A new genus of Staphylinidae (Coleoptera) from the Lower Cretaceous: the first fossil rove beetles from the Southern Hemisphere

The fossil genus of rove beetles Apticax gen.n . with two new species, A. volans sp.n . and A. solidus sp.n., is described from the Nova Olinda Member of the Crato Formation in north‐eastern Brazil (Aptian–Albian, dated as 125–99.6 Ma old). Both species belong to the clade Staphylininae + Paederinae in the staphylinine group of subfamilies and are the first fossil true Staphylinidae to be described from the entire Southern Hemisphere. Although they resemble Paederinae, the low number of satisfactorily preserved characters do not allow definite placement of either A. volans or A. solidus in any of the subfamilies of Staphylinidae.     

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n. , has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796‐6E70‐4D81‐BB34‐0FEEA765DC25 .     

Mitogenomic phylogenetics of Diochus occultus n. sp., a palaeoendemic endogean species within the tribe Diochini (Coleoptera: Staphylinidae: Staphylininae)

The tribe Diochini has a worldwide distribution, with 2 and 74 epigean species within the genera Antarctothius and Diochus, respectively. Recent phylogenetic studies suggest a sister relationship of Diochini and a lineage formed by Xantholinini, Maorothiini, and Othiini, within the subfamily Staphylininae. Here, we describe the first known endogean representative of Diochini, Diochus occultus n. sp., and provide the first two complete mitogenomes for the tribe, corresponding to the two European Diochus species: Diochus occultus n. sp. and Diochus staudingeri. These sequences were combined with 40 additional mitogenomes from representatives within Staphylininae, Paederinae, Silphidae, and Aleocharinae, and COI sequences from 5 additional species of Diochus to conduct a series of mitogenomic phylogenetic and dating analyses. The estimated molecular phylogeny is fully consistent with previous studies based on morphology and molecular data, finding a sister relationship of Diochini with a clade formed by Xantholinini and Othiini (Maorothiini not sampled). Dating analyses inferred an early split of the tribe Diochini at 140–156 Mya. Morphology shows clear differences in the aedeagal and external morphology of D. occultus n. sp. and D. staudingeri, whereas a sister relationship of these taxa is found in the phylogenetic analyses, with the split dated at 48–61 Mya. Although the study of additional Palaearctic Diochus species will be required to conclusively establish that D. occultus n. sp. is a palaeoendemic taxon sister to D. staudingeri, associated with forests of Abies pinsapo in the south of the Iberian Peninsula, this conclusion is consistent with the ancient estimated age of speciation, endogean habitat specificity, low dispersal capacity (flightless species), and microendemicity of D. occultus. This is also consistent with the continued emersion of the Betic sub‐plate along its tectonic evolution. The estimated ages of diversification of the Paederinae‐Staphylininae lineage are also discussed.     

Early evolution of Cupedidae revealed by a mid‐Cretaceous reticulated beetle from Myanmar (Coleoptera: Archostemata)

Cupedidae, the most species‐rich family of the archaic suborder Archostemata, were abundant, diverse and widespread in the Mesozoic, yet little is known about the early evolution and biogeography. This stems, in part, from a lack of exceptionally preserved fossils from the Mesozoic and of formal phylogenetic study of both extant and extinct taxa. Here we describe and illustrate a new fossil from mid‐Cretaceous Burmese amber, and provide a phylogeny combining both fossils and all known extant genera of Archostemata. A dataset of 43 ingroup taxa and four outgroup taxa based on 110 morphological characters was analysed under parsimony. The results indicate that Priacma LeConte and Paracupes Kolbe, as well as the Cretaceous genera Barbaticupes Jarzembowski et al. and Mallecupes Jarzembowski et al., together form a sister clade to the rest of Cupedidae. Priacma megapuncta sp.n. is attributed to the relict North American Priacma by the presence of distinct subtruncate elytral apices, lateral elytral margins with two rows of sharp teeth, and peculiar fixing epipleural folds near the elytral apices. Our discovery of the first fossil species of Priacma in Burmese amber reveals the antiquity and wider distribution of the genus in the late Mesozoic. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:313565C2‐4F42‐48BD‐8720‐F379DE202868 .     

New fossil Progonocimicidae (Hemiptera: Coleorrhyncha: Progonocimicoidea) from the Upper Mesozoic of northeastern China,with a phylogeny of Coleorrhyncha

A new Progonocimicidae species, Cicadocoris anisomeridis sp.n. , with asymmetrical tegmina is described from the Middle Jurassic Jiulongshan Formation in northeastern China. This is the fifth report of Coleorrhyncha from China. A cladistic analysis based on a combination of fossil and extant taxa clarifies the phylogenetic status of the new fossils and allows the reconstruction of inter‐subfamily relationships within the suborder Coleorrhyncha. Coleorrhyncha is monophyletic and divided into two main clades. Progonocimicidae comprises a monophyletic lineage, to which the new fossils belong. The broadly conceived Progonocimicinae and Cicadocorinae, as recognized by earlier authors, are not supported. The monophyly of the family Karabasiidae is also not supported, and its two constituent subfamilies Hoploridiinae and Karabasiinae are raised to family rank. Hoploridiidae is found to be sister group to all extant moss bugs, and Karabasiidae is found to be the monophyletic sister group to Hoploridiidae + all extant moss bugs. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:30BC0498‐3B8A‐4650‐BC9D‐02A0C8D870B2 .     

Multilocus phylogeny defines a new classification of Staphylininae (Coleoptera,Staphylinidae), a rove beetle group with high lineage diversity

We provide the first multilocus molecular phylogeny of a group corresponding to the former subfamily Staphylininae. Results are corroborated by the morphological, biogeographical and palaeobiological evidence to serve as a baseline for an updated suprageneric classification. The former subfamily Staphylininae is proven to be a lineage sister to the monophyletic Paederinae and reclassified according to a robust phylogeny resolving a number of long-standing controversies. The subfamily Xantholininae (revised status) is reinstated to contain the tribes Xantholinini, Othiini, Maorothiini and Diochini. Subfamily Platyprosopinae (revised status) is reinstated for the tribes Platyprosopinini, Arrowinini and †Thayeralinini. For a highly peculiar genus Coomania Cameron, formerly in Diochini, a new subfamily Coomaniinae subfam.n. is established and the composition of Diochini (revised status) is changed accordingly. The subfamily Staphylininae (revised status) is reduced to contain the former tribe Staphylinini only. Elevating this mega-diverse tribe to the subfamily rank opened up an opportunity for its more fractional classification by raising several subtribes to the tribal level as follows: Acylophorini, Afroquediini, Amblyopinini, Antimerini, †Baltognathini, Cyrtoquediini, Erichsoniini, Hyptiomini, Indoquediini, Quediini and Tanygnathinini (revised status for all). As a result, the most species-rich tribe Staphylinini (revised status) is reduced to the more homogeneous lineage containing the subtribes Algonina, Anisolinina, Philonthina, Philothalpina, Staphylinina and Xanthopygina. Morphological synapomorphies and diagnostic characters supporting all newly defined higher taxa are provided. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:DED8B042-83C9-4D10-B0CB-B50372B067A9 .     

Long‐proboscid brachyceran flies in Cretaceous amber (Diptera: Stratiomyomorpha: Zhangsolvidae)

The monophyletic family Zhangsolvidae comprises stout‐bodied brachyceran flies with a long proboscis and occurring only in the Cretaceous, originally known in shale from the Early Cretaceous Laiyang Formation (Fm.) in China (Zhangsolva Nagatomi & Yang), subsequently from limestones of the Early Cretaceous Crato Fm. of Brazil. Cratomyoides Wilkommen is synonymized with Cratomyia Mazzarolo & Amorim, both from the Crato Fm.; Cratomyiidae is synonymized with Zhangsolvidae. Two genera and three species of Zhangsolvidae are described: Buccinatormyia magnifica Arillo, Peñalver & Pérez‐de la Fuente, gen. et sp.n. and B. soplaensis Arillo, Peñalver & Pérez‐de la Fuente, sp.n. , in Albian amber from Las Peñosas Fm. in Spain; and Linguatormyia teletacta Grimaldi, gen. et sp.n. , in Upper Albian–Lower Cenomanian amber from Hukawng Valley in Myanmar. Buccinatormyia soplaensis and Linguatormyia teletacta are unique among all Brachycera, extant or extinct, by their remarkably long, flagellate antennae, about 1.6× the body length in the latter species. A phylogenetic analysis of 52 morphological characters for 35 taxa is presented, 11 taxa being Cretaceous species, which supports placement of the family within Stratiomyomorpha, although not to any particular family within the infraorder. This published work has been registered in Zoobank, http://zoobank.org/urn:lsid:zoobank.org:pub:F32CF887‐7C37‐45D5‐BD6B‐135FE9B729A7 .     

The first Ironomyiidae from mid‐Cretaceous Burmese amber provides insights into the phylogeny of Phoroidea (Diptera: Cyclorrhapha)

Macalpinomyia jiewenae gen. et sp.n. is described from the mid‐Cretaceous (～99 Ma) amber of Myanmar. Macalpinomyia jiewenae is the first Oriental representative of the enigmatic family Ironomyiidae (Diptera: Phoroidea), currently known from a single extant genus restricted to southeastern Australia, plus a monotypic genus from Canadian amber and three controversial genera based on impression fossils from China, Mongolia and Russia. A phylogenetic analysis of all Phoroidea families, including all ironomyiid extant and extinct genera, corroborates the monophyly of Ironomyiidae, and Macalpinomyia gen.n. is assigned to the subfamily Sinolestinae. Cretonomyiinae subfam.n. , is erected to accommodate the basal lineage of Ironomyiidae. Lebambromyia acrai Grimaldi & Cumming, previously placed in Ironomyiidae, is supported as an early branching lineage of Platypezidae. Our topology proposes that Ironomyiidae is sister to the remaining Phoroidea. The phylogenetic results, in combination with the fossil ages and relevant molecular divergence time analysis, suggests that Ironomyiidae probably originated at least in the Berriasian of the Early Cretaceous (～140 Ma). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:5DFFC944‐1350‐418E‐BCDC‐BB87FC013D5D .     

Phylogeny,divergence times and biogeography of window flies (Scenopinidae) and the therevoid clade (Diptera: Asiloidea)

The evolution of the ‘therevoid’ clade, with an emphasis on window flies (Scenopinidae), is presented by combining DNA sequence data with morphological characters for living and fossil species. The therevoid clade represents a group of four families (Apsilocephalidae, Evocoidae, Scenopinidae and Therevidae) of lower brachyceran Diptera in the superfamily Asiloidea. A comprehensive phylogenetic analysis using parsimony and likelihood methods was undertaken using extensive taxon sampling from all families and subfamilies, and compared with outgroup taxa sampled from the related families Asilidae, Mydidae, Apioceridae and Empididae. Fifty‐nine morphological characters (adult, larval and pupal) were combined with 6.4 kb of DNA sequences for two ribosomal genes (16S and 18S ribosomal DNA) and three protein‐encoding genes [cytochrome oxidase I (COI), triose phosphate isomerase (TPI) and the CPSase region of carbamoyl‐phosphate synthase‐aspartate transcarbamoylase‐dihydroorotase (CAD)]. Results from combined analyses of morphological and molecular data for 78 taxa representing all families of the therevoid clade are presented. Specific hypotheses of the relationship between respective families and subfamilies were tested statistically using four‐cluster likelihood mapping. The therevoid clade is a well‐supported monophyletic group within Asiloidea, with Evocoidae sister to Apsilocephalidae and Therevidae sister to Scenopinidae. Temporal and zoogeographical aspects of therevoid clade evolution were investigated using Bayesian divergence time estimates and Lagrange ancestral range scenarios. The effect of inclusion of fossils as terminal taxa on phylogenetic and divergence time estimation was investigated, with morphological scoring for fossil representatives included in the analyses rather than used simply as minimum age constraints. In each analysis there was either improvement in estimation, or only marginal and localized loss in tree resolution, and with younger estimates of divergence time across the tree. The historical biogeography of the therevoid clade was examined with multiple trans‐Antarctic vicariance events between Australasia and South America evident during the Late Cretaceous to early Palaeogene. Scenopininae is newly subdivided into two tribes, Metatrichini trib.n. and Scenopinini Fallén stat.r. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4974EBF8‐3117‐4189‐B6DE‐7D5BF9B23E53 .     

The oldest known clerid fossils from the Middle Jurassic of China,with a review of Cleridae systematics (Coleoptera)

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 .     

Phylogeny of Stephanidae (Hymenoptera: Apocrita) with a new genus from Upper Cretaceous Myanmar amber

A new wasp species, Lagenostephanus lii Li, Rasnitsyn, Shih and Ren gen. et sp.n. , is assigned to Stephaninae (Hymenoptera: Stephanidae). The specimen is described from the lowermost Upper Cretaceous Myanmar (Burmese) amber. This is the second stephanid wasp described from Myanmar amber after Kronostephanus zigrasi Engel & Grimaldi, representing the earliest fossil occurrences of Stephanidae. Based on morphological characters of all extant and extinct genera, a phylogenetic analysis of Stephanidae was conducted. The results indicate that Schlettereriinae and Stephaninae are monophyletic, whereas Electrostephanus Brues, as the only genus within Electrostephaninae, belongs to Stephaninae. Therefore, we propose a synonymy of Electrostephaninae with Stephaninae. Based on the fossil evidence, we infer that the family of Stephanidae was significantly diverse in the Late Cretaceous and that they originated during the Early Cretaceous or late Jurassic. A key to genera of Stephanidae is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:C082C245-8FD3-42B0-880F-94333286500B .     

Early evolution of Nemopteridae illuminated with the first and oldest thread‐winged lacewing in Cretaceous amber

Nemopteridae are a charismatic family of lacewings distinguished by specialization of morphology in both larvae and adults as well as in their biology and behaviour. The evolutionary history of Nemopteridae is largely unknown due to the scarcity of fossil records. Here we report a new genus and species of Nemopteridae (Cretocroce xiai gen. et sp.n.) based on an exquisitely preserved adult specimen in mid‐Cretaceous Burmese amber. We place the new genus in the subfamily Crocinae, which is characterized by filamentous hindwings with a sensory function as an adaptation to their cavernicolous habitats. The new genus represents the first nemopterid in amber and the oldest crocine known to date. The new fossil crocine is nearly morphologically unchanged over 99 Ma compared with its modern counterparts. Moreover, it is estimated to be closely related to some extant crocines endemic to South America based on morphological phylogenetic analysis. The new finding provides direct evidence to demonstrate the antiquity of Crocinae morphological innovation and related biology, such as pollenophagous habits, and further corroborates Gondwanan origin of the subfamily. The published work has been registered on ZooBank: http://zoobank.org/urn:lsid:zoobank.org:pub:3D29DE48‐DBD8‐4614‐8387‐D9FCF90B0640 .     

New fossils from China elucidating the phylogeny of Praesiricidae (Insecta: Hymenoptera)

A new subfamily of Praesiricidae (Pamphilioidea), Decorisiricinae subfam.n. , is erected based on three new genera: Decorisiricius gen.n. , Limbisiricius gen.n. and Brevisiricius gen.n. Two new species – Decorisiricius patulus gen. et sp.n. and D. longus sp.n. – from the Lower Cretaceous Yixian Formation and three species –Limbisiricius aequalis gen. et sp.n. , Limbisiricius complanatus sp.n. and Brevisiricius partialis gen. et sp.n. – from the Middle Jurassic Jiulongshan Formation, are described. Based on these well‐preserved new fossil specimens and previously published data, the nonmonophyly of Praesiricidae is confirmed and the phylogenetic relationships of species of Praesiricidae are analysed for the first time. Two main clades within Praesiricidae are recognized from the cladistic analysis: Decorisiricinae subfam.n. forms a monophyletic lineage, with the remaining members of Praesiricidae plus Megalodontes (Megalodontesidae) forming its sister group. The two subfamilies Archoxyelydinae and Praesiricinae are discarded with no strong supported synapomorphic characters based on phylogenetic research. A key to all genera of Praesiricidae is provided. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:38D703ED‐127A‐4DB0‐8153‐8D78AF4AC212 .     

Of teeth and trees: A fossil tip‐dating approach to infer divergence times of extinct and extant squaliform sharks

Fossil tip‐dating allows for the inclusion of morphological data in divergence time estimates based on both extant and extinct taxa. Neoselachii have a cartilaginous skeleton, which is less prone to fossilization compared to skeletons of Osteichthyans. Therefore, the majority of the neoselachian fossil record is comprised of single teeth, which fossilize more easily. Neoselachian teeth can be found in large numbers as they are continuously replaced. Tooth morphologies are of major importance on multiple taxonomic levels for identification of shark and ray taxa. Here, we review dental morphological characters of squalomorph sharks and test these for their phylogenetic signal. Subsequently, we combine DNA sequence data (concatenated exon sequences) with dental morphological characters from 85 fossil and extant taxa to simultaneously infer the phylogeny and re‐estimate divergence times using information of 61 fossil tip‐dates as well as eight node age calibrations of squalomorph sharks. Our findings show that the phylogenetic placement of fossil taxa is mostly in accordance with their previous taxonomic allocation. An exception is the phylogenetic placement of the extinct genus ?Protospinax , which remains unclear. We conclude that the high number of fossil taxa as well as the comprehensive DNA sequence data for extant taxa may compensate for the limited number of morphological characters identifiable on teeth, serving as a backbone for reliably estimating the phylogeny of both extinct and extant taxa. In general, tip‐dating mostly estimates older node ages compared to previous studies based on calibrated molecular clocks.     

Small but not ephemeral: newly discovered species of Aphelinidae and Trichogrammatidae (Insecta: Hymenoptera: Chalcidoidea) from Eocene amber

New species of fossil Aphelinidae and Trichogrammatidae are described from middle Eocene (Lutetian) Baltic amber (41.3–47.8 Ma). A new subfamily, two new genera and three new species of Aphelinidae are described, with comments on their placement: Phtuaria fimbriae gen.n. , sp.n. in Phtuariinae subf.n. , Glaesaphytis interregni gen.n. , sp.n. and Centrodora brevispinae sp.n. These represent the first described true fossil Aphelinidae. Four new species of Trichogrammatidae are described: Mirufens illusionis sp.n. , Palaeogramma eos gen.n. , sp.n. , Pterandrophysalis plasmans sp.n. and Szelenyia terebrae sp.n. , thus expanding our knowledge of fossil Trichogrammatidae beyond the single previously described species. The presence of recognizable extant genera of Aphelinidae and Trichogrammatidae in the Eocene suggests that the morphology of these genera has been relatively invariant despite highly variable conditions during and since the Eocene. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:E9AD60B6‐3D56‐4E74‐AA54‐F7B91F4FDC79 .     

A new species of Pseudotettigonia Zeuner (Orthoptera: Tettigoniidae) with an intact stridulatory field and reexamination of the subfamily Pseudotettigoniinae

Pseudotettigonia leona sp.n. is described from a single left forewing collected from the Middle Eocene Kishenehn Formation in northwestern Montana, U.S.A. P. leona is the second of only two fossil macropterous Tettigoniidae that preserve an intact stridulatory apparatus. The subfamily Pseudotettigoniinae is reexamined. The specific epithet bricei is assigned to the late Miocene fossil tettigoniid Tettigonia bricei sp.n. , originally described as the extant Tettigonia viridissima. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:E69E83CE‐9046‐4BBD‐AA11‐E0AD0C382700 .     

Phylogeny of the rove beetle tribe Gymnusini sensu n. (Coleoptera: Staphylinidae: Aleocharinae): implications for the early branching events of the subfamily

The rove beetle subfamily Aleocharinae is the largest subfamily of animals known in terms of species richness. Two small aleocharine tribes, Gymnusini and Deinopsini, are believed to be a monophyletic clade, sister to the rest of the Aleocharinae. Although the phylogenetic relationships of the extant lineages have been well investigated, the monophyly of Gymnusini has been questioned due to a series of previous studies and the recent discovery of the aleocharine †Cretodeinopsis Cai & Huang (Deinopsini) from mid‐Cretaceous Burmese amber. Using an additional specimen of †Cretodeinopsis and well‐preserved specimens of †Electrogymnusa Wolf‐Schwenninger from Eocene Baltic amber, we present here two types of morphology‐based phylogenetic analyses, employing all extant/extinct genera of Gymnusini and Deinopsini for the first time. The maximum parsimony and Bayesian analyses recovered a monophyletic clade of the two tribes combined, but each analysis suggested nonmonophyly of Gymnusini. In agreement with the results of the present study, we synonymize Deinopsini syn.n. under Gymnusini sensu n. , by priority. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:F09EB444‐C6CA‐4525‐A986‐3CFC826F5877 .