correspond to those in the drawing-reconstruction: [67]: 6, Figure 1; Pl. 1, Figure 1a,b). In contrast to Many permocupedines (including Kaltanicupes), Pintolla ponomarenkoi comb. nov. has not any clear triangle of cells between Cu and A1 at the elytral base. Besides, the prescutellar cells of its elytra demonstrate more than one row of cells and there are two rows of cells between R and M, M and Cu, and also Cu and A1.
Etymology. The name of the new genus is formed from the name of the descriptor of the type species (Irajá Damiani Pinto) and suffix "lla". Gender feminine.

Notes on the Subfamily Permocupedinae
The taxon Afrocupes was synonymized with Permocupes on-line by Ponomarenko and Kirejtshuk [29] because its elytral venation is very similar to that in Martynov's Permocupes and "Permocupoides", although the cells in the distal part of the elytron seem to become smaller more gradually.
The genus Archicupes was initially proposed for two species (type species, A. jacobsoni, and A. reichardti), whose holotypes were collected in the same locality (Kaltan) in the Kuznetzk Basin. The holotype of the first species is represented only by the left elytron, while that of the second one has the body and some parts of the appendages. Ponomarenko [26] synonymized Archicupes jacobsoni with Palaeocupes kaltanicus, but the second initial congener of Archicupes (A. reichardti) was excluded from this genus by this author, because A1 (A2 after Ponomarenko [26]) of the second species is Markedly longer than in the type species. Nevertheless, elytra of both species of Archicupes included by Rohdendorf show considerable similarity, except for the length of the mentioned anal vein. Ponomarenko [26] wrote that Archicupes reichardti could belong to Kaltanicupes and Eocupes without providing any additional argument. It is a good idea to revise these three genera or perhaps all permocupedines to determine the relation between the genera currently included in this subfamily.
According to Ponomarenko [26] the type specimen of Kaltanocoleus pospelovi has the subcostal area and base of elytron missing, but its anal field is similar to that in species of Kaltanicupes and Eocupes. Indeed the holotype of this species at the time of the Rohdendorfian description had a clear venation and cells that were drawn and pictured ( [2]: Text- Figure 319 and Pl. XXIX, Figure 190) demonstrating the rather distinct R branch. This genus and species can be diagnosed with the Rohdendorf's [2] keys.
The holotype elytron of "Taldycupes" chachlovi Rohdendorf, 1961 [2] (Russia: Kemerovo Region (Suriekova), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5-268.0 Ma) is longer, less convex and finely microtuberculate in comparson with other species of Taldycupes. It has eight rows of large cells, instead of ten as in other species of this genus. Thus, the later species seems to belong to another genus. However the holotype of this species is represented only by the apical part of the elytron bearing the same characteristics as Taldycupedidae and Permocupedidae. Ponomarenko [26] put this species in the latter family because of the "primary" veins of its elytra are different from other interspaces between longitudinal rows of cells.
The genus Protocupes Rohdendorf, 1961 [2] is represented by two species, type species: Protocupes Martynovi Rohdendorf, 1961 [2] and another congener: P. ogloblini Rohdendorf, 1961 [2]. Both species originated from the same locality (Kaltan) in the Kuznetzk Basin (Russia: Kemerovo Region (Kaltan), Permian, Cisuralian/Guadalupian, Kungurian/Roadian (Cisuralian/Biarmian, Kungurian/Kazanian), −272.5-268.0 Ma) and illustrated only by the drawing-reconstructions. The holotype of the former species is represented by a considerable distal part of the elytron ( [2]: figures 330,331), while the holotype of the latter species shows only a small part of the elytral base ([2]: figure 332). Ponomarenko [26] regarded these species separately as genera incertae sedis. The last author published another drawing-reconstruction of the type species of this "genus" different from that published by Rohdendorf, but again without providing this reconstruction with a photograph. Thus, it is impossible to clarify which of these reconstructions is more correct or more adequate for the remainder of the specimen. As a result, the genus Protocupes should be considered as a taxon incertae sedis among members of Permocupedidae and with unclear composition (one or two species).
Stegocupes efremovi Rohdendorf 1961 [2] was described together with the type species of this genus (S. fedotovi Rohdendorf 1961 [2]). Both species originated from the same locality (Kaltan) in the Kuznetzk Basin and have some similarity in the body shape, microtuberculate integument of the elytra and thickened apical 5-6 antennomeres. Ponomarenko [26] redescrided Stegocupes efremovi taking into consideration its pronotum as different from that in other taldycupedids but the dorsal integument of the elytra is not clear in its holotype; as a result, he transferred this species to the permocupedids (i.e., permocupedines), although the later description does not include the peculiar antennal structure, microtuberculate integument of the elytra and uniform elytral venation (first descriptor considered that in Stegocupes a trend to reduction of venation and cells together with sclerotization of the elytral integument is to be observed [2]). Both descriptions of Stegocupes efremovi [2,26] do not contain the photographs and, therefore, it is difficult to choose the more correct one without re-examination of the holotype. This species has been preliminarily re-transferred back to the taldycupedine genus Tecticupes (which is provisionally regarded as the senior synonym of Stegocupes). Rohdendorf, 1961 Notes. Afrotaldycupes gen. nov. is proposed after the current observation of characteristics of the elytral structure and, therefore, it should be regarded as a "formal" or "collective" genus ("morphogenus") as in Many cases of permocupedids. In contrast to Afrotaldycupes gen. nov., the Eurasian members of Taldycupes demonstrate elytra with regular longitudinal rows of subquadrate to subpolygonal cells, and only the elytron of T. cellulosus shows subquadrate to a little suboval cells disposed in the regular and somewhat curved rows and veins ending to the elytral apex.

Subfamily Taldycupedinae
Etymology. The name of this new genus is formed from the name of the continent (Africa) and generic name Taldycupes. Gender masculine.  [96] indicated that this generic taxon is based on a part of the elytron with destroyed upper surface and, therefore, it cannot be identified as a member of any family, although it differs from other early Mesozoic beetles in the peculiar pattern of the elytral punctation. The genus Penecupes Ren, 1995 [98] with the type species: Penecupes rapax Ren, 1995 [98] (China: Beijing Shi (Chongqing Reservoir), Lower Cretaceous, Late/Upper Aptian, −122.5-112.6 Ma) was transferred to the subfamily Protorabinae Ponomarenko, 1977 [30].

Family Cupedidae Laporte, 1836
Type genus: Cupes Fabricius, 1801 [99] (Three subfamilies, 47 genera, 249 species) Comments. This family has a very distinct structural base in Many body sclerites and appendages, including the elytral venation, which is Maintained or can be traced in different phyletic branches, including two of them reaching the modern epoch (Figures 4-24). This feature seemed to combine with parallel structural transformations, on one hand, and very frequent disappearance in specialized groups existed in various geological periods. As a result, the groups slightly structurally modified are present in the Recent fauna, while the more advanced apotypic representatives left only in the fossil record. Ponomarenko [26] elaborated a key to the cupedid subfamilies, which was later emended by Kirejtshuk et al. [33]. Nevertheless the characteristics in them should still be regarded as tentative because their diagnostic value is not enough reliable. The separated or contiguous procoxae represent the alone feature which can be used for discrimination of cupedines and ommatines. If these characteristics are not observable an assignement of fossils to any supraspecific taxon becomes more or less depending on available additional characteristics associated with certain groups but not diagnostic or restrictedly diagnostic ones for these groups.
In addition, the genera and species included in this review, the descriptions of two new ommatine genera with four species recently described are put in the Appendix A (see below). Comments. This subfamily is distinct among fossil cupedids mostly in the contiguous procoxae.

Subfamily Ommatinae Sharp et Muir, 1912
The key to tribes of this subfamily for fossil beetles was elaborated by Ponomarenko [26], however later the description of Many new generic taxa of Ommatinae Makes it necessary to revise Many taxa with a wide comparison of all groups of the subfamily. Perhaps, the ommatine genera without prothoracic lateral carinae ( Figure 4C-F, 13 and 14) are more closely related and should be united in one tribe (Ommatini sensu stricto). On the other hand, it is necessary to take into consideration that some structures are characterized by a strong trend to similar modifications in a definite group (i.e., elytral cells and punctation) or parallel transformations in different and not closely related groups (i.e., some structural changes in elytral venation).
Other congeners: Allophalerus antiquus (Ponomarenko, 1964)     Notes. Ponomarenko ([110]: 89) after re-examination of the holotype of Blapsium egertoni represented by only the underside of metathorax, metacoxae, abdomen and elytral epipleura came to the conclusion that it belongs to the archostematan tribe Notocupedini because of the characteristic metepisterna, metacoxae and short overlapping abdominal ventrites. He also considered that this species was apterous because of its rather short metathorax.  [113]. . Taking into consideration other species here included in this genus, it can be defined that Brochocoleus is characterized by the rather wide and comparatively robust body with the elytra having Sc curved along the elytral lateral edge so that the other veins without fusion terminate on it apically, and by the longitudinal rows of more or less clearly expressed cells and extremely widely explanate elytral sides with very large cells arranged in 2-3 rows of large cells (typically two long rows and at base one short row of cells between them). The head in species of this genus is long and with long temples. Their prothorax is narrow, arcuate at sides and with more or less expressed lateral carinae. The abdominal ventrites of this group are co-planar (abutting). Another group with somewhat similar elytral venation, but with a slenderer body, prothorax distinctly carinate and explanate at sides, elytra having fused A1 with Cu, and M with R just at the elytral apex, and also double rows of cells, irregular rows of microtubercles or with diffuse microtubercles (or punctures) along the not very widely explanate elytral sides and shoulder corner not projecting anteriorly has a sufficient morphological hiatus with Brochocoleus in order to regard it as a separate taxon. Ren [98] proposed the generic taxon, Diluticupes Ren, 1995 [98], for one species of this group (Diluticupes impressus Ren, 1995 [98]) after his study of the specimen with the dorsal and underside body sclerites, Mandibles, one antennae and left anterior leg. Its holotype elytron has five veins, two rows of cells between them, and very wide explanate sides with dense microtuberculation. Besides, the original drawing shows subparallel veins ( [98]: 77: Figures 3-37), although his photographs of Diluticupes impressus ( [98]: Pl. 11, 1-2) Make it possible to trace fused A1 and Cu. The additional specimen of this species studied by Jarzembowski et al. [108] clearly shows this fusion. Nevertheless, the name Diluticupes was synonymized with Brochocoleus to no purpose [11,29]. Tan et al. [115] divided the species mixture of Brochocoleus with Diluticupes into two groups as the "Brochocoleus punctatus series" and "B. minor series", which more or less correspond to the division into the genera here accepted. Thus, the genus Diluticupes can reasonably include all members of the "Brochocoleus minor series" sensu Tan et al. [115].

Genus
The re-examined holotype of "Brochocoleus indibili" Soriano et Delclòs. 2006 [116] (Spain, Sierra del Montsec (La Cabrua outcrop), Lower Cretaceous, Early/Lower Barremian, −130.0-125.5 Ma) is so badly preserved that it is impossible to be sure that this species is really a member of Diluticupes because of the absence of any strict evidence for its archostematan attribution. The prothorax of the latter species with the very widely explanate sides and strongly excised anterior edge is very different from that in the species of both Brochocoleus and Diluticupes. The procoxae of the above-mentioned specimen are scarcely traceable to clarify if they are contiguous for a supposition on a probable similarity with ommatines (as was supposed by the descriptors). Therefore, "Brochocoleus indibili" should be regarded as a species without generic, family and subordinal attribution (incertae sedis).
Finally, Brochocoleus angustus Tan, Ren et Shih, 2007 [115] should be regarded as a member of the genus Pareuryomma [Pareuryomma angustum, comb. nov.] because of its rather slender body, characteristic head structure, narrow prothorax and reduced cells and venation on its elytra, and also because of the comparatively large cells on the explanate elytral sides. See also the Notes to the genus Pareuryomma.
Genus Burmocoleus Kirejtshuk, gen. nov. (Figures 5 and 6    Notes. After the current studies of Jarzembowski et al. [118] the position of this group became rather clarified. This genus is characterized by the elongate body with finely and diffusely microreticulated integument, comparatively long head with rather small eyes and slightly expressed neck, prothorax subspherical with incarinate sides, elytra somewhat explanate at sides and their lateral edges smooth (not densely serate) and without clear veins or at most reduction or lack of cell development on the elytra, abdominal ventrites co-planar (abutting). The polygonal (to subquadrate) cells are apparently located along the elytral lateral edges and arranged in weakly expressed longitudinal rows. The prothorax of some species of Cionocoleus seems to have no or apparently only weak remnants of lateral carinae.
Cionocoleus ommamimus does not fit with other congeners as its head has prominent temples, its elytra are extremely narrowly explanate at the sides, and its abdominal ventrites could be overlapping. The descriptor drawing ( [105]: 54, Figure 13) shows a prothorax different from his photograph ( [105]: Pl. VI, 16), but it seems that the prothorax of Cionocoleus ommamimus is indeed incarinate. Jarzembowski et al. [118] placed Cionocoleus punctatus in the genus Cionocoleus, although its longitudinal seriation in the elytral sculpture and apparently extremely narrowly explanate sides of the elytra give reasons to doubt of this attribution (some longitudinal seriation is also visible in the photograph of Cionocoleus ommamimus). At the same time Cionocoleus tanae looks like a quite typical member of this genus despite strongly reduced explanate sides of the elytra.   considerably longer than pro-and mesotarsi, flat (co-planar) abdominal ventrites. Clessidromma palmeri is very similar to Lepidomma tianae and differs from the latter only in the outlines of body sclerites giving an impression that the first is the second, which was strongly extended in length. The third species recovered during the recent studies looks like somehow intermediate between the first and second species described before, although every of these three species has some additional specific diagnostic features [101,121] (see below the Descriptions of new taxa of the subfamily Ommatinae).       Genus Diluticupes Ren, 1995 Figures 4 and 5); nevertheless, it is clear that the elytra of the holotype have a fused A1 and Cu at the apex. The head of the type species also has differences in the author's illustrations mentioned above and is needs to be checked by a re-examination of the type specimen. Thus, Monticupes is characterized by the moderately oval body, subtriangular or long (subparallelsided) head with large eyes and well raised temples, pronotum carinate and with subexplanate sides, veins well expressed with fused A1 and Cu at apex and their common vein ending on Sc, explanate elytral sides moderately wide and with diffuse small microtubercles, and abdominal ventrites co-planar (abutting). The head of the type species also has differences in the author's illustrations mentioned above and also needs to be checked by a re-examination of the type specimen. The members of this genus, in contrast to those of Allophalerus gen. nov., demonstrate the fusion of A1 and CuA at the elytral apex and different shape of their head (see below the comparison of Allophalerus gen. nov.) Genus Notocupoides Ponomarenko, 1966 [102]. Type species Notocupoides triassicus Ponomarenko, 1966 [102], by original designation. Three species.
Notes. The most peculiar feature of this genus is that the elytra of its species have well expressed veins and longitudinal rows of cells reaching the sutural edge without fusion. Besides, Notocupoides is characterized by the long head with short temples and long frons before the eyes, rather wide pronotum with widely explanate and carinate sides, intervenal stripes of the elytra with two cells in longitudinal rows, explanate sides with one row of rather large cells, rather short antennae, and abdominal ventrites overlapping. Notes. This genus is rather similar to Diluticupes, but the veins of Odontomma elytra are not fused at the apices and the head of its members, in contrast to species of Diluticupes, is longer and not narrowed anteriorly. Thus Odontomma is characterized by the moderately oval body, moderately more or less long and subparallel or oviform head with comparatively small eyes, rather wide pronotum with the widely explanate carinate sides, all veins ending on Sc along the lateral elytral edge, veins well expressed, explanate elytral sides with diffuse small microtubercles, and abdominal ventrites co-planar (abutting) or looking like bordered along the posterior edge.     Notes. This genus still remains rather problematic because of difficulties with analysis of the accessible characteristics in the descriptions and illustrations of its fossil members and comparison with the characteristics of the type species and other recent representatives. Therefore, most fossil species included here in this genus need to be re-examined. The extinct genus that is most closely related to Omma is Polyakius gen. nov., which differs from the extant genus mostly in some level of dorsoventral compression of the body, and also in the presence of one row of very large cells along the lateral elytral edges and very dense and fine dorsal vestiture.
Ponomarenko [129] put Procarabus zitteli in Omma, but in his redescription of this species there are no reliable characteristics to support such an attribution. According to the original descriptions, Omma aberrantum has explanate elytral sides, while O. jurassicum demonstrates the narrowly explanate elytral sides; besides, O. antennarum Ponomarenko, 1997 [105] has the narrow and quadrangular prothorax (apparently with lateral carinae) and the very narrowly explanate elytral sides.
On the other hand, "Pareuryomma" ancistrodontum shows all the diagnostic characteristics to be transferred to Omma (characteristics of its head and elytra), although its pronotum has a rather large anterior orifice. Omma daxishanense was temporally placed in Omma, but at least its head with very wide frons, very prominent temples and distinct antennal grooves provide evidence to put it in Allophalerus gen. nov. rather than to another ommatine genus. Probably the same pertains to Omma brevipes, whose attribution can be clarified only after re-examination of the type of this species. Omma elongatum was initially described in the genus Carabus [126], however later Handlirsch [1] proposed to put it in a separate genus [Chalepocarabus], Crowson [131] transferred it to Omma and Ponomarenko [110] restored the genus Chalepocarabus for this unique species ("elongatus") in the subfamily Cupedinae and prepared the incorrect drawing with the prosternal process and separated procoxae in the holotype of Omma elongatum ( [110]: 97, Figure 5). Kirejtshuk et al. [33] followed the latter opinion of Ponomarenko. The current assessment of the last two redescriptions and illustrations of this species [110,128] Make it possible to find only the characteristics that can be interpreted as ommatine ones (the outlines of the body sclerites (particularly, subglobular prothorax and shape of head), also, the venation and cells of elytra in the photograph of Crowson ([131]: Pl. IV, Figure 9) fit better with those of the genuine members of Omma in comparison with the photograph in Ponomarenko ( [110]: Pl. 10, Figure 7)). "Tetraphalerus" grandis better fits with Omma by the outlines of its body sclerites, than with Tetraphalerus or Allophalerus gen. nov. and, therefore, it is preliminarily put in the genus under consideration, and also "Tetraphalerus" longicollis apparently like Omma has the very narrow explanate elytral sides and prothorax clearly without lateral carinae. Ponomarenko [130] put Procarabus zitteli in Omma, but in his redescription of this species there are no reliable characteristics to support such an attribution. According to the original descriptions, Omma aberrantum has explanate elytral sides, while O. jurassicum demonstrates the narrowly explanate elytral sides; besides, O. antennarum Ponomarenko, 1997 [105] has the narrow and quadrangular prothorax (apparently with lateral carinae) and the very narrowly explanate elytral sides.
On the other hand, "Pareuryomma" ancistrodontum shows all the diagnostic characteristics to be transferred to Omma (characteristics of its head and elytra), although its pronotum has a rather large   moderately widely explanate sides of the elytra, and abdominal ventrites co-planar (abutting). Pareuryomma angustum comb. nov. differs from other congeners by the shorter and rather narrowing head and seems to be able to be considered as a separate genus. The generic name Euryomma was erroneously synonymyzed with Notocupes by Kirejtshuk et al. in 2010 [11]. Later Tan et al. in 2012 [125] proposed a new name Pareuryomma for the mentioned junior homonym.    The recent photograph of the holotype of Omma zitteli ( [129]: Pl. VII, 1) shows the remnants of a specimen similar to Allophalerus gen. nov. rather than to any other genus, although because of a lack of reliable character for generic diagnostics it can be regarded as attributed to genus incertus. See also below the description of this new genus and comments on it. Notes. According to the described holotype of the type species, this genus is represented in the positive and negative print and counterprint only showing the metatorax and abdomen with the most part of the elytra and characterized by an elongate oval body with the elytra finely and diffusely punctured as on disc as on explanate sides, very weakly traced veins and co-planar (abutting) abdominal ventrites. To clarify other characteristics of this genus it is necessary to find some Cenozoic congener with a better preservation. Notes. If the characteristics used in this publication for other generic diagnoses of fossils are applied to the modern species, this genus can be characterized by the elongate body, elongate and slightly anteriorly narrowed head with small eyes or eyes covered with processes of the frons, very long and prominent temples, narrow prothorax without sharp lateral carinae (modern species of Tetraphalerus have lateral carinae on the prothorax, although they are obscured by the microtubercles), elytra with more or less expressed veins (A1 and Cu fused at apex or independently ending on Sc), explanate elytral sides which are very narrow and without cells, and abdominal ventrites co-planar (abutting). At the same time, it is scarcely reasonable to expect fossil relatives of the modern South American species of Tetraphalerus sensu stricto to be found among Mesozoic cupedids and most species which were formerly put in this genus are here considered in the composition of Allophalerus gen. nov. (see below the Description of the latter genus). Some Mesozoic species are still temporally remain in this genus with a question Mark and their attribution needs to be checked by a further re-examination of type material.

Genus
According to the original description, (?) Tetraphalerus brevis has the pronotum shorter than head, flattened and with sharp lateral edges ([103]: 53: "переднеспинкa короче головы, поперечнaя, ее боковой крaй уплощенный и острый") and, on the other hand, a rather narrow epipleuron ("эпипплеврa довольно узкaя"), although its photograph Makes it possible to suppose that prothorax seems to have rounded sides ( [103]: Pl.VI, 3) but not subquadrangular as drawn in the original description ([103]: 52, 2b). (?) Tetraphalerus collaris has the characteristics of the modern species in its head, its pronotum and elytral base fit better with the modern species than those in other Mesozoic species. However in order to be sure of its attribution it is necessary to study similar fossil representative with a better preservation. (?) Tetraphalerus glabratus somewhat corresponds to Tetraphalerites oligocenicus, but in the original description it was mentioned that its elytral "cells" (not punctures) are scarcely observable. The holotype of (?) Tetraphalerus mongolicus has its body outlines like those in other "? Tetraphalerus" species, but it shows no peculiar important character for a reliable generic diagnostics, although the descriptor compared it with "T. verrucosus" [139]. The type series of (?) Tetraphalerus notatus includes different imprints. The descriptor did not indicate the holotype in the list of the examined specimens, but in the caption to " Figure 6" there is mentioned as "the holotype" the number which was not included in the list of the examined specimens. The description and illustrations of (?) Tetraphalerus notatus contain characteristics from different imprints which do not give any reason to assign this species to any precise genus. Thus, this species name should be reasonably considered without generic attribution.
Genus Zygadenia Handlirsch, 1906 [1] (Figures 18 and 19      Notes. This genus with its wide elytral explanate sides is similar to Brochocoleus and Jarzembowskiops gen. nov., but quite different from them in its prothorax. It is characterized by the very wide body in the elytra, moderately long and subtriangular head with slightly prominent eyes and very prominent temples, comparatively narrow to moderately wide and sharply carinate prothorax, elytral veins not expressed, more or less irregular cells of the elytral disc and explanate elytral sides with four or more rows of very large cells (more or less visible at least at the elytral base), and abdominal ventrites apparently co-planar (abutting). Notes. This genus is diagnosed based on the following characteristics: head elongate, with longitudinal grooves and slightly narrowing anteriorly (frons before eyes more or less long); antennae short and scape usually longest among antennomeres; pronotum wide and distinctly carinate at sides, its anterior and posterior angles with distinct top; elytra with explanate sides bearing one long row of large cells along Sc, A1 and Cu fused before apex and common vein ending on suture, intervenal stripes with two rows of cells. The group with this diagnosis is most abundant and rather variable in Mesozoic deposits and, therefore, it acquired so Many synonyms. However, its variability is so great that it is a reasonable to expect that a detailed revision of this group in the current sense will Make it possible to find some groups of close relatives with expressed hiatus, for which some clear diagnostic characteristics can be found to propose new generic taxa or restore some names treated here as synonyms. In particular, the Australian members of Zygadenia (Z. Martinae and Z. westraliensis) have peculiar cells of the elytra and seem to be enough separated from Eurasian ones [160] to propose for them a new palaeoendemic Australian genus.
The junior homonym Curculionites Giebel, 1856 [140], non Heer, 1847 [141] was proposed for the group with four species proposed together with the genus (syrichtus, westwoodi, tuberculatus and marginatus) and, Curculionites tuberculatus Giebel, 1856, which has enough the more or less distinct characteristics for a reliable taxonomic interpretation, was chosen by Handlirsch as the type species of Zygadenia. The same was Made for the generic name Procarabus including three species names proposed together with the generic one (reticulatus, tripartitus and zitteli), which were put in Omma (zitteli) and Notocupes (reticulatus, tripartitus) [129].
The generic name Forticupes is based on the description of a badly preserved specimen (Forticupes laiyangensis) which was put in the subfamily Ommatinae, although the characteristics Making such supposition really true are not clearly expressed (nevertheless the elytral apex looks ommatine rather than cupedine). The name Conexicoxa was proposed for the description of a species (Conexicoxa homora) interpreted as a member of Trachypachidae and represented only by the underside of the metathorax with posterior legs and abdomen. It was transferred at first to Zygadenia [110] and later automatically to the restored Notocupes [96], however the imprint of the holotype of Conexicoxa homora can scarcely be considered as Notocupes or Zygadenia. The synonymization of these names is also here recognized as proposed by Ponomarenko et al. [96].
The genus Chengdecupes with the type species Chengdecupes jurassicus was preliminarily synonymized with Notocupes by Kirejtshuk et al. [11] and later it was also assigned to Ommatinae [33]. To this genus after its proposal other three members were added by Hong: Chengdecupes baojiatunensis, C. kezuoense and C. shiluoense. The additional estimation of the characteristics of the above mentioned species Made it possible to specify some details visible in the photograph of the holotypes of all members of this genus and to support the previous synonymization of this name with Zygadenia by Kirejtshuk et al. [11], although four species described as members of Chengdecupes could belong to different generic groups.
[Tetraphalerus] has the structure of pronotum and elytra certainly characteristic of Notocupes but not Tetraphalerus. Both species described as "Notocupes" undatabdominus Lin, 1980 [161] and "N." (?) multituberatus Lin, 1980 [161] (China: Zhejiang (Laozhu, Lishui); Lower Cretaceous, Lower Aptian, −125.5-122.5 Ma) have no characterictics Making it possible to assign them to archostematans and, therefore, these two species are regarded without subordinal, family and generic attribution (incertae sedis). Both species (?) Zygadenia semen and (?) Z. sibirica were described after study of the underside sclerites of pterothorax, abdomen and elytral epipleura as members of Zygadenia. However, in the original descriptions of both these species a very narrow epipleuron is pointed out, but there is no mention any diagnostic character. Therefore, the attribution of these species is also used here with question Mark. In other cases the question Mark was put if any described member of Zygadenia (=Notocupes) has no characteristics for a reliable generic diagnostics.
Nevertheless, the large taxon Zygadenia in sense here accepted, on the one hand, is rather variable and, on the other hand, includes some members, which seem to be enough different, to consider them as a member of another genus. In particular, the drawing of the distal part of the elytron designated as the holotype of Zygadenia tuberculata (type species of the genus) [1] shows the obscure fusion A1 and CuA and strongly approached to the elytral apex. The description of the type species of Notocupes (N. picturalis) [103] was described with mention on the elytral venation in the text and with unclear picture and drawing of the holotype. Besides, Ponomarenko described Zygadenia sinitzae [73] with completely different elytral venation (according to the drawing), and some congeners were described without information on elytral venation at all. Finally, Ponomarenko gave a general scheme of the elytral venation of the genus Zygadenia [110] which is rather different different from that in known congeners. Thus, the Many species of this taxon need to be revised to clarify their attribution. Diagnosis. Body elongate; head long and usually subparallelsided, eyes located at middle level, temples very long; prothorax rather narrow with sides explanate and distinctly carinate; veins more or less expressed (including A2), subparallelly terminated on Sc along lateral elytral edge; cells variable but usually expressed; explanate elytral sides moderately to rather wide and with diffuse microtubercles or small punctures; abdominal ventrites co-planar (abutting).

Notes.
Usually this genus is associated with Tetraphalerus, although the species of this genus in the modern fauna are distributed in South America (Tetraphalerus bruchi Heller, 1913 [162] and T. wagneri [104] and at the moment it is scarcely possible to explain their phyletic links with Asian Mesozoic species. Moreover, the modern species of the mentioned group sharing some external similarity with some extinct Mesozoic ones differ from the latter groups in the lateral elytral sides not being explanate or very narrowly explanate, prothorax without sharp lateral carinae and scutellum widened apically (although the prothorax of the modern representatives slightly dorsoventrally compressed). The two mentioned recent species of Tetraphalerus have different elytral venation and it Makes it possible to suppose that their phylogenetic separation happened rather long ago. It is here supposed that a considerable similarity of the mentioned modern species and Mesozoic ones are mostly convergent and the Asian Mesozoic species of Allophalerus gen. nov. have not close relationship with the modern representatives of Tetraphalerus.
Comparison. This new genus is somewhat similar to other genera including slender ommatines with comparatively narrow prothorax, namely: Omma, Pareuryomma, Polyakius gen. nov. and Rhabdocupes; however the unique combination of the diagnostic characteristics can Make easy recognition of the new genus which was frequently described as "Tetraphalerus". Besides, the elytra and general appearance of congeners of this genus are similar to those in Monticupes, but the elytral veins of species of Allophalerus gen. nov. are separately ended to Sc, while A1 and CuA of congeners of Monticupes are fused in distal part. In addition to this diagnostic feature, the head of both groups is different (longer and subparallelsided in Allophalerus gen. nov. and shorter and subtriangular in Monticupes). Comparison. This new genus is somewhat similar to other genera including slender ommatines with comparatively narrow prothorax, namely: Omma, Pareuryomma, Polyakius gen. nov. and Rhabdocupes; however the unique combination of the diagnostic characteristics can make easy recognition of the new genus which was frequently described as "Tetraphalerus". Besides, the elytra and general appearance of congeners of this genus are similar to those in Monticupes, but the elytral veins of species of Allophalerus gen. nov. are separately ended to Sc, while A1 and CuA of congeners of Monticupes are fused in distal part. In addition to this diagnostic feature, the head of both groups is different (longer and subparallelsided in Allophalerus gen. nov. and shorter and subtriangular in Monticupes).

Etymology. The name of this genus is formed from the Greek
Etymology. The name of this genus is formed from the Greek ἄ λλος (another) and stem of the generic name "Tetraphalerus". Gender masculine. Diagnosis. Body wide; head subquadrate and subparallelsided or subtriangular and moderately long, eyes located at base, temples very short; prothorax subspherical with incarinate sides; veins scarcely expressed, longitudinal rows of cells not quite regular and subparallelly terminated on Sc along lateral elytral edge; cells variable but more or less expressed; explanate elytral sides rather wide and with very large cells arranged in more or less expressed 2-3 longitudinal rows (two complete rows and one short row between them at elytral base), abdominal ventrites co-planar (abutting).

Notes.
A certain similarity of the elytral structure of the first described species of this new genus and that of the elytron described by Hong [71] as "Brochocoleus punctatus" was a reason to join them in one genus but the elytra of these species have only a sketchy similarity, while the difference in other organs between two groups here considered as two genera, Brochocoleus and Burmocoleus gen. nov. is rather great (see the comparison below).
Comparison. This new genus has the elytra similar to those in Brochocoleus, Jarzembowskiops gen. nov. and Stegocoleus (with widely explanate elytral sides bearing very large cells); however Burmocoleus gen. nov. differs from them in the subglobular (subspherical) prothorax without lateral carinae (not explanate and incarinate at sides), and also from: -Brochocoleus in the very large and almost irregular cells and lack of trace of veins on elytral disk, very narrow "neck" and very short temples and subquadrate scutellum; -Jarzembowskiops gen. nov. in the irregular cells and lack of trace of veins on elytral disk, seriate large cells on explanate elytral sides; -Stegocoleus in the irregular cells and lack of trace of veins on the elytral disc (not indistinctly striate and not with small cells between striae), seriate large cells on explanate elytral sides, and subquadrate scutellum.
The prothorax of the new genus subglobular and uncarinate, reminiscent of that in species of Bukhkaliusgen. nov., Cionocoleus, Cionocups gen. nov., Omma, Polyakius gen. nov. and Rhopalomma whose elytra, however, are completely different from that in Burmocoleus gen. nov. (see above notes to the mentioned genera and the diagnosis of Polyakius gen. nov.). Diagnosis. Body wide; head subquadrate and subparallelsided or subtriangular and moderately long, eyes located at base, temples very short; prothorax subspherical with incarinate sides; veins scarcely expressed, longitudinal rows of cells not quite regular and subparallelly terminated on Sc along lateral elytral edge; cells variable but more or less expressed; explanate elytral sides rather wide and with very large cells arranged in more or less expressed 2-3 longitudinal rows (two complete rows and one short row between them at elytral base), abdominal ventrites co-planar (abutting).

Notes.
A certain similarity of the elytral structure of the first described species of this new genus and that of the elytron described by Hong [71] as "Brochocoleus punctatus" was a reason to join them in one genus but the elytra of these species have only a sketchy similarity, while the difference in other organs between two groups here considered as two genera, Brochocoleus and Burmocoleus gen. nov. is rather great (see the comparison below).
Comparison. This new genus has the elytra similar to those in Brochocoleus, Jarzembowskiops gen. nov. and Stegocoleus (with widely explanate elytral sides bearing very large cells); however Burmocoleus gen. nov. differs from them in the subglobular (subspherical) prothorax without lateral carinae (not explanate and incarinate at sides), and also from: The prothorax of the new genus subglobular and uncarinate, reminiscent of that in species of Bukhkalius gen. nov., Cionocoleus, Cionocups gen. nov., Omma, Polyakius gen. nov. and Rhopalomma whose elytra, however, are completely different from that in Burmocoleus gen. nov. (see above notes to the mentioned genera and the diagnosis of Polyakius gen. nov.).
Etymology. The name of this new genus is formed from the ancient name of the country (Burma) and stem "coleus" (κoλεóς). Gender masculine.
Burmocoleus prisnyi Kirejtshuk, sp. nov. (Figures 5 and 6) urn: lsid:zoobank.org:act:164F4C0B-0F43-4DF3-90BA-89E17C7AD276 Holotype. ZIN Cuped 002. The specimen with removed posterior parts of elytra and abdomen, also with a missing part of the right mesotarsus, and part of the metatibiae and metatarsi is included in a flattened piece of amber (rectilinearly cut from one side (26 mm) and with suboval outline from another side with diameter 15 mm). Also, the part of the underside of the prothoracic and head integument is also missing. The beetle is located along the oval edge of the amber piece with the broken posterior part near to the oval edge of the amber piece and beyond. Because of optical reason the sculpture of the integument is poorly visible, particularly from above. The piece of amber with the beetle also includes spores and pollen grains, Many pieces of litter and organic Matter different in shape and size, and also the chitinized fragments of insect bodies, including one separate Mandible. The amber piece has also some cracks from the surface plane, from where the upper side of the beetle body accessible for observation. Diagnosis. This new species is distinct from Burmocoleus zhiyuani comb. nov. in the clearly projecting part of the anterior edge of the elytra at the middle (but not the shoulder angle as in its congener), and also in the much narrower (subtriangular) frons, other proportions in the antennomeres, widely rounded anterior edge of the pronotum (like that in species of Omma) and only two long rows of large cells on the explanate elytral sides (without a short row of cells between them).
Notes. The name Burmocoleus zhiyuani comb. nov. was proposed for the holotype which can be compared with the holotype of the new species. Another specimen was used for the description of this species as a paratype. However this paratype is Markedly slenderer than the holotype and has the structure of the head, antennae, pronotum, elytral outline and cells rather different from those in the holotype. The latter specimen certainly is a third congener of this new genus. Nevertheless, the head and pronotum of this paratype is somewhat similar to those in Burmocoleus prisnyi sp. nov., but not in B. zhiyuani comb. nov.
Description (holotype). Length without missing posterior part of body 14.8, width 6.2 mm. Body slightly to moderately convex dorsally and subflattened ventrally, dark brownish; sclerites of dorsal surface and thoracic underside with very dense and uniform small microtubercles and with long and very fine hairs, forming ciliation along elytral sides (visible at high Magnification).
Head subtriangular and rather narrowed apically, eyes located at base of head near distinctly isolated "neck", temples very short. Mandibles rather stout and rather developed and tridentate at apex. Antennae very thin, slightly shorter than head and prothorax together; their scape moderately long and thick (much thicker than tibial apex), antennomere 2 (pedicel) small (slightly longer than thick), antennomere 3 longest (about four time as long as antennomere 2 and twice as long as antennomere 4), antennomeres 4-10 subtriangularly widened apically and gradually becoming shorter, ultimate antennomere lanceolate and about as long as antennomere 7. Prothorax subglobular, wide basal orifice and narrow apical one from above, anterior and lateral outline forming contiguous contour with subuniform serration. Scutellum subquadrangular, slightly longer than wide and with slightly convex posterior edge. Elytra moderately convex along body and steeply sloping at widely explanate sides; lateral edge densely serrate, anterior edge with projection at middle; elytral disk apparently with almost irregular large cells (but Markedly smaller than those on explanate sides) and without trace of veins, but explanate sides with two longitudinal rows of particularly large cells; anterior and lateral edges densely serrate.
Thoracic underside sclerites with somewhat obliterated microtubercles, and also with coarse and rather dense punctures having diameter about 3-5 times as great as eye facets. Procoxae and metacoxae subcontiguous, mesocoxae very narrowly separated. Metaventrite slightly longer than mesoventrite, medially depressed, with discrimen and parametacoxal sutures. Metacoxae with mesal parts strongly projecting posteriorly. Abdomen with abutting junction of ventrites, ventrite 1 slightly extending behind apices of mesal part of metacoxae and about 1.5 times as long as each of ventrites 2-4 and ventrite 5 considerably longest (even its basal part without missing apical one).
Legs moderately long and with comparatively narrow tibiae and tarsi. Pro-and metafemora thickened at middle (nearly three times as thick as scape) and rather thin in distal third, mesofemur of usual shape, about twice as thick as scape. All tibiae very thin and gradually thickening apically, and well developed spur. Protibia with long and thin process at outer apical angle. Meso-and metatarsi with tarsomeres 1-4 slightly subconical and gradually shortening distally, tarsomere 5 longest. Protarsus about 3/4 as long as protibia, protarsomere 1 slightly longer than each of protarsomeres 2-4, protarsomere 5 about as long as protarsomeres 1-4 together. Claws simple and thin, very long. Diagnosis. Body elongate oval; head about as wide as long, subtriangular and strongly narrowed at base (neck), eyes very large and located at base, temples very short; prothorax subspherical with incarinate sides; dorsal integument with diffuse, dense and comparatively coarse microtubercles; elytra with microtubercles intermixed by small oval cells comparable in size with microtubercles; explanate elytral sides moderately and with densely serrate along edges, abdominal ventrites apparently co-planar (abutting).
Comparison. This new genus is rather similar to Cionocoleus, but, in contrast to the latter, has the peculiar shape of the head with very large eyes and rather narrow neck, much longer antennae, dorsal integument with dense and high coarse microtubercles, on the elytra microtubercles are intermixed with small oval cells apparently, and elytral sides with distinctly serrate lateral edges Composition. Type species (Cionocups Manukyani sp. nov.) only.
Etymology. The name of this new genus is formed from the name of the closest taxon "Cionocoleus" and stem of the generic name "Cupes". Gender masculine.
Cionocups manukyani Kirejtshuk, sp. nov. (Figures 7 and 8) urn:lsid:zoobank.org:act:014722E5-D1DE-4A55-95A2-D1D7FA8A90A7 Holotype. GPIH 4984, coll. C. Gröhn 11,186. The complete beetle is included in a flat amber piece of elliptic shape with the greater diameter 20.0 mm and smaller one 8.0 mm and height 3.3 mm. The beetle is obliquely located on one side of this piece. Besides, this amber piece also has Many transparent and Mat gas bubbles (most of them below the beetle), some cracks in different directions and size (some of them passing through the beetle), and also some spores and pieces of organic Matter of different size.
Notes. Some dorsoventral compression of the prothorax of this holotype examined could be at least partly lifetime, but mostly appeared as a sequence of postmortem deformation (it is clearly visible in on somewhat depressed pronotal base and irregularly subflattened elytra). The current careful observation showed that the prothoracic sides of this specimen are certainly incarinate, but the serration along the lateral edges of the pronotum looks like that along lateral the elytral sides rather than microtuberculation on the upper body sclerites. Description (holotype). Length 7.3, width 2.7, height 0.8 mm. Body subflattened dorsally (partly postmortem) and subflattened ventrally, dark chestnut brownish; sclerites of dorsal surface and thoracic underside with dense subuniform microtubercles and with dense rather conspicuous hairs, forming also short ciliation along elytral sides.
Head transversely subtriangular and rather narrowed apically, eyes strongly projecting and located at base of head near moderately expressed "neck", temples very short; transversely elevated at level of eyes and depressed before anterior edge, with slight covering over antennal insertion. Mandibles moderately long and gently curved to bidentate apices. Antennae very thin, somewhat longer than head and prothorax together; their scape moderately long and thick (much thicker than tibial apex), antennomere 2 (pedicel) moderately small (about twice longer than thick), antennomere 3 longest (about 2.5 times as long as each of antennomeres 2 and 4), antennomeres 4-10 subconical, ultimate antennomere lanceolate and about one and one quarter as long as each of antennomeres 4-10. Pronotum subflattened (partly as postmortem event) and with anterior edge forming line continuing arcuate lateral outlines, sides arcuately widening anteriorly, visible lateral contour with subuniform crenellation, base nearly straight. Scutellum apparently subquadrangular and with slightly convex posterior edge. Elytra moderately convex along body and apparently gently sloping to narrowly explanate sides; about 1.5 times as long as wide combined; lateral edge densely and sharply serrate, anterior edge almost straight; elytral disc apparently with diffuse microtubercles (without remnants of cells) and without veins, explanate sides with finer punctures; anterior and lateral edges densely serrate.
Genal processes of head rather wide, far projecting and exposed over level of mouthparts. Thoracic underside sclerites with somewhat obliterated microtubercles and also with coarse and moderately dense punctures having diameter somewhat greater than eye facets. Procoxae and metacoxae subcontiguous, mesocoxae very narrowly separated. Abdomen with abutting ventrites, ventrite 5 (hypopygidium) about 1.5 times as long as each of ventrites 2-4 and apparently only slightly longer than ventrite 1.
All femora and tibiae of more or les usual shape, but very long and extremely narrow, every femur seemingly comparable in length with corresponding tibia. Tarsi very thin and very long, ultimate tarsomere longer than combined length of antennomeres 2-4; claws simple and thin, moderately long.
Etymology. The epithet of this new species is dedicated to Andranik Rafaelievich Manukyan for friendly professional communication and cooperation during Many years, particularly for his help in preparation of Many inclusions of fossil beetles.
Clessidromma zengi Kirejtshuk, sp. nov. (Figures 9 and 10) urn:lsid:zoobank.org:act:2A59B74F-A5C4-415C-8C2D-40B169F2039D Holotype. ZIN Cuped 003. The complete specimen is included in the amber piece with a shape of parallepiped (with facets about 10.0 × 6.0 × 2.1 mm) having one facet (upper surface of the beetle) not completely flattened. This amber piece also includes some gas bubbles and some pieces of organic Matter and litter with different size and shape. Diagnosis. This new species in the general outlines of its body sclerites has a somehow intermediate position between two other congeners (Clessidromma palmeri and C. tianae comb. nov.) demonstrating also the Mandibles being longer than that in both these species, peculiar shape of the pronotum, peculiar proportions of the antennomeres, smaller scales on the dorsal surface and along the pronotal and elytral sides. Clessidromma zengi sp. nov. differs from the slenderer member of the genus (Clessidromma palmeri) in the shorter head, "neck"-shaped pronotal base and elytra, and, on the other hand, differs from the more robust species (C. tianae comb. nov.) in the longer head, pronotum and elytra.
Description (holotype). Length 6.2, width 1.9, height 0.9 mm. Body slightly to moderately convex dorsally and subflattened ventrally, dark brownish; sclerites of dorsal surface and thoracic underside with very dense uniform and small microtubercles, and also with thick and very conspicuous hairs of different colouration (yellowish, brownish and blackish), intermixed by thinner hairs; dense and conspicuous pubescence Masking integument and forming also ciliation along pronotal and elytral sides; dorsal vestiture on head, prothorax, mesoventrite and partly on base of elytra and profemora yellowish to reddish, while that on other parts Markedly darker.
Head subtriangular and slightly narrowed apically, eyes strongly projecting and located at base of head near moderately expressed "neck", temples very short; upper surface rather transversely elevated at level of eyes and with elevated tubercle over place of each antennal insertion. Mandibles very long and sharply curved at blunt apices. Antennae very thin, somewhat longer than head and prothorax together; their scape moderately long and thick (much thicker than tibial apex), antennomere 2 (pedicel) moderately small (about twice longer than thick), antennomere 3 longest (nearly three times as long as each of antennomeres 2 and 4), antennomeres 4-10 gradually becoming thicker, ultimate antennomere lanceolate and almost twice as long as each of antennomeres 4-10. Pronotum subflattened and with short "neck"-shaped base, anterior edge rather convex and explanate at middle and with slight median emargination of explanate portion, sides widening anteriorly, rounded anterior angles and very stump posterior ones, contour with subuniform serration. Scutellum rather widened apically, slightly shorter than wide and with widely rounded posterior edge. Elytra moderately convex along body and gently sloping to moderately explanate sides; almost twice as long as wide combined and broadly arcuate at sides; anterior edge almost straight; elytral disk apparently with regular medium-sized cells and scarcely expressed veins, Masked by dense vestiture, explanate sides with one longitudinal row of medium-sized cells; anterior and lateral edges densely and finely serrate.
Genal processes of head rather wide, far projecting and exposed over level of mouthparts. Labrum moderately exposed and with dense setae along transverse anterior edge. Mentum transverse, suboval to subpentagonal. Ultimate Maxillary palpomere subconical and slightly narrowing apically, about 2.5 times as long as thick. Thoracic underside sclerites with somewhat smoothed microtubercles and also with coarse and rather dense punctures having diameter somewhat greater than eye facets. Procoxae and metacoxae subcontiguous, mesocoxae very narrowly separated. Metaventrite slightly longer than mesoventrite, medially depressed and apparently with parametacoxal sutures. Mesal processes of metacoxae very moderately extending posteriorly. Abdomen with abutting ventrites, ventrite 5 (hypopygidium) about 1.5 times as long as each of ventrites 2-4 and slightly shorter than ventrite 1.
Anterior legs very long and significantly longer than posterior ones, intermediate legs somewhat shorter than anterior ones and Markedly longer than posterior ones. Pro-and mesofemora long and narrow, rather thicker in proximal half and very narrow at apex, profemur about 0.8 and mesofemur about 0.6 as long as antenna; metafemur moderately thickened in proximal half and about half as long as antenna. All tibiae narrow and comparable in length with corresponding femora, gradually thickening to apex, and with small spur. Protibia with moderately prominent and acute outer apical angle. Pro-and mesotarsi slightly longer and metatarsus Marked longer than corresponding tibiae; tarsomere 1 Markedly longer than each of tarsomeres 2-4 and tarsomere 5 longer than tarsomeres 2-4 together; claws simple and thin, moderately long.
Etymology. The epithet of the new species is dedicated to Zeng Xiantai (Zhengzhou, Henan, China) for friendly relation in cooperation with the author, particularly for his help in receipt of inclusions of fossil beetles for the collection of ZIN.
Diagnosis. Body wide oval; head long and subparallelsided, eyes located at base, temples very short; prothorax dorsoventrally compressed and widely subexplanate at carinate sides; elytra with shoulder angle slightly projecting anteriorly, veins very weakly expressed, longitudinal rows of rather small regular cells between veins and subparallelly terminated on Sc along lateral elytral edge; explanate elytral sides rather wide and with very large irregular cells, abdominal ventrites co-planar (abutting).
Comparison. This new genus has the charactertistic very widely explanate elytral sides with large cells, more or less similar to those in Brochocoleus, Burmocoleus gen. nov., Polyakius gen. nov. and Stegocoleus, however Jarzembowskiops gen. nov. differs from all mentioned genera in the very widely subexplanate pronotum and strongly projecting eyes, and also from: -Brochocoleus in the finer cells and more expressed trace of veins on the elytral disc, diffuse very large cells on explanate elytral sides, very narrow "neck" and very short temples and subquadrate scutellum; -Burmocoleus gen. nov. in the regular cells and exposed trace of veins on the elytral disc, diffuse large cells on explanate elytral sides; -Polyakius gen. nov. in the more exposed trace of veins on the elytral disc, diffuse large cells on more widely explanate elytral sides, very narrow "neck", very short temples and subquadrate scutellum; -Stegocoleus in the long head (not subtriangular) with very short temples, narrower explanate part of elytra, diffuse large cells on explanate elytral sides, and subquadrate scutellum.
Etymology. The name of this generic taxon is dedicated to Edmund Aleksander Jarzembowski, who Made Many discoveries of fossil archostematans and other fossil insects. Gender feminine.
Omma janetae Kirejtshuk, sp. nov. (Figures 13 and 14) urn:lsid:zoobank.org:act:C60194E1-D91B-4764-856C-2A39CC1ACF34 Holotype. ZIN Cuped 004. The complete specimen is included into the amber piece with shape of parallepiped (with facets about 12.7 × 7.3 × 2.8 mm) having one facet (at side of the underside of the beetle) not flattened and spherically curved. This amber piece also includes the remnants of leaves (one of them on the right part of the beetle body), Many small gas bubbles and some small pieces of organic Matter of different size and shape. This amber piece has some cracks of different size, obliquely oriented to the beetle body and one crack along the lower plane of the beetle. The species regarded as questional members of Omma and Marked in the above list with Mark (?), including also O. brevipes are not mentioned in this comparison (see also the above Notes to the genus Omma).
Description (holotype). Length 8.6, width 2.6, height 1.1 mm. Body moderately convex dorsally and ventrally, dark brownish; sclerites of dorsal surface and thoracic underside with very dense and subuniform microtubercles, and also with hairs of different thickness, colouration and conspicuousness, Masking integument (particularly on elytra) and forming also ciliation along elytral sides; underside vestiture darker, with less raised and less conspicuous pubescence.
Head subtriangular and rather narrow apically, eyes strongly projecting and located near well expressed "neck", temples extremely short; upper surface rather transversely elevated at level of eyes and this elevation extending to level above place of antennal insertion, and slightly depressed frons. Labrum well exposed and far projecting anteriorly, its anterior edge transverse. Mandibles very long and arcuately curved at blunt apices. Antennae moderately thin, somewhat shorter than head and prothorax together; their scape moderately long and thick (somewhat thicker than tibial apex), antennomere 2 (pedicel) smallest and nearly globular, antennomere 3 longest (nearly three times as long as antennomere 2 and about 2.5 times as long as antennomere 4), antennomeres 4-10 subequal and subconical, ultimate antennomere lanceolate and almost twice as long as each of antennomeres 4-10. Pronotum nearly globular and incarinate, anterior outline rather convex and forming one continuous arc with lateral sides; posterior edge subrectilinear and posterior angles stumpy. Scutellum subtriangular, about as long as wide, with arcuate sides and rounded at apex. Elytra moderately convex along body and steeply sloping to narrowly explanate sides; somewhat more than 1.5 times as long as wide combined and very broadly arcuate at sides; anterior edge gently rounded; elytral disk apparently with almost regular sculpture (probably small suboval cells) and slightly expressed veins, Masking by dense vestiture, narrow explanate sides with diffuse small microtubercles and small punctures; anterior and lateral edges finely and densely serrate.
Genal processes comparatively short and rather wide. Ultimate Maxillary palpomere slightly thickening apically, about twice as long as thick. Thoracic underside sclerites with somewhat obliterated microtubercles and also with coarse and rather dense punctures having diameter Markedly greater than eye facets. Procoxae and metacoxae subcontiguous, mesocoxae very narrowly separated. Metaventrite slightly longer than mesoventrite, medially depressed. Mesal processes of metacoxae very slightly extending posteriorly. Abdomen with abutting ventrites, ventrite 5 (hypopygidium) about 1.5 times as long as each of ventrites 2-4 and only slightly longer than ventrite 1.
Anterior and intermediate legs rather long and significantly longer than posterior ones, intermediate legs somewhat shorter than anterior ones and Markedly longer than posterior ones. Proand mesofemora long and narrow, profemur about 0.8 and mesofemur about 0.7 as long as antenna; moderately thick at base and moderately thin at apex. Metafemur rather thickened in proximal half and about half as long as antenna. All tibiae narrow and somewhat shorter than corresponding femora, with apical spur; protibia with rather prominent outer apical angle. Protarsus much shorter than protibia; protarsomeres 1-4 subcylindrical, moderately thick and with gradual shortening, protarsomere 5 about as long as protarsomeres 1 and 2 together. Meso-and metatarsi significantly longer than corresponding tibiae (metatarsus about 1.3 times as long as metatibia), meso-and metatarsomeres 1-4 rather narrow. Claws simple and thin, moderately long.
Notes. The sculpture and vestiture of this new species and the species of Omma, and also proportions of sclerites of different legs of Omma janetae sp. nov. are somewhat reminiscent of those in species of Clessidromma. These peculiarities could reflect some relationship of these ommatines.
Etymology. The epithet of this new species is dedicated to the wife (Janet) of Albert Allen who donated the holotypes of two new ommatine species described in this paper. Composition. Type species and Polyakius pubescens sp. nov.

Genus
Diagnosis. Body comparatively wide; head moderately long and subtriangular, eyes located at base, temples extremely short; prothorax slightly dorsoventrally compressed but incarinate at sides; elytra with shoulder angle slightly projecting anteriorly, veins apparently very weakly expressed, longitudinal rows of moderately large cells regular between veins and subparallelly terminated on Sc along lateral elytral edge; explanate elytral sides moderately wide and with large regular cells arranged in one row, abdominal ventrites co-planar (abutting).
Comparison. This new genus is similar and closely related to the genus Omma, differing from it mostly in some level of dorsoventral compression of the body, and also the presence of a regular row of large cells along the lateral elytral edge and very dense and comparatively long dorsal vestiture consisting of very thin hairs. Polyakius gen. nov. differs from Burmocoleus gen. nov., Cionocoleus, Cionocups gen. nov. and Bukhkalius gen. nov. in the regular cells on the elytral disc and only with one row of large cells along the lateral edge of the elytron; differs also from Rhopalomma in the short head with distinct neck and very short temples, independent ending of A1 and Cu (without fusion of A1 and Cu at elytral apex) and with one row of large cells along lateral edge of elytron (but not with diffuse microtubercles).
Etymology. The name of this new genus is devoted to my class Mate, Vladimir Aronovich Polyak, with whom the author was in close and friendly relationship for more than than 50 years. Holotype. ZIN-Cuped-01. The elliptic flat amber piece (25.0 × 16.0 mm) with height of 3.6 mm includes the beetle with more or less good preservation of the upper surface and rather bad one of its underside. The beetle specimen is almost complete but with missing left antenna, preapical right antennomeres, distal parts of profemora, protibiae and protarsi. It looks somewhat dorsally compressed because some deformation is observable on the pronotum and probably of the elytral base, a part of the upper surface and most of the beetle underside has a "milky" cover and it is difficult to observe structural details. This amber piece has some cracks in different directions, including the largest one in the plane of the elytral lateral edges of the beetle, and contains Many small pieces of organic Matter different in colour, size and shape, and also Many small gas bubbles. Diagnosis. This new species differs from another congener (P. pubescens sp. nov.) in the smaller and slenderer body, shape of the pronotum and steeply sloping elytral sides, comparatively longer head, proportions of the basal antennomeres, shorter pubescence on the elytra, somewhat larger cells on disk and explanate sides of elytra.

Polyakius alberti
Description (holotype). Length 14.6, width 4.9, height 1.1 mm. Body slightly convex to subflattened dorsally and subflattened ventrally, dark brownish; sclerites of head and pronotum densely microtuberculate, prosternum distinctly and densely punctured; body integument with dense hairs Masking sculpture and colouration, upper surface with much longer and more conspicuous hairs and with Many small gas bubbles between them forming along elytral lateral edges distinct ciliation, underside with shorter hairs.
Head subtriangular, about 0.7 times as long as pronotum, slightly longer than wide and rather narrowed apically, eyes rather prominent and located at base of head, "neck" well expressed and moderately narrow, temples very short; upper surface rather transversely elevated at level of eye base and with tubercle above each antennal insertion. Mandibles moderately raised and sharply curved at apparently sharp apices. Antennae moderately long and moderately thin, somewhat shorter than head and prothorax together, their scape of usual proportion and thickness (poorly visible but probably thicker than tibial apex), antennomere 2 (pedicel) very small (about twice longer than thick), antennomere 3 longest (nearly five times as long as antennomere 2 and about 2.5 times as long as antennomere 4), antennomeres 4-7 (antennomeres 8-11 missing) subequal and slightly thickened apically. Prothorax apparently initially subglobular; pronotum about as wide as long, subflattened (in life more or less convex but postmortem deformed) and with uniformly arcuate anterior edge, subflattened at posterior angles. Scutellum subquadrate to subpentagonal, widest at apex, apparently somewhat longer than wide. Elytra subflattened above body and rather steeply sloping to widely explanate sides; slightly more than twice as long as wide combined and broadly arcuate at sides; anterior edge gently arcuate; elytral disc apparently with regular medium-sized cells and probably veins (not visible because of dense vestiture and extremely small gas bubbles between hairs), explanate sides with one longitudinal rows of large cells; anterior and lateral edges densely serrate.
Procoxae and metacoxae subcontiguous, mesocoxae extremely narrowly separated. Intermediate legs Markedly longer than posterior ones. Mesofemur thickened in proximal half, about 1.3 times as long as metafemur and about 1.3 times as long as head. Meso-and metatibiae comparable in length with corresponding femora. Mesotarsus slightly longer than mesotibia, mesotarsomere 1 about as long as mesotarsomere 5 and about as long as mesotarsomeres 2-4 together. Metatarsus about 1.5 times as long as metatibia, metatarsomere 5 about as long as metatarsomeres 3 and 4. Claws simple and thin, moderately long.
Etymology. The epithet of this new species is dedicated to Albert Allen who donated the holotypes of two new ommatine species described in this paper.
Polyakius pubescens Kirejtshuk, sp. nov. (Figure 17) urn:lsid:zoobank.org:act:4CFAA645-24DA-4C05-929C-FBC9D93FC000 Holotype. ALDC0450/ALD.Bu.195 for deposition in ZMUC. Suboval to subtriangular flat unclear amber piece with one rectiliner facet (24.0 × 17.0 mm) with height of 3.6 mm includes the beetle with more or less good preservation of upper surface and rather bad preservation of its underside, which is covered with Mat and not transparent amber layer. The beetle specimen seems to be almost complete but only the upper integument can be observable, although Many small gas bubbles over the abdomen and a considerable part of the right elytron Make observation of these structures very complicated. The specimen could be somewhat postmortem compressed. Description (holotype). Length 21.0, width 7.7 mm. Body slightly convex dorsally and apparently subflattened ventrally, dark brownish; sclerites of head and pronotum densely microtuberculate, upper integument with dense straw reddish hairs Masking sculpture and colouration, upper surface with much longer and more conspicuous hairs and with Many small gass bubbles between them forming along elytral lateral edges distinct ciliation, underside with shorter hairs.
Head subtriangular, about 0.6 times as long as pronotum, slightly shorter than long and rather narrowed apically, eyes rather prominent and located at base of head near moderately narrow and well expressed "neck", temples almost absent; upper surface rather transversely elevated at level of eyes base. Antennae apparently moderately long and moderately thin, apparently much shorter than head and prothorax together, antennomere 3 longest (could be 1.5 times as long as antennomere 4), antennomeres 4-7 (antennomeres 8-11 missing) subequal and scarcely thickened apically. Prothorax apparently initially subglobular; pronotum transverse (more than 1.5 times as wide as long), somewhat subflattened (intravitally more or less convex but postmortally deformed) and with uniformly arcuate anterior edge, not subflattened at posterior angles. Scutellum, widest at rounded apex, apparently somewhat wider than long. Elytra subflattened above body and rather steeply sloping to widely explanate sides; slightly more than 1.5 times as long as wide combined and very broadly arcuate at sides; anterior edge gently arcuate; elytral disc apparently with regular medium-sized cells and veins and other interspaces between longitudinal rows of cells comparably raised), explanate sides with one longitudinal row of large cells; anterior and lateral edges densely serrate.

Notes on the Subfamily Ommatinae
The genus Fuscicupes Hong and Wang 1990 [145] (type species: Fuscicupes parvus Hong et Wang, 1990 [145]; China: Shandong Province (Tuanwang Village, Laiyang City); Lower Cretaceous, Aptian/Albian, -125.5-112.6 Ma) was proposed after the description of one specimen with very poor preservation and lack of characteristics for supposition on a connecting it with any suborder or family, although a general body outline of the print is partly reminiscent of elateroids.

Subfamily Triadocupedinae Ponomarenko, 1966
Type species:       over antennal insertion, and with large and rather projecting temples. Antennae apparently slightly extending behind middle of elytra, rather thick (scape and flagellomeres with comparatively small difference in thickness); scape longest and thickest, apparently at least three times as long as pedicel (antennomere 2) and more or less longer (apparently 1.5 to 2.0 times) than each of flagellomeres gradually becoming shorter apically. Pronotum slightly less than twice as wide as long, subpentangular and with gently and weakly convex posterior edge, moderately to rather arcuately convex anterior edge; subrectilinearly widened lateral edges between distinct stump posterior angles and also distinct, slightly acute and slightly projecting anterior angles. Elytra one and three-fourths as long as combined width, slightly to moderately arcuately curved along lateral edge, moderately and conjointly acuminate at apices, with well expressed primary veins, large and apparently subquadrate cells; epipleura comparatively narrow (comparable with those in modern species). Pygidium very widely rounded to transverse at apex. Profemora comparatively short and apparently rather thick, slightly longer than pronotum and with apices moderately extended beyond lateral edges of pronotum.
Etymology.The epithet of this new species is dedicated to Sergey Il'ich Golovatch, outstanding expert in myriapods and friend of the author during Many years. Diagnosis. This new species is characterized by the comparatively wide body, very wide and comparatively short prothorax with arcuate lateral edges. These features distinguish Cupes legalovi sp. nov. from other species known from all comparatively contemporary European outcrops in Eckfeld and Messel, and also from other congeners.

Cupes legalovi
Description (holotype) ( Figure 20C,D). Length 11.7 mm (better measureable in counterpart). Elongate and apparently rather convex dorsally. Head transverse, about twice as wide as long, subtriangular and rather narrowed anteriorly, with moderately large eyes, and with large and rather projecting temples somewhat extending posteriorly. Antennae with visible two first antennomeres, scape twice longer than thick; also twice as long as and about 1.5 times as thick as pedicel (antennomere 2). Prothorax about 2.5 times as wide as long; prosternum with shallowly bi-emarginate anterior edge and with moderately narrow prosternal process; pronotum with regularly and weakly convex posterior edge, arcuately widened lateral edges between distinct stump posterior angles and also distinct, slightly acute and slightly projecting anterior angles. Meso-and metathoracic sclerites, including metacoxae, very similar to those in modern species, although metaventrite Markedly shorter. Elytra with very narrow epipleura and cells rather large and somewhat transverse. Abdomen with the overlapping junctions of ventrites and clear transverse roller at base of ventrites 2-5, hypopygidium longer than each of other ventrites and widely rounded at apex. Legs moderately long and rather thick, profemur much longer than prothorax, protibiae widened apically.
Etymology. The epithet of this new species is dedicated to Andrey Alexandrovich Legalov, close colleague of the author, with whom he published Many papers on fossil beetles. Diagnosis.This new species is characterized by the comparatively wide body, comparatively short prothorax and one longitudinal row of large cells along lateral edge of elytra. Cupes lutzi sp. nov. is very similar to C. eckfeldensis, but distinct from it in the outlines of the prothoracic sides.

Cupes lutzi
Description (holotype) ( Figure 21). Length 10.6 mm. Elongate and apparently moderately convex dorsally. Head transverse, about twice as wide as long, subtriangular and rather narrowed anteriorly, apparently with comparatively large eyes, large tubercle over antennal insertion, and with large and rather projecting temples. Antennae apparently slightly extending behind distal third of elytra, moderately thick to thin (scape and flagellomeres with rather great difference in thickness); scape longest and thickest, more than three times as long as pedicel (antennomere 2) and more or less longer (about 1.5 times) than each of flagellomeres subequal in length. Pronotum about twice as wide as long, subpentangular to subhexangular, rather convex at posterior edge with more prominent its median part (forming tri-sinuate outline: median emargination and and two paramedian sinuations), rather arcuately convex anterior edge (also forming somewhat bi-sinuate outline); very slightly subrectilinearly widened lateral edges between distinct stump posterior angles and also distinct, nearly right and not projecting anterior angles. Scutellum somewhat transverse, subpentagonal with rounded apex. Elytra about 2.4 times as long as combined width, slightly arcuately curved along lateral edge, broken apices, with weakly expressed primary veins, large and apparently subquadrate to somewhat transversely quadrangular cells; somewhat (sub) explanate lateral sides with one row of large cells, epipleura comparatively narrow (comparable with those in modern species). Pro-and mesofemora moderately long and apparently rather thick, slightly longer than pronotum and with apices moderately extended beyond lateral edges of pronotum; protibia somewhat widened to apex.
Etymology. The epithet of this new species is dedicated to Herbert Lutz, curator of fossil beetles in LNNR.
Paratype. Represented by one print MeI11220-01 (SFNFM). The remains of the specimen show the almost complete outline dorsal integument with destroyed some places of the elytra in the middle and along their lateral edges; besides, the apices of the profemora and bases of protibiae are partly exposed beyond the pronotal sides, and also a fragment of the left intermediate leg is visible. Diagnosis. This new species is characterized by the comparatively large eyes, subparallelsided pronotum with regularly rounded anterior and posterior edges, and also elytra much wider than pronotum. The mentioned characterictics Makes it possible to recognize this new species among the species known from comparatively contemporary European outcrops in Eckfeld and Messel.
Description (holotype) (Figure 22A,B). Length 5.3 mm. Elongate oval and apparently moderately convex dorsally. Head transverse, about 1.5 times as wide as long, subtriangular and moderately narrowed anteriorly, apparently with rather large eyes, and with moderately raised temples. Antennae apparently slightly extending behind middle of elytra, moderately thick (scape and flagellomeres with small difference in thickness); scape longest and thickest, about 2.5 times as long as pedicel (antennomere 2) and more or less longer (about 1.5 times) than each of flagellomeres subequal in length. Pronotum about twice as wide as long, subquadrangular, slightly and regularly convex at posterior edge and rather arcuately convex anterior edge; subrectilinearly parallelsided atsides, with distinct stump anterior and posterior angles. Scutellum very large and transverse, subtriangular with widely rounded apex. Elytra about twice as long as combined width, slightly arcuately curved along lateral edge, apices subangular separated, with well expressed primary veins, large and apparently subquadrangular to subpolygonal cells; somewhat (sub) explanate lateral sides with dense and diffuse microtubercles, epipleura apparently rather wide. Exposed part of pygidium showing widely rounded apex. Profemora moderately long and apparently rather thick, Markedly longer than pronotum and with apices moderately extended beyond lateral edges of pronotum; protibia somewhat widened to apex.
Paratype. Length 8.5 mm. Visible outlines of dorsal surface similar to those in holotype, but head about twice as wide as long and with traceable paired tubercles: one pair over antennal insertions and one pair behind them; pronotum with median carina; sculpture of elytra less clear than in holotype.
Etymology. The epithet of this new species is dedicated to Maxim Vital'evich Nabozhenko, close colleague of the author, with whom he published Many papers on fossil beetles.
Cupes wedmannae Kirejtshuk, sp. nov. (Figure 22C,D) urn:lsid:zoobank.org:act:B1716D2E-86E1-43A0-9D12-6566DF045F99 Holotype. Represented by the part MeI4386A (SFNFM) and counterpart MeI4386B (SFNFM). The part shows the dorsal integument of the head with the left scape and right antennomeres 1-4, pronotum, most part of elytral discs and exposed apex of left profemur and base of left protibia; sculpture of elytral integument is more or less observable. The counterpart has the observable the same organs, but with rather obscure outlines more destructed antennae, and, alternatively, with some traces of the elytral parts (including the outlines of elytral apices) absent in the holotype part and very weak and obscure traces of some thoracic sclerites and legs.
Notes. This new species is assigned to the genus Cupes despite the missing characterictics of the sculpture of the elytral apices because the good clarity of the primary veins and the presence of the large and regularly seriate cells in elytra (not diffuse and small cells or punctures). All other important characterictics of both Cupes and Miocupes are rather similar. Diagnosis. This new species is characterized by the very peculiar shape of comparatively long pronotum with the almost straight posterior edge of pronotum and also the anterior edge rather far projecting anteriorly. This feature very distinguishes this species among all the congeners. Besides, it distinct among the contemporary European congeners from Eckfeld and Messel in the comparatively long head with small eyes and strongly projecting temples.
Description (holotype) ( Figure 22C-D). Length of part 9.8, length of counterpart 5.1 mm. Elongate and apparently moderately convex dorsally. Head transverse, about one and one-seventh as wide as long, subtriangular and very narrowed anteriorly, with comparatively small eyes, two pair of tubercles (over antennal insertions and along eyes) and with strongly raised temples, represented by only four right antennomeres, with moderately thick (scape and flagellomeres with small difference in thickness); scape longest and thickest, about 2.5 times as long as pedicel (antennomere 2) and more or less longer (about1.5 times) than each of flagellomeres 3 and 4 subequal in length. Pronotum about 1.5 times as wide as long, subpentangular; subrectilinear to shallowly subemarginate at posterior edge and strongly arcuately convex anterior edge (strongly projecting anteriorly); subrectilinearly widened from distinct posterior angles to distinct anterior angles. Scutellum not visible. Elytra about 2.5 times as long as combined width, very slightly arcuately curved to rectilinear along lateral edge, apices subangular separated, with well expressed veins, large and apparently subquadrangular to subpolygonal cells; apparently somewhat (sub) explanate lateral sides with one row of large cells Etymology. The epithet of this new species is dedicated to Sonja Wedmann, curator of fossil beetles in SFNFM. Holotype. GPIH 4985, coll. Gröhn 11192. The beetle specimen with missing right eye and abdomen and desegmented in the pro-mesothoracic junction is included into the flat amber piece with an suboval shape which is rectilinearly cut at one longer diameter (16.0 × 13.0 mm) and with height 1.3 mm. This amber piece some different amber layers, and it is also included some small pieces of organic Matter and litter with different size and shape, and apparently some fungal hyphae. Diagnosis. This new species is similar to another congener (Mallecupes qingqingae) and differs from the latter at least in the large and rather prominent temples, and shape of pronotum with the sharp and comparatively long lateral processes. Besides, Mallecupes prokini sp. nov. has the extremely steeply sloping elytral sides forming angle between flat plane of the disc of the elytron and sloping plane almost or even less than 90%, apparently Markedly lesser than that in Mallecupes qingqingae.

Mallecupes prokini
Notes. The type series of Mallecupes qingqingae consists of some specimens which seem to belong to at least two species. The pictured holotype NIGP 157008 and drawn paratype NIGP 164791 have the same characteristics different from those in Mallecupes prokini sp. nov. (see above), however each of these specimens looks enough different to regard them as separate species.
Description (holotype) (Figures 23 and 24). Length 6.3, width 1.3, height 0.7 mm. Body rather convex dorsally and subflattened ventrally, very dark brown; sclerites of dorsal surface and thoracic underside with very dense uniform and extremely small microtubercles, and also with thin and slightly conspicuous hairs, on dorsal integument apparently intermixed by narrow scales. Head subtriangular, strongly transverse and rather narrowed apically, eyes rather large and located between rather prominent temples and very short frons; "neck" distint and short, nearly one third as thick as head wide; temples very short; upper surface apparently rather smoothed. Mandibles comparatively short and sharply curved at acuminate apices. Antennae very thin, somewhat shorter than body length; its scape moderately long and comparatively thin (almost as thin as femora); antennomere 2 (pedicel) rather small (about three times as thick as long and one third as long as each of antennomeres 1 and 2); other antennomeres subcylindrical, very long and very thin; antennomere 3 apparently longest and next antennomeres gradually becoming shorter, ultimate antennomere gently narrowing at acute apex. Pronotum subflattened, anterior part from very sharp processes (anterior angles) gently narrowing to transverse anterior edge, sides widening anteriorly from subrectilinear posterior edge, posterior angles stump, contour not even but not serrate. Scutellum somewhat widened apically, slightly shorter than wide and with widely rounded posterior edge. Elytra very flat at discs and very steeply (subvertically) sloping to sides (very narrow epipleura), about 2.5 times as long as wide combined and subparallelsided in three-fourths of length; elytral disk and sloping sides with regular comparatively large cells and well subparallel and expressed veins ending on Sc; anterior and lateral edges smooth (not serrate).
Genal processes of head very short and wide. Thoracic underside sclerites of usual shape and proportions. Procoxae distinctly separated by narrow process somewhat widened at transverse apex. Mesocoxae about as narrowly separated as procoxae. Metacoxae subcontiguous. Metaventrite about as long as mesoventrite, medially depressed and apparently with parametacoxal sutures. Mesal processes of metacoxae very far extending posteriorly, rather wide at base and sharply narrowing posteriorly, forming deep and comparatively widely triangular interspace between them.
Anterior legs very long and Markedly longer than intermediate and posterior ones. All femora long and narrow, of usual subelliptic shape, thickest along midlength. All tibiae extremely narrow and comparable in length with corresponding femora, subparallelsided, and with small spur. Pro-and mesotarsi slightly longer and metatarsus nearly 1.5 times as long as corresponding tibiae; protarsomere 1 Markedly longer (almost twice) than each of tarsomere 2-4 and tarsomere 5 longer than tarsomeres 2-4 together. Meso-and metatarsomeres 1 slightly longer than each of corresponding meso-and metatarsomeres 2-4. Claws simple and thin, moderately long.
Etymology. The epithet of this new species is dedicated to Alexander Alexandrovich Prokin, colleague of the author, with whom he published Many papers on fossil beetles.

Conflicts of Interest:
The author declares no conflict of interest.

Appendix A
After preparation of this Manuscript the descriptions of three species appeared and all of them belong to a new genus not included in this review. An appendix is therefore necessary. Also, one Tetraphalerus species, the description which was published in 2017, was missed by the author and is added to this review. E.A. Jarzembowski kindly provided detailed discussion on the Manuscript and great assistance in improving the text contributing very significant ideas and facts, including some data on the following two genera (Echonocups Kirejtshuk et Jarzembowski, gen. nov. and Lobanovia Kirejtshuk et Jarzembowski, gen. nov.). Therefore, the co-authorship of E.A. Jarzembowski is appropriate. Composition. Echinocups neli comb. nov., and also Echinocups ohmkuhnlei (Jarzembowski, Wang et Zheng, 2020) [184], comb. nov. [Notocupes] and Echinocups denticollis (Jiang, Li, Song, Shi, Liu, Chen et Kong, 2020) [185], comb. nov. [Notocupes].
Diagnosis. Body elongate oval; head more or less transverse, subtriangular and weakly narrowed at base (neck), eyes medium-size to large and located at base, temples very short and retracted into prothoracic segment; prothorax subflattened, with widely explanate and carinate sides, its lateral edges strongly dentate, procoxae subcontiguous; dorsal integument with diffuse, dense and comparatively coarse microtubercles; elytra with CuA and M fused and curved towards 1A subapically, cells large and arranged in longitudinal rows, longitudinal interspaces between these rows of cells bearing rather high tubercles (sharp spines), explanate stripe at least with one row of large cells or with additional rows of smaller cells and lateral edge densely dentate; abdominal ventrites 1-3 overlapping and others co-planar (abutting).
Notes. All three members of Echonocups gen. nov. were described as congeners of Notocupes because of some general body similarity. Nevertheless these three species are very different from all Notocupes sensu lato in the characteristics mentioned in the comparison bellow. Although the drawing of Echinocups denticollis comb. nov. shows the venation as should be the case in true congeners of Notocupes, it is here treated as erroneous, as all other characteristics of this species completely correspond with those in other species of Echonocups gen. nov.
The important feature of Echonocups gen. nov. is a discovery of the exposed aedeagus from the genital capsule in the holotype of Echinocups denticollis comb. nov. (Jiang et al., 2020: Figure 4) which is very reminiscent of that in Priacma and Gracilicupes [33] in the subfamily Cupedinae (but not Ommatinae). Other fossil cupedids showing exposed aedeagus fit with the traditional classification of the family Cupedidae [26,33], etc. Nevertheless, the attribution of Echinocups denticollis comb. nov. to the ommatines at the moment is based on the subcontiguous procoxal cavities.
Another feature of this new genus is its overlapping abdominal ventrites 1-3. The type of junction of abdominal ventrites is usually one of the group feature characterizing closely related species and very frequently used as diagnostic for supraspecific taxa. In contrast to the congeners of the new genus, the cupedine Priacma and Gracilicupes have co-planar ventrites.
Comparison. This new genus is characterized by the general body outline somewhat similar to that in Many representatives of Notocupes, but, unlike the latter, the three species considered as congeners of Echinocups gen. nov. have strongly dentate lateral edges of the prothorax, sharp spines on elytra, different elytral venation and overlapping abdominal ventrites. The co-planar abdominal ventrites are characteristic of most ommatines, including relatives of Notocupes (Notocupedini sensu Ponomarenko, 1969), although not infrequently the posterior edge of the abdominal ventrites in different ommatines somewhat elevated over the base of the following ventrites. The true overlapping (tegular) junction of abdominal ventrites is more characteristic of cupedines than ommatines, but the cupedine representatives with the Priacma-type aedeagus (Priacma and Gracilicupes), as mentioned above, have the co-planar abdominal ventrites. Thus, assignement of Echinocups gen. nov. to ommatines can be regarded as preliminary and could be revised after obtaining more data.
Etymology. The name of this new genus is formed from the Greek "Echinus" (spiny animal) and stem of the generic name "Cupes". Gender masculine. Composition. Only type species (Bukhkalius lindae comb. nov.) Diagnosis. Body elongate; head somewhat elongate, subtriangular and strongly narrowed at base (neck) and anteriorly, eyes large and located at base, temples very short; prothorax subglobular with neck-like anterior projection and incarinate, procoxae subcontiguous; dorsal integument with diffuse, dense and comparatively coarse microtubercles; elytra with indistinct and not fused veins, cells somewhat elongate and arranged in irregular longitudinal rows, explanate lateral stripe apparently with two rows of cells and lateral edge densely dentate; abdominal ventrites co-planar (abutting).

Genus
Comparison. This new genus is characterized by the general body outlines somewhat similar to that in Many representatives of Tetraphalerus and Allophalerus gen. nov., but Bukhkalius gen. nov. has the head rather narrowing anteriorly, narrow neck and very large eyes, prothorax with narrow "neck", and densely serrate elytra sides. Besides, this new genus differs from the genera with subglobous prothorax (Cionocoleus, Cionocups gen. nov., Omma, Polyakius gen. nov. and Rhopalomma) in the clear "neck"of the head, and also from Cionocoleus and Cionocups gen. nov. also in the longitudinal rows of cells on elytra.
Etymology. The name of this new genus is dedicated to entomologist Sergey Pеtrovich Bukhkalo, who graduated from the Kharkov State University together with the author of this paper and who passed away in the summer of 2019. Gender masculine.