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Maximum-likelihood tree (part 2 of 6). The full topology is shown to the left of the figure with the emboldened region enlarged and coloured for discussion. Branches are coloured red for Cucujoidea following the legend in the previous figures. Families are coloured as indicated to the right of the corresponding terminals. Nomenclatural changes proposed in this study are denoted to the far right of the tree with grey bars. Nodes supported by bootstrap support ≥ 90 are indicated by black circles, and nodes with support between 70 and 89 are indicated by grey circles.

Maximum-likelihood tree (part 2 of 6). The full topology is shown to the left of the figure with the emboldened region enlarged and coloured for discussion. Branches are coloured red for Cucujoidea following the legend in the previous figures. Families are coloured as indicated to the right of the corresponding terminals. Nomenclatural changes proposed in this study are denoted to the far right of the tree with grey bars. Nodes supported by bootstrap support ≥ 90 are indicated by black circles, and nodes with support between 70 and 89 are indicated by grey circles.

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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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... early diverging lineages of core Cucujoidea recov- ered in the present study include three major clades: (i) Monotomidae + nitidulid series, (ii) erotylid series (Helotidae-Protocucujidae-Protosphindus Sen Gupta & Crowson-Erotylidae), (iii) Boganiidae + Hobartiidae (see Fig. 5); all three lineages comprise taxa that have been con- sidered as early diverging cucujoids and plesiomorphic from a morphological perspective (Crowson, 1955(Crowson, , 1960(Crowson, , 1990). For example, Crowson (1955) postulated that Nitidulidae, Smicrip- idae and Monotomidae were closely related to Protocucujidae and Sphindidae ...
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... of the Nitidulidae group ( Leschen et al., 2005) -or nitidulid series -including Kateretidae, Smicripidae and Nitidulidae, formed a monophyletic group, albeit with only weak support (<50) (Fig. 5). Whereas morphology strongly supports this grouping ( Leschen et al., 2005;Cline, 2010;but see also Lawrence et al., 2011 for an exception), the present study is the first phylogenetic analysis based on molecular data to recover a monophyletic nitidulid series; previous studies did not include sufficient representation of the series, ...
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... was weakly supported (<50) as the sister taxon to the nitidulid series ( Fig. 5), another relationship formally recovered for the first time in the present study yet consis- tent with previous views (e.g. Crowson, 1955). Monotomidae are an enigmatic group with dubious phylogenetic affinity within Cucujoidea. Crowson (1955) noted the similarity of Monotomidae and the nitidulid group based on shared adult anatomical ...
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... sister grouping of Ericmodes Reitter (Protocucujidae) and Protosphindus (Sphindidae) is strongly supported in the present study (97) (Fig. 5). A close relationship of these taxa has long been recognized (Crowson, 1955;Thomas, 1984b) and is well supported by morphological data (McHugh, 1993;Leschen et al., 2005). Along with Ericmodes and Protosphin- dus, Helota Maclay has been considered to be a relatively early diverging cucujoid. Helotidae in particular is an enigmatic ...
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... & Crowson were far removed from each other. The present study is the first molecular phylogenetic analysis to include members of both Boganiidae and Hobartiidae. The recovery of these taxa forming a sister group (Fig. 5) is therefore intriguing, yet this relationship is only weakly supported (<50) and warrants further ...
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... recovered a strongly supported clade (95) comprising Cryptophagidae and the remaining core cucujoid families (Fig. 5). In terms of the families included, this clade is nearly consistent with the cucujid series of Hunt et al. (2007) andBocak et al. (2014), although the series is not consistent between those two studies: Hunt et al. (2007) does not include Silvanidae in the cucujid series, whereas Bocak et al. (2014) does not include Pas- sandridae. Our ...
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... represents the first molecular analysis to include exemplars of these enigmatic, species-poor families. We recovered a weakly supported clade comprising south temperate taxa including Agapytho Broun (Agapythidae), Priasilpha Broun (Priasilphidae), Taphropiestes Reitter (= Cavognatha Crowson) (Cavognathidae) and Hymaea Pascoe (Phloeostichidae) (Fig. 5). Within this clade Agapytho formed the sister group to Priasilpha obscura Broun (97); the remaining nodes were only weakly supported. The position of Taphropiestes in the present study was unstable. Although Taphropiestes was recovered as the sister group of Hymaea (<50), it often placed within the clade comprising Myrabolia Reitter ...
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... analyses consistently recovered a sister grouping of Cucujidae + Silvanidae ( Fig. 5) with moderate support (78). Both families were at one time classified together with Lae- mophloeidae and Passandridae in a broadly defined Cucujidae (Cucujidae s.l.). Crowson (1955) recognized at the familial level both Passandridae (with reservation) and Silvanidae, thereby removing them from Cucujidae s.l. The laemophloeids were ...
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... (Endomychidae). The spurious affiliation of Phalacridae, Hypo- dacnella and Bystus in Lawrence et al. (2011) is not consistent with previous views (Leschen et al., 2005;Hunt et al., 2007), nor is it consistent with the present study. In our analyses Cyclaxyra was consistently recovered as the sister group to Myrabolia Reitter (Myraboliidae) (Fig. 5) with weak support (50), a grouping that has never been suggested previously; this study is the first molecular analysis to include exemplars of either family. Myraboliidae represents another species-poor, monogeneric family with 13 species restricted to Australia. The inclusion of Myrabolia in multiple cucujoid families historically ...
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... Laemophloeidae, Propalti- cidae, Phalacridae and Passandridae form a natural lineage based on a number of morphological features including unequal protibial spurs, structural similarities of the male genitalia, and the presence of pronotal lines and elytral cells. Our results support this grouping (hereafter referred to as the laemophloeid group) (Fig. 5) as the above four families form a clade with moderately high branch support (89). Interestingly, previous molecular (Hunt et al., 2007;Bocak et al., 2014;McElrath et al., 2015) and morphological phylogenetic studies (Leschen et al., 2005;Lawrence et al., 2011) have not recovered the lae- mophloeid group as monophyletic; the present ...
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... closely related to Cyclaxyridae and Myraboliidae than to the remaining lae- mophloeid group taxa. Laemophloeidae and Propalticidae have been considered to be sister taxa (e.g. Lawrence & Newton, 1995;Leschen et al., 2005;Hunt et al., 2007;Lawrence et al., 2011). Interestingly, our results recovered Propalticus Sharp nested within Laemophloeidae (Fig. 5). These results concur with recent findings by Bocak et al. (2014) and McElrath et al. (2015), and support the proposal by McElrath et al. (2015) to subsume Propalticidae within ...
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... Cucujiformia and therefore not allied with any of the existing superfamilies of Cucujiformia, including the remaining Cucujoidea, was one of the most significant results of the present study. The support for the CS clade was high (97), as was the support for Cucujiformia (98) and the clade comprising the remaining cucujiform lineages (83) (Fig. 5). Although the exact placement of the CS in Hunt et al. (2007) and Bocak et al. (2014) is not concordant with our results, both studies independently demonstrated the isolated position of the CS clade relative to the remaining cucujiform lineages. Given our resulting topology and previously published results (Hunt et al., 2007;Bocak et ...

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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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... 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. ...
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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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... 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. ...
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... 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. ...
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... 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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... 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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... 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. ...
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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.