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Phloiophilus edwardsi: 15 – phallus, 16 – tegmen ventrally (transparent view), 17 – ovipositor ventrally, 18 – ovipositor dorsally, 19 – female internal reproductive organs (oviducts and ovaries removed, gut cut off).
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Phloiophilus edwardsi Stephens, 1830 (Coleoptera, Cleroidea) is redescribed and its character states inserted into a character matrix of ten families of Coleoptera and into a matrix of Trogossitidae genera. The matrices are analyzed by NONA. The resulting trees reveal relations of P. edwardsi to Cleroidea and to the tribes Thymalini and Colydiopelt...
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... To evaluate the systematic placement of the new genus, a morphological phylogenetic analysis was performed. The data matrix (Supplemental Material Datas S1-S3) was derived from a previously published dataset [30,31], and includes 91 morphological characters of both adults and larvae. Gracilenticrus gen. ...
A new genus and species of the cleroid family Lophocateridae are described and illustrated from the mid-Cretaceous amber of northern Myanmar. Gracilenticrus burmiticus Yu, Kolibáč & Ślipiński gen. et sp. nov. is unique among Lophocateridae in the tiny body size, frontoclypeal suture and antennal grooves absent, symmetrical antennal clubs, protrochantin reduced, tarsal claws small and widened at base. A key to the species of Mesozoic Lophocateridae is also provided. Morphological characters of the newly discovered Gracilenticrus were analyzed together with representatives of 43 extant genera of Cleroidea (broadly defined Trogossitidae) in a matrix of 91 characters. Gracilenticrus burmiticus was resolved as a member of Lophocateridae. The discovery of a diverse fauna of Lophocateridae in the mid-Cretaceous sheds a new light on the early evolution of superfamily Cleroidea.
... The former beetle family Trogossitidae (e.g., Barron, 1971;Kolibáč, 2006Kolibáč, , 2008Ślipiński, 1992) (Cleroidea) had been revealed to be paraphyletic in several molecular phylogenies (e.g., Bocak et al., 2014;Bocakova et al., 2016;McKenna et al., 2015McKenna et al., , 2019Zhang et al., 2018), and it has been recently divided into several families in the most comprehensive analysis by Gimmel et al. (2019). Under the traditional broad definition, the family comprises about 600 extant species in over 50 genera distributed worldwide (Kolibáč & Leschen, 2010;Kolibáč, 2013). ...
... The morphological character matrix was modified based on Kolibáč (2006Kolibáč ( , 2008, who conducted a phylogenetic analysis Parsimony analyses using implied weighting (K = 12) yielded a single most parsimonious tree ( Figure 5). The overall support values of key nodes of the tree were quite low, suggesting morphological information alone is not sufficient to solve the phylogeny of Trogossitidae. ...
... Traditionally, five tribes were included in the subfamily Trogossitinae (or Trogossitidae in the modern sense): Calityini, Egoliini, Larinotini, Gymnochilini, and Trogossitini (e.g., Kolibáč, Kolibáč (2006Kolibáč ( , 2008. However, in the molecular phylogeny of Gimmel et al. (2019), Calityini, Egoliini, and Larinotini did not form a clade together with Gymnochilini and Trogossitini, but rather occupied a position closer to Peltidae. ...
The first fossil representative of the cleroid family Trogossitidae is described from mid‐Cretaceous Burmese amber. Microtrogossita qizhihaoi Li & Cai gen. et sp. nov. is unique among Trogossitidae in the relatively widely separated procoxal and mesocoxal cavities, weakly asymmetrical antennal clubs, coarsely facetted eyes, coarse sculpture of dorsal and ventral surfaces of thorax in comparison with tiny body size, and the absence of spines along side margin of tibiae. Morphological characters of the fossil were analyzed together with representatives of 44 extant genera of Cleroidea (Peltidae, Lophocateridae, and Trogossitidae) in a matrix of 93 characters. Microtrogossita qizhihaoi was resolved as a member of Trogossitini within Trogossitidae. The tribal composition of Trogossitidae is discussed in light of our re‐analysis of a previously published four‐gene dataset under a site‐heterogeneous model. The recently described lophocaterid Mesolophocateres pengweii Yu, Leschen & Ślipiński syn. nov. from Burmese amber is suggested to be a junior synonym of Burmacateres longicoxa Kolibáč & Peris. The first fossil representative of the cleroid family Trogossitidae is described from mid‐Cretaceous Burmese amber. Microtrogossita qizhihaoi Li & Cai gen. et sp. nov. has a unique character combination among Trogossitidae. The fossil was resolved as a member of Trogossitini within Trogossitidae based on a morphological phylogenetic analysis. The tribal composition of Trogossitidae is discussed in light of our re‐analysis of a previously published four‐gene dataset under a site‐heterogeneous model.
... The new genus belongs to a group of lophocateride genera with distinct mola and deeply emarginate frontoclypeal suture ('Lophocaterini' sensu Kolibáč 2006Kolibáč , 2008Kolibáč , 2013. It shares divided prementum and structure of lacinia with Indopeltis Crowson, 1966, while five elytral carinae and distinct dorsal pubescence resemble Trichocateres Kolibáč, 2010. ...
... et sp. nov. is a member of Lophocaterini sensu Kolibáč (2006Kolibáč ( , 2008Kolibáč ( , 2013, however, the same author (Kolibáč 2013) had speculated about paraphyly of the tribe. A molecular analysis by Gimmel et al. (2019) showed Lophocaterini as polyphyletic, with the specimens g0085 and P1123 denoted as "Lophocateres cf. ...
... The tribal system of Lophocateridae should certainly be adapted and the first step should be to include the common pest Lophocateres pusillus Klug, 1833, the type genus and species of the group, in an analysis. The morphological analyses (Kolibáč 2006(Kolibáč , 2008 partially supported by molecular data (gimmel et al. 2019) preliminarily suggest existence of two groups (A, B; see below) in Lophocateridae that are unambiguously morphologically recognisable by the structure of the mandible and mostly also by the structure of the frontoclypeal suture (compare Kolibáč 2013: Fig. 26, Gimmel et al. 2019: Fig. 5). ...
Carinicateres merkli gen. et sp. nov. of Lophocateridae from eastern Thailand is described. The monotypic genus is distinct in its unique synapomorphies, namely the tarsal formula 4-4-4, elytron with 2 complete and 3 incomplete carinae, antenna 10-segmented with 2-segmented club. The emarginate frontoclypeal suture, mandible with mola, prementum divided into two parts, cranium dorsally with large impressions, each with seta in centre, crenulate lateral margins of pronotum, and distinct dorsal vestiture are shared with lophocateride genera Indopeltis Crowson and Trichocateres Kolibáč. Two specimens affiliated to Lophocateres Olliff and recently used in three molecular phylogenetic analyses are unambiguously identified as members of the clade Ancyrona + Neaspis and associated with Neaspis Pascoe.
... Two tribes, Lophocaterini and Ancyronini, have been described within the former Lophocaterinae (Crowson, 1964;Kolib a c, 2006). However, although the groups seem distinct, based mostly on the morphology of mouthparts, the advanced ancyronines could be derived from the paraphyletic lophocaterines (Kolib a c, 2008(Kolib a c, , 2013. Although the lophocateride clade in Gimmel et al. (2019) is not completely sampled and is partly unresolved, the tree indicates the possibility of multiple evolution of the advanced ancyronine features and confirms paraphyly of Lophocaterini. ...
... Cretamerus vulloi was the first lophocaterid beetle from Cretaceous amber to be described (Peris et al., 2014). However, an original placement of similarly basal Decamerini in the former Lophocaterinae (Kolib a c, 2006(Kolib a c, , 2008 was called into question by Bocakova et al. (2016) and Gimmel et al. (2019), who related the decamerine genus Diontolobus to Peltidae or Thymalus, respectively. Considering these molecular studies, the classification of C. vulloi, originally placed at the very base of the lophocateride tree, is, as yet, unclear. ...
Burmacateres longicoxa gen. et sp. nov. is described as the first definite representative of Lophocateridae coming from Cretaceous amber deposits, based on a fossil specimen of Upper Cretaceous Kachin amber, from northern Myanmar. Cleroidea has undergone dramatic taxonomic changes, following the most recent phylogenetic analysis of the group, some of which still need further research and better support. Burmacateres longicoxa gen. et sp. nov. is a well-preserved fossil possessing the appearance and visible morphological features of modern lophocaterids. A saproxylic mode of life, possibly predatory, is inferred for the fossil species, based on similar extant representatives of the family.
... This Trogossitidae s.l. has been divided into three subfamilies, i.e. Peltinae, Lophocaterinae and Trogossitinae (Kolibáč, 2008;Kolibáč & Zaitsev, 2010;Lawrence et al., 2014). Several extensive molecular phylogenetic analyses have been presented in the last decade, either focused on Coleoptera (Bocak et al., 2014), Cucujiformia or Cleroidea (Bocakova et al., 2012(Bocakova et al., , 2016Gunter et al., 2013). ...
... Temnoscheila caerulea was selected as the outgroup in the matrix of morphological characters (Supporting Information, Table S1) following the molecular phylogenetic analysis presented herein, and results of earlier molecular and morphological phylogenies (Kolibáč, 2006(Kolibáč, , 2008Bocak et al., 2014;Bocakova et al., 2016). The tree is rooted at Temnoscheila as a presumed sister to the clade of other studied taxa of Tenebroides. ...
... Remarks: Parallelodera comprises four insular species distributed from New Caledonia to Polynesia and the Marquesas and, following the presented morphological analysis, one species from Panama and Colombia. Because of the lack of material, the genus was not included in the analyses by Kolibáč (2006Kolibáč ( , 2008. Following morphological analysis (Fig. 3), Parallelodera is nested in Tenebroides and is, therefore, classified as a subgenus of the latter genus. ...
A new subgenus of Tenebroides, Polynesibroides subgen. nov., is established for six potentially extinct beetles, Tenebroides atiu, T. moorea, T. raivavae, T. rimatara, T. tubuai and T. mihiura spp. nov., described from the Cook, Austral and Society Islands based on subfossil fragments, and the extant T. tahiti sp. nov. from the island of Tahiti, Society Islands. Here we present the first detailed time-scaled phylogeny of the superfamily Cleroidea inferred from our four-gene dataset, including T. tahiti. Bayesian tip-dating, incorporating 17 fossils, estimated that Cleroidea appeared at the end of the Triassic (~210 Mya). The split of Biphyllidae + Byturidae + Phloiophilidae was reconstructed at ~204 Mya, followed by Early Jurassic separation of Trogossitidae (~200 Mya), Acanthocnemidae (~197 Mya), stem Cleridae-Thanerocleridae-Chaetosomatidae (~194 Mya), Peltidae, Lophocateridae, Decamerinae and the melyrid lineage (~191 Mya). Trogossitidae diversified by separation of Kolibacia (~136 Mya) in the Cretaceous, followed by parting Temnoscheila + Nemozoma and Airora + Tenebroides lineages (~113 Mya). Tenebroides radiated ~75 Mya and T. tahiti diverged from Panamanian taxa in the Middle Eocene (~40 Mya). Fourteen morphological characters of Pacific trogossitids were analysed. Parallelodera, including the Panamanian species, is nested within Tenebroides being sister to Polynesibroides. Therefore, Parallelodera is classified as a subgenus of Tenebroides. Tenebroides fairmairei is placed in the subgenus Parallelodera. A Neotropical origin for Parallelodera and Polynesibroides is inferred.
... This Trogossitidae s.l. has been divided into three subfamilies, i.e. Peltinae, Lophocaterinae and Trogossitinae (Kolibáč, 2008;Kolibáč & Zaitsev, 2010;Lawrence et al., 2014). Several extensive molecular phylogenetic analyses have been presented in the last decade, either focused on Coleoptera (Bocak et al., 2014), Cucujiformia or Cleroidea (Bocakova et al., 2012(Bocakova et al., , 2016Gunter et al., 2013). ...
... Temnoscheila caerulea was selected as the outgroup in the matrix of morphological characters (Supporting Information, Table S1) following the molecular phylogenetic analysis presented herein, and results of earlier molecular and morphological phylogenies (Kolibáč, 2006(Kolibáč, , 2008Bocak et al., 2014;Bocakova et al., 2016). The tree is rooted at Temnoscheila as a presumed sister to the clade of other studied taxa of Tenebroides. ...
... Remarks: Parallelodera comprises four insular species distributed from New Caledonia to Polynesia and the Marquesas and, following the presented morphological analysis, one species from Panama and Colombia. Because of the lack of material, the genus was not included in the analyses by Kolibáč (2006Kolibáč ( , 2008. Following morphological analysis (Fig. 3), Parallelodera is nested in Tenebroides and is, therefore, classified as a subgenus of the latter genus. ...
A new subgenus of Tenebroides, Polynesibroides subgen. nov., is established for six potentially extinct beetles, Tenebroides atiu, T. moorea, T. raivavae, T. rimatara, T. tubuai and T. mihiura spp. nov., described from the Cook, Austral and Society Islands based on subfossil fragments, and the extant T. tahiti sp. nov. from the island of Tahiti, Society Islands. Here we present the first detailed time-scaled phylogeny of the superfamily Cleroidea inferred from our four-gene dataset, including T. tahiti. Bayesian tip-dating, incorporating 17 fossils, estimated that Cleroidea appeared at the end of the Triassic (~210 Mya). The split of Biphyllidae + Byturidae + Phloiophilidae was reconstructed at ~204 Mya, followed by Early Jurassic separation of Trogossitidae (~200 Mya), Acanthocnemidae (~197 Mya), stem Cleridae–Thanerocleridae–Chaetosomatidae (~194 Mya), Peltidae, Lophocateridae, Decamerinae and the melyrid lineage (~191 Mya). Trogossitidae diversified by separation of Kolibacia (~136 Mya) in the Cretaceous, followed by parting Temnoscheila + Nemozoma and Airora + Tenebroides lineages (~113 Mya). Tenebroides radiated ~75 Mya and T. tahiti diverged from Panamanian taxa in the Middle Eocene (~40 Mya). Fourteen morphological characters of Pacific trogossitids were analysed. Parallelodera, including the Panamanian species, is nested within Tenebroides being sister to Polynesibroides. Therefore, Parallelodera is classified as a subgenus of Tenebroides. Tenebroides fairmairei is placed in the subgenus Parallelodera. A Neotropical origin for Parallelodera and Polynesibroides is inferred.
... Kolibáč (2006), in an analysis of adult and larval characters, divided the Trogossitidae into two subfamilies, Trogossitinae [Calityini (as Calityni), Larinotini, Egoliini, Gymnochilini, Trogossitini] and Peltinae (Peltini, Thymalini, Colydiopeltini, Decamerini, Ancyronini, Lophocaterini). This classification was amended by Kolibáč (2008), who included Phloiophilini in Peltinae, and by Kolibáč & Zaitsev (2010) to include a third subfamily, Lophocaterinae (Decamerini, Ancyronini, Lophocaterini) (see Appendix 1B). However, the subsuming of Phloiophilidae into Trogossitidae has not gained general acceptance (see Lawrence & Leschen, 2010a;Bouchard et al., 2011;Kolibáč, 2013;Lawrence et al., 2014). ...
... However, the topology of the ML tree recovered Phloiophilidae sister to Rentoniidae, but this relationship was also unsupported. This enigmatic family with one species endemic to Europe was recently transferred as the tribe Phloiophilini to Trogossitidae: Peltinae by Kolibáč (2008), a placement which has not been adopted by subsequent authors Bouchard et al., 2011;Kolibáč, 2013). Our results reject Kolibáč's (2008) placement in the Peltinae, the core of which are recovered as a distinct monophyletic lineage at the base of the latter clade in our BA tree. ...
... This enigmatic family with one species endemic to Europe was recently transferred as the tribe Phloiophilini to Trogossitidae: Peltinae by Kolibáč (2008), a placement which has not been adopted by subsequent authors Bouchard et al., 2011;Kolibáč, 2013). Our results reject Kolibáč's (2008) placement in the Peltinae, the core of which are recovered as a distinct monophyletic lineage at the base of the latter clade in our BA tree. Kolibáč's (2008) proposed sister groups to Phloiophilini were Colydiopeltini and Thymalini, which were fragmented into multiple lineages throughout our BA and ML trees. ...
The first thorough molecular phylogeny of the superfamily Cleroidea, represented by 377 taxa, and the first with an emphasis on Trogossitidae, was undertaken. Maximum likelihood and Bayesian analyses were performed on a four‐gene dataset (18S, 28S, cox1, cytb) of 395 taxa (along with 18 outgroups), including all 16 currently recognized families of Cleroidea and all current and formerly recognized tribes of Trogossitidae. The superfamily as a whole received strong support in Bayesian analyses. On the basis of phylogenetic results, 18 families in Cleroidea are recognized, including three taxa elevated to family for the first time and two reinstated families. The former tribe Rentoniini (Trogossitidae: Peltinae) was strongly supported as a monophyletic group apart from the remainder of Trogossitidae, and is herein elevated to family status, Rentoniidae stat.n. Protopeltis was also found to be an isolated lineage and becomes Protopeltidae stat.n. Peltini + Larinotini were recovered as a weakly supported sister grouping; Peltini (including only Peltis) becomes Peltidae stat.rest. The trogossitid subfamily Lophocaterinae, to the exclusion of Decamerini, formed a clade which is here designated Lophocateridae stat.rest. and sensu n. The Trogossitinae tribes Calityini, Egoliini (represented by Egolia) and Larinotini were recovered apart from core Trogossitidae but showed no strong affinities to other taxa or congruence between analyses; they are here conservatively retained in Trogossitidae as Calityinae stat.rest., Egoliinae stat.rest. and Larinotinae stat.rest. The genus Thymalus of the peltine tribe Thymalini was indicated with moderate to strong support as the sister group of the Decamerini (Trogossitidae: Lophocaterinae); together these represent Thymalidae stat.n. and sensu n. with subfamilies Decamerinae stat.rest. (new placement) and Thymalinae stat.n. The remainder of Trogossitinae, the tribes Trogossitini and Gymnochilini, formed a well‐supported clade which comprises the Trogossitidae: Trogossitinae sensu n. The tribe Gymnochilini syn.n. is synonymized with Trogossitini. The monotypic family Phloiophilidae was recovered, contradicting a recent placement within Trogossitidae. The melyrid lineage was recovered with moderate (maximum likelihood) to strong (Bayesian analyses) support and includes the families Phycosecidae, Rhadalidae, Mauroniscidae, Prionoceridae and Melyridae (including Dasytidae and Malachiidae). The genus Dasyrhadus is tentatively transferred from Rhadalidae to Mauroniscidae. The genus Gietella, once proposed as a distinct family but recently placed within Dasytidae, was recovered as strongly sister to Rhadalidae sensu n., and we transfer it to that family as Gietellinae new placement. Attalomiminae (formerly Attalomimidae) syn.n. is synonymized with Melyridae: Malachiinae: Lemphini sensu n. Melyridae sensu n. includes only Dasytinae, Malachiinae and Melyrinae. Metaxina is returned to the Chaetosomatidae sensu n., of which Metaxinidae syn.n. becomes a junior synonym. Resolution within Cleridae was generally poor, but a broadly defined Korynetinae stat rest. + Epiclininae received high support (Bayesian analyses). Outside of Trogossitidae, the main focus of this study, major rearrangements of the classification of Cleroidea were not undertaken, despite evidence indicating such changes are needed. We constructed the first molecular phylogeny of Cleroidea to include all families in the superfamily. The heterogeneous family Trogossitidae is restructured into six separate families, Rentoniidae, Protopeltidae, Peltidae, Lophocateridae, Trogossitidae and Thymalidae. A number of other genus‐ to family‐level classificatory rearrangements are undertaken within the superfamily.
... The family Trogossitidae, the bark-gnawing beetles, constitute one of the lesser families of the superfamily Cleroidea, with about 600 recent species described to date (Kolibáč 2013). The modern classification of the Trogossitidae was developed from the classic work of Reitter (1876), later studied by Crowson (1964Crowson ( , 1966Crowson ( , 1970, Barron (1971) and Ś lipiń ski (1992) and, most recently, by Kolibáč (2005Kolibáč ( , 2006Kolibáč ( , 2008 and Kolibáč & Leschen (2010). ...
... Both amber pieces have been deposited at the Senckenberg Deutsches Entomologisches Institut in Seidlitzella is situated at the very base of the gymnochiline clade, closely related to Melambia, Phanodesta, Leperina (incl. present Kolibacia species) in the phylogenetic analysis of Trogossitidae by Kolibáč (2008), whilst the genus forms a polytomy with the cluster Kolibacia (Phanodesta þ Leperina) in an analysis of Gymnochilini by Leschen & Lackner (2013). Yoshitomi (2014) suggests a relation Seidlitzella ((Kolibacia þ Leperina) þ Phanodesta) in his analysis of the gymnochiline ''normal-eyed group''. ...
... Yoshitomi (2014) suggests a relation Seidlitzella ((Kolibacia þ Leperina) þ Phanodesta) in his analysis of the gymnochiline ''normal-eyed group''. Following the above-mentioned morphologically-based phylogenetic analyses (Kolibáč 2008;Leschen & Lackner 2013;Yoshitomi 2014), Palaearctic Kolibacia and Seidlitzella are related to Gondwanan Leperina and Phanodesta, not to the Afro-Asian ''split-eyed group''. External morphological characters of S. hoffeinsorum sp. ...
Based on two well-preserved specimens from late Eocene Baltic amber, a new fossil species belonging to the family Trogossitidae, Seidlitzella hoffeinsorum sp. nov., is described. This is the second known fossil species of the tribe Gymnochilini and the second known species of the genus Seidlitzella . The systematic and biogeographical relations of the genus to other members of the Gymnochilini are discussed. It is hypothesised that the extant eastern Mediterranean species Seidlitzella procera may be phylogenetically related to the genus Phanodesta , today distributed in New Zealand, New Caledonia, Lord Howe Island, Juan Fernandez Island and Sulawesi.
... All trees were modified and annotated in Adobe Photoshop CS6. For illustrations of morphological and anatomical characters of analysed extant cleroid representatives, see Kolibáč (1987Kolibáč ( , 1989aKolibáč ( ,1989bKolibáč ( , 1992Kolibáč ( , 1997Kolibáč ( , 1998Kolibáč ( , 1999Kolibáč ( , 2003Kolibáč ( , 2004Kolibáč ( , 2005Kolibáč ( , 2006Kolibáč ( , 2008 and Majer (1994). The terminology of morphological structures used herein follows Lawrence et al. (2011), character scoring is explained in Table S1 for the matrix, and a list of character states appears in Table S2. ...
Three fossil beetles recorded from Inner Mongolia, China (Callovian, Middle Jurassic, approximately 165 Ma) are assigned to Cleridae and constitute the earliest known representatives of this family. Two of the fossils are described as Protoclerus korynetoides gen. et sp.n. and the third as Wangweiella calloviana gen. et sp.n. Ninety‐six morphological characters of the larvae and adults of 17 extant clerid representatives and seven cleroid families as outgroups were analysed under maximum parsimony with, and without, fossil species included in the matrix. The results indicate that Protoclerus gen.n. is a separate lineage within the family, whereas Wangweiella gen.n. is sister to the rest of Epiclininae. The resulting trees are compared with the most recent morphological and molecular phylogenies of Cleridae. The trees presented appear congruent with the latest molecular phylogenies. The evolution of Cleridae, as well as their biogeography and systematics, are briefly reviewed. A complete list of clerid fossil species described to date is provided.
This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:32240D3D-2A7A-41F6-B485-A66F2265CB8A.
... Today, two species of Peltis (P. ferruginea Linnaeus and P. grossa Linnaeus) are among the most abundant representatives of Trogossitidae in Central Europe; however, there is still much to do in a systematics and phylogeny of the monotypic tribe Peltini, even the subfamily Peltinae (Bocák et al. 2014;Hunt et al. 2007;Kolibáč 2008;Lawrence et al. 2011). Heer (1847Heer ( , 1865 considered the specimen as belonging to Silphidae (Heer 1847(Heer , 1865. ...
A new species of the genus Peltis (Coleoptera, Trogossitidae, Peltinae), P. antehercynica sp. nov., is described from the Pliocene lake sediments of the former clay pit of Willershausen (Lower Saxony, Germany). The clearly visible antennae and elytra sculpturing of fossil provide clear morphological information about generic position, and other features, some of them unique in the Peltinae, show that it is an independent new way. In addition, the importance, geology and stratigraphy of the Willershausen Fossil Lagerstätte is briefly discussed.
