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Habitus photographs of cucujoid (A–H), cleroid (I) and coccinelloid (J–L) taxa. Taxonomy follows proposed classification introduced in the text. (A) Paracucujus rostratus (Boganiidae). (B) Megauchenia sp. (Nitidulidae). (C) Psammoecus sp. (Silvanidae). (D) Pharaxonotha sp. (Erotylidae). (E) Ericmodes sylvaticus (Protocucujidae). (F) Cryptophagus sp. (Cryptophagidae). (G) Anthonaeus agavensis (Kateretidae). (H) Hobartius sp. (Hobartiidae) (I) Diplocoelus sp. (Biphyllidae). (J) Teredolaemus sp. (Teredidae stat.n.). (K) Bystus sp. (Anamorphidae stat.n.). (L) Bicava sp. (Latridiidae). Scale bars = 1 mm. Photos by JAR.

Habitus photographs of cucujoid (A–H), cleroid (I) and coccinelloid (J–L) taxa. Taxonomy follows proposed classification introduced in the text. (A) Paracucujus rostratus (Boganiidae). (B) Megauchenia sp. (Nitidulidae). (C) Psammoecus sp. (Silvanidae). (D) Pharaxonotha sp. (Erotylidae). (E) Ericmodes sylvaticus (Protocucujidae). (F) Cryptophagus sp. (Cryptophagidae). (G) Anthonaeus agavensis (Kateretidae). (H) Hobartius sp. (Hobartiidae) (I) Diplocoelus sp. (Biphyllidae). (J) Teredolaemus sp. (Teredidae stat.n.). (K) Bystus sp. (Anamorphidae stat.n.). (L) Bicava sp. (Latridiidae). Scale bars = 1 mm. Photos by JAR.

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A large-scale phylogenetic study is presented for Cucujoidea (Coleoptera), a diverse superfamily of beetles that historically has been taxonomically difficult. This study is the most comprehensive analysis of cucujoid taxa to date, with DNA sequence data sampled from eight genes (four nuclear, four mitochondrial) for 384 coleopteran taxa, including...

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... superfamilies, Cucujoidea is the most problematic with regard to classification and no synapomorphies supporting its monophyly have been identified (Leschen et al., 2005;Leschen & ´ Slipi´nskiSlipi´nski, 2010). Cucujoidea is a heterogeneous group of beetles which have a similar appearance (e.g. small, drab colouration, clubbed antennae) ( Fig. 1) that could not be placed satisfactorily elsewhere. The group was established for convenience and represents the largest taxonomic dumping ground among the superfamilies of Coleoptera. Cleroidea in particular shares many characters with certain groups of Cucujoidea such that these two superfamilies are difficult to separate (Crowson, ...

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The Neotropical tribe Dorynotini is characterized by a conspicuous tubercle or spine adorning the elytra, which, along with a few other characters, has been used to differentiate its recognized five genera and two subgenera. However, relationships among these taxa and the evolutionary origin of the pronounced tubercle remain speculative. Here we pr...
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The Neotropical tribe Dorynotini is characterized by a conspicuous tubercle or spine adorning the elytra, which, along with a few other characters, has been used to differentiate its recognized five genera and two subgenera. However, relationships among these taxa and the evolutionary origin of the pronounced tubercle remain speculative. Here we pr...

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... The classification of beetle families is given according to the publications [40,41]. At the same time, we have taken into account changes in names from the Catalogue of Palaearctic Coleoptera [42][43][44][45][46][47][48], as well as for Cucujoidea from the publication of Robertson et al. [49], for Curculionoidea-from the publication of Alonso-Zarazaga et al. [50]. To clarify the nomenclature, the above publications were used, as well as the Catalogue of Palaearctic Coleoptera [51,52]. ...
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Monitoring Coleoptera diversity in protected areas is part of the global ecological monitoring of the state of ecosystems. The purpose of this research is to describe the biodiversity of Coleoptera studied with the help of baits based on fermented substrate in the European part of Russia (Smolny National Park). The research was conducted April–August 2018–2022. Samples were collected in traps of our own design. Beer or wine with the addition of sugar, honey, or jam was used for bait. A total of 194 traps were installed. The dataset contains 1254 occurrences. A total of 9226 Coleoptera specimens have been studied. The dataset contains information about 134 species from 24 Coleoptera families. The largest number of species that have been found in traps belongs to the family Cerambycidae (30 species), Nitidulidae (14 species), Elateridae (12 species), and Curculionidae and Coccinellidae (10 species each). The number of individuals in the traps of these families was distributed as follows: Cerambycidae—1018 specimens; Nitidulidae—5359; Staphylinidae—241; Elateridae—33; Curculionidae—148; and Coccinellidae—19. The 10 dominant species accounted for 90.7% of all detected specimens in the traps. The maximum species diversity and abundance of Coleoptera was obtained in 2021. With the installation of the largest number of traps in 2022 and more diverse biotopes (64 traps), a smaller number of species was caught compared to 2021. New populations of such species have been found from rare Coleoptera: Calosoma sycophanta, Elater ferrugineus, Osmoderma barnabita, Protaetia speciosissima, and Protaetia fieberi.
... Historically, it is essentially a group of families without clear diagnostic characteristics of other superfamilies (especially Tenebrionoidea; Crowson, 1955;Lawrence and Newton, 1982). The coccinelloid group, once regarded as the cerylonid series, was recognized based on multiple lines of morphological (Crowson, 1955;Ślipiński and Pakaluk, 1991) and molecular evidence (Hunt et al., 2007;Robertson et al., 2008Robertson et al., , 2015Bocak et al., 2014), and formally removed from Cucujoidea and elevated to its superfamilial status by Robertson et al. (2015). The phylogenetic relationships within the remaining Cucujoidea vary dramatically among various morphological and molecular studies (e.g., Leschen et al., 2005;Robertson et al., 2008Robertson et al., , 2015Lawrence et al., 2011;McElrath et al., 2015;Timmermans et al., 2016;Zhang et al., 2018;McKenna et al., 2019). ...
... Historically, it is essentially a group of families without clear diagnostic characteristics of other superfamilies (especially Tenebrionoidea; Crowson, 1955;Lawrence and Newton, 1982). The coccinelloid group, once regarded as the cerylonid series, was recognized based on multiple lines of morphological (Crowson, 1955;Ślipiński and Pakaluk, 1991) and molecular evidence (Hunt et al., 2007;Robertson et al., 2008Robertson et al., , 2015Bocak et al., 2014), and formally removed from Cucujoidea and elevated to its superfamilial status by Robertson et al. (2015). The phylogenetic relationships within the remaining Cucujoidea vary dramatically among various morphological and molecular studies (e.g., Leschen et al., 2005;Robertson et al., 2008Robertson et al., , 2015Lawrence et al., 2011;McElrath et al., 2015;Timmermans et al., 2016;Zhang et al., 2018;McKenna et al., 2019). ...
... The coccinelloid group, once regarded as the cerylonid series, was recognized based on multiple lines of morphological (Crowson, 1955;Ślipiński and Pakaluk, 1991) and molecular evidence (Hunt et al., 2007;Robertson et al., 2008Robertson et al., , 2015Bocak et al., 2014), and formally removed from Cucujoidea and elevated to its superfamilial status by Robertson et al. (2015). The phylogenetic relationships within the remaining Cucujoidea vary dramatically among various morphological and molecular studies (e.g., Leschen et al., 2005;Robertson et al., 2008Robertson et al., , 2015Lawrence et al., 2011;McElrath et al., 2015;Timmermans et al., 2016;Zhang et al., 2018;McKenna et al., 2019). Although some molecular analyses either based on a few gene markers or a larger dataset (95 nuclear protein-coding genes) under a site-homogeneous substitution model supported a monophyletic Cucujoidea sensu Robertson et al. (2015), recent studies using transcriptomic data (McKenna et al., 2019) or a better-fitting site-heterogeneous model (Cai et al., 2022) have consistently demonstrated the paraphyly of Cucujoidea sensu Robertson et al. (2015). ...
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An enigmatic cucujiform beetle, Alloterocucus atratus Li, Leschen, Liu, and Cai gen. et sp. nov. , is reported from mid-Cretaceous Burmese amber. The character combination of the new fossil is not completely consistent with any of the known cucujoid or erotyloid families. Based on our phylogenetic analyses, Alloterocucus is assigned to Cucujoidea and may be allied to Lamingtoniidae, which contains a single Australasian genus in the extant fauna. Alloterocucus shares with Lamingtoniidae a similar habitus and a series of characters, including the absence of postocular constriction, 3-segmented antennal club, externally open procoxal cavities, laterally open mesocoxal cavities, exposed pro- and mesotrochantins, and the absence of epipleural fovea and pronotal setose cavities, but differs from extant Lamingtoniidae in its apically truncate terminal maxillary palpomeres, 5-5-4 tarsi in male and absence of distinct dorsal punctation.
... The laterally expanded clypeus is also present in other unrelated groups of ladybird beetles such as Telsimiini or Chilocorini. However, various recent molecular studies (Seago et al. 2011, Robertson et al. 2015, Che et al. 2021 demonstrated that they are unrelated, and are positioned in different places of the Coccinellidae phylogenetic tree. The peculiar shape of the clypeus is probably related to the behaviour of the beetles, how they feed, and on what type of Sternorrhyncha they prey. ...
Article
A new remarkable ladybird beetle Platycrus laotanus gen. et sp. nov. is described from Laos. It is placed in the tribe Platynaspini, but it differs from the remaining members of the tribe by having unusually expanded legs, a peculiar pocket-like structure to accommodate the tarsi in repose, and the antennae consisting of 11 antennomeres. Detailed morphological description and illustrations are provided. Taxonomic placement of the newly described taxon is discussed, and a transfer of the genera Crypticolus Strohecker and Hornious Weise from Platynaspini to Coccinellinae as incerte sedis.
... The classification of the family-group taxa used predominantly follows Cai et al. [48] and McKenna et al. [49]. The lists of species were verified according to the Catalogue of Palaearctic Coleoptera [50][51][52][53][54][55][56][57][58], to Robertson et al. [59], and to Alonso-Zarazaga et al. [60]. The years of description of some beetle species are specified according to Bousquet [61]. ...
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(1) Background: Coleoptera is one of the most diverse insect lineages. Coleoptera species live in many ecosystems around the globe and their role in ecosystems is very diverse. To study the number and distribution of species, lists of species were compiled and then added into the database generation platforms. The aim of the work was to describe the modern fauna of Coleoptera based on a recently published dataset. (2) Methods: Studies were conducted from 1971 to 2022. Insects were collected by different means (searching under the bark of trees and stumps, sifting litter and wood dust, soil samples, caught in the light, in soil traps, window traps, etc.). For each observation, the coordinates of the place of discovery, the number of individuals, dates were noted. (3) Results: The dataset presents data on 2416 species and subspecies of Coleoptera from 89 families found in the Lipetsk region. In total, the number of studied specimens in the dataset was 16,184, the number of occurrences was 6192. The largest families in terms of species diversity were Staphylinidae (541 species), Curculionidae (416), Chrysomelidae (315) and Carabidae (285). (4) Conclusions: In addition, based on the analysis of additional references, 452 more species and 2 families are indicated. Thus, the biodiversity of Coleoptera of the Lipetsk region is 2868 species from 89 families.
... The superfamily Coccinelloidea was proposed by Robertson et al. (2015) for a group of 15 families whose members were formerly placed in Cucujoidea (or Clavicornia) as part of the Cerylonid series (Robertson et al. 2007). Older works on this group are cited in the following chapters of the DeGruyter Handbook of Zoology: Ślipiński et al. 2010b (Bothrideridae) Cline & Ślipiński, 2010 (Discolomatidae), Tomaszewska, 2010. ...
... Based on the cladogram in Robertson et al. (2015), coccinelloids form 12 clades: 1) Bothrideridae, with Derataphrus sister to the remaining genera and Bothrideres sister to a clade containing Acetoderes, Dastarcus and all other genera; 2) Murmidiidae and Discolomatidae, treated as families in the present work; 3) Teredidae with Oxylaemus and Anommatus sister to the remaining genera, Euxestidae with Metacerylon sister to the other genera and Cerylonidae with Ostomopsis sister to the remaining genera; 4) Latridiidae, with separate cortricariine and latridiine clades; 5) and 6) containing the wingless genera Acalyptoischion and Sphaerosoma, respectively; 7) eight genera of Anamorphidae; 8) Corylophidae, with the basal Periptyctinae and three main clades for Holopsis, Foadia + Priamima and the 12 remaining corylophine genera; 9) a clade with two main groups: the merophysiine complex (subfamilies Pleganophorinae, Leiestinae and Merophysiinae), and the endomychine complex with the remaining Endomychidae; 10) Mycetaeidae, 11) Eupsilobiidae, and 12) Coccinellidae, with subclades for Microweiseinae and Coccinellinae. Coccinelloid wings illustrated here represent the following families and subfamilies: Bothrideridae (8), Murmidiidae (1), Discolomatidae (1), Teredidae (4), Euxestidae (3), Cerylonidae: Ostomopsinae (1), Ceryloninae (2); Latridiidae (2), Anamorphidae (3), Corylophidae: Periptyctinae (1), Corylophinae (1); Endomychidae (3), Eupsilobiidae (1), Coccinellidae: Micro weiseinae (1), Coccinellinae (6). ...
... This last series was further divided into several groups: Hobartiidae + Boganiidae, Cryptophagidae, the Phloeostichid group (Agapythidae, Priasilphidae, Cavognathidae and Phloe ostichidae), Cucujidae + Silvanidae, Cyclaxyridae + Myraboliidae, and the Laemophloeid group (Passandridae, Phalacridae and Laemophloeidae (with Propalticidae as one of its four subgroups). In the phylogenetic study of Cai et al. (2022) Cucujoidea is divided into three superfamilies, Erotyloidea, Nitiduloidea and Cucujoidea, the families included in each often corresponding to those of Robertson et al. (2015), which will be used here as a template for discussing wing characters. ...
Article
Part 2 of this work includes a review of morphological and systematic work on Histeridae (G07, revision), Bostrichoidea (G15), Coccinelloidea (G16), Lymexyloidea + Tenebrionoidea (G17), Cleroidea (G18), Cucujoidea (G19), Chrysomeloidea (G20) and Curculionoidea (G21), discussions of hind wing structure in each group based on 702 wing images, references to additional published figures and comments on wing morphology and, if possible, how these wing features may or may not be correlated with recent phylogenetic hypotheses. The introduction is followed by brief discussions of some important works not mentioned in Part 1, particularly those dealing with relationships of extinct taxa.
... The following references were used for the taxonomic placement and comparison with recent and fossil taxa: Sasaji (1995), Tomaszewska (2000aTomaszewska ( , 2000bTomaszewska ( , 2015, Shockley and Alekseev (2014), Robertson et al. (2015), Alekseev and Tomaszewska (2018) and Esser (2019). Specimens of extant Leiestes seminiger (Gyllenhal, 1808) collected by the first author in the Kaliningrad region ( Figure 1) were also used for morphological comparison. ...
... According to Tomaszewska (2000a) and Robertson et al. (2015), the fossil specimen under consideration is placed in Endomychidae based on the following combination of characters: head with distinct fronto-clypeal suture, subantennal groove absent, antennae clubbed, pronotum with basal and paired lateral sulci, procoxae subglobular, mesocoxal cavities open, elytral epipleura well developed, abdominal ventrite 1 as long as two following ventrites combined, abdominal ventrite 1 without postcoxal lines and tarsi 4-segmented. ...
... The increasing number of described fossil endomychid taxa (Motschulsky 1856; Kirejtshuk and Nel 2009;Shockley and Alekseev 2014;Alekseev and Tomaszewska 2018;Tomaszewska et al. 2018;Reike et al. 2020;Li et al. 2022Li et al. , 2022 needs to be catalogued and kept up-to-date for goals of possible use in further studies. A brief updated list of the described extinct species of handsome fungus beetles, Endomychidae sensu Robertson et al. (2015), from fossil resins (excluded copals) is compiled and presented below. The present described endomychid diversity of the Eocene epoch consists of 10 extinct species belonging to eight genera (5 extinct, marked with †; and three extant) from 4 subfamilies. ...
Article
A new extinct handsome fungus beetle species of the extant Palaearctic genus Leiestes Chevrolat, L. tomaszewskae Alekseev and Bukejs sp. nov., is described from the Baltic amber. The new species is studied and illustrated in detail using X-ray micro-computed tomography (μCT). A list of described Endomychidae known from fossil resins (Baltic, Bitterfeld, Oise and Burmese ambers) is compiled.
... Historically, Lymexylidae had once been associated with Cleroidea or Cucujoidea [10,11], but molecular studies have recovered an affinity with Tenebrionoidea. In three of the studies, Lymexylidae appeared to be nested within basal Tenebrionoidea, with various positions [12][13][14], although only a few gene fragments were sampled in these studies. Other studies, including three recent phylogenomic ones, suggested Lymexylidae as the sister group of (the remaining) Tenebrionoidea [15][16][17][18][19]. Wheeler [20] concluded that the maxillary palporgan is the strongest autapomorphy for this family, and considered its loss in Australymexylon Wheeler as secondary. ...
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The fossils once assigned to Raractocetus Kurosawa from the Mesozoic and Cenozoic amber deposits differ from extant Raractocetus in the longer elytra, the more strongly projecting metacoxae, and the hind wing with vein 2A forked. Thus, these fossils should be removed from Raractocetus. Cretoquadratus engeli Chen from Kachin amber appears to be conspecific with R. fossilis Yamamoto. As a result, R. fossilis and R. extinctus Yamamoto from Kachin amber, R. balticus Yamamoto from Baltic amber, and R. sverlilo Nazarenko, Perkovsky & Yamamoto from Rovno amber are transferred to Cretoquadratus Chen, as C. fossilis (Yamamoto) comb. nov., C. extinctus (Yamamoto) comb. nov., C. balticus (Yamamoto) comb. nov., and C. sverlilo (Nazarenko, Perkovsky & Yamamoto) comb. nov., and C. engeli syn. nov. is suggested to be a junior synonym of C. fossilis.
... Corylophidae, also known as the minute hooded beetles, is a moderately diverse and cosmopolitan family in the superfamily Coccinelloidea (Robertson et al. 2015), with about 285 extant species in 27 genera (Robertson et al. 2013). Corylophids generally have a minute body, and the ones with further miniaturization occur in several inde-pendent lineages (Robertson et al. 2013;Polilov 2016;Yavorskaya and Polilov 2016). ...
... The postcoxal lines on metaventrite and abdominal ventrite 1 are important diagnostic characters for Xenostanus. These postcoxal lines are usually present in Coccinellidae and some other related taxa (Ślipiński and Tomaszewska 2010;Robertson et al. 2015). However, most corylophids do not possess such lines (e.g., Furukawa 2010: fig. ...
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The family Corylophidae is a moderately diverse coccinelloid beetle family. The fossil record of corylophid beetles is extremely sparse, with only one species formally described from the Eocene Baltic amber. Here we report a new corylophid genus and species, Xenostanus jiangkuni Li, Szawaryn & Cai gen. et sp. nov. , from mid-Cretaceous amber from northern Myanmar (ca. 99 Ma). Xenostanus is most distinctly characterized by the antenna with 10 antennomeres and the presence of metaventral and abdominal postcoxal lines. Our phylogenetic analysis suggested Xenostanus as sister to tribe Stanini. Based on its distinctive morphology and the phylogenetic results, Xenostanus is placed in the tribe Xenostanini Li, Szawaryn & Cai trib. nov.
... Coleoptera families were classified according to Bouchard and co-authors [38], with additions [39]. We took into account changes from the Catalog of Palaearctic Coleoptera [40][41][42][43][44][45][46], publications of Robertson and co-authors [47] for Cucujoidea, and Alonso-Zarazaga and co-authors [48] for Curculionoidea. The nomenclature of beetles was standardized according to the publications cited above, with addition of the Catalog of Palaearctic Coleoptera [49,50]. ...
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Research on the Arthropoda distribution in temperate forest ecosystems has shown that communities are distributed vertically in unequal numbers. However, this issue still has research gaps in relation to the vertical stratification of Coleoptera in pine forests of the temperate zone. This study was conducted in the Republic of Mordovia situated in central part of European Russia. We used fermental traps to collect Coleoptera (the bait is fermenting beer with sugar and honey). The sampling was conducted from May to September 2021 in five sites of pine forests. One hundred and twenty-five species from 36 families were identified. The highest species richness was found in Nitidulidae and Cerambycidae (19 species each), Elateridae (13), Curculionidae (7) and Scarabaeidae (6). The highest number of species (84 species) was obtained at the height of 1.5 m, while the smallest species richness (44 species) was found at the height of 12.0 m. At the height of 12 m, we also registered the minimum number of specimens. Twenty-four species from 11 families were common to all the heights studied. Cryptarcha strigata and Glischrochilus grandis preferred heights of 3 and 1.5 m. Cryptarcha undata and Protaetia marmorata were more common at a height of 7 m with a frequency of 61.4–79.6% and 68.2–79.6%, respectively. The absolute number of saproxyl and anthophilic beetle species was higher in the ground layer and at the undergrowth level. These studies expand our understanding of the vertical distribution of beetles in pine forests of the temperate zone of European Russia.
... Traditionally, Endomychidae contained 12 subfamilies [1] and was classified in the superfamily Cucujoidea [2][3][4][5], in the derived group called 'Cerylonid Series' [6]. However, the most recent, comprehensive molecular research on Cucujoidea by Robertson et al. [7] has resulted in the formal recognition of the Cerylonid Series as an independent superfamily Coccinelloidea and the redefinition of Endomychidae by removing Anamorphidae, Mycetaeidae, and Eupsilobiidae as separate families. ...
... Endomychidae currently contains over 1600 described species classified in about 90 genera distributed in all zoogeographical realms, with the highest diversity in the tropical and subtropical regions of the world [1,3,4]. The study of Robertson et al. [7] recovered two main clades within the family, the 'merophysiine complex' and the 'endomychine complex'. The merophysiine complex includes the subfamilies Leiestinae, Merophysiinae, and Pleganophorinae, the basal lineages of the family according to Tomaszewska [4], while the endomychine complex includes Cyclotominae, Endomychinae, Epipocinae, and Lycoperdininae, and corresponds to 'Higher Endomychidae' sensu Tomaszewska [4]. ...
... The endomychine complex is supported morphologically by the pseudotrimerous tarsi in adults and the V-or U-shaped frontal arms on the head, and four pairs of stemmata in larvae [4]. The study of Robertson et al. [7], however, did not include exemplars of subfamilies Danascelinae and Xenomycetinae, this last one a sister group to 'Higher Endomychidae' in the analysis of Tomaszewska [4], so their relationships with the rest of the handsome fungus beetles remain unclear. Moreover, some current subfamilies, e.g., Endomychinae in the new sense, which now includes the genus Endomychus plus all genera of the former subfamily Stenotarsinae, are an anatomically heterogeneous group with no potential supporting synapomorphies. ...
Article
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A new genus and species of the family Endomychidae (Coleoptera: Coccinelloidea): Cretostenotarsus striatus Tomaszewska, Szawaryn and Arriaga-Varela gen. et sp. nov. are described, diagnosed and illustrated from the mid-Cretaceous amber from northern Myanmar. To test the systematic placement of the new extinct genus and species within the family, a phylogenetic analysis was conducted. A dataset of 38 morphological characters scored for 29 species (including the new fossil taxon), members of Endomychidae sensu stricto and representatives of Coccinelloidea as outgroups were analyzed using maximum parsimony. The results of the analysis indicate unequivocally that Cretostenotarsus striatus is a member of the Stenotarsus clade within a monophyletic ‘endomychine complex’ sensu Robertson et al. (2015), which corresponds to ‘Higher Endomychidae’ sensu Tomaszewska (2005). The present discovery confirms at least the Jurassic origin of Coccinelloidea and indicates a much older origin of Endomychidae than previously hypothesized.