On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera,Insecta)

The relationships of extant and extinct lineages of Adephaga were analysed formally for the first time. Emphasis is placed on the aquatic and semiaquatic groups and their evolution in the Mesozoic. ?Triadogyrus and ?Mesodineutus belong to Gyrinidae, the sister group of the remaining families. ?Triaplidae are the sister group of the following groups (Haliplidae, Geadephaga, Dytiscoidea incl. ?Liadytidae, ?Parahygrobiidae and ?Coptoclavidae [major part]). The lack of a ventral procoxal joint and a very short prosternal process are plesiomorphies of ?Triaplidae. ?Coptoclavidae and ?Timarchopsinae are paraphyletic. ?Timarchopsis is placed in a geadephagan clade. In contrast to other coptoclavids, its metathorax is close to the condition found in Haliplidae, with a complete transverse ridge and coxae with large plates and free mesal walls. ?Coptoclavidae s.str., i.e. excl. ?Timarchopsis, is a dytiscoid subgroup. The mesal metacoxal walls are fused, the coxal plates are reduced, and the transverse ridge is absent. ?Stygeonectes belongs to this dytiscoid coptoclavid unit and is therefore misplaced in ?Timarchopsinae. ?Liadytidae belongs to a dytiscoid subgroup, which also comprises the extant families Aspidytidae, Amphizoidae, Hygrobiidae and Dytiscidae. ?Parahygrobia is the sister group of Hygrobiidae. The larvae are characterized by a broad gula, the absence of the lacinia, retractile maxillary bases and very long urogomphi set with long setae. ?Liadytiscinae is the sister group of extant Dytiscidae. There is no support for a clade ?Eodromeinae and for Trachypachidae incl. ?Eodromeinae. ?Fortiseode is nested within Carabidae. The exclusion of fossil taxa has no effect on the branching pattern. The evolution of Adephaga in the Mesozoic is discussed. Possible reasons for the extinction of ?Coptoclavidae are the rise of teleost fish and the competition of Gyrinidae and Dytiscidae, which possess efficient defensive glands and larval mandibular sucking channels.     

A new ripiphorid beetle from Upper Cretaceous Burmese amber sheds light on early evolution of the extant subfamily Ripidiinae (Coleoptera: Ripiphoridae)

Fossils belonging to Ripidiinae are rare. Here, we describe and figure a remarkable new genus and species of Ripidiinae, Protoripidius burmiticus gen. et sp. nov., from the Upper Cretaceous amber of Myanmar. Protoripidius can be placed in the extant subfamily Ripidiinae based on the widely separated and abbreviated elytra (in male), unfolded hind wings, simple claws and the absence of tibial spurs. Protoripidius bears many transitional features shedding new light on the relationships of the two extant tribes of Ripidiinae. The discovery also highlights the palaeodiversity of the subfamily Ripidiinae in the mid-Cretaceous.     

Phylogeny of the Saprininae reveals interesting ecological shifts in the history of the subfamily (Coleoptera: Histeridae)

A morphological data set for the histerid beetle subfamily Saprininae comprising 95 adult morphological characters scored (multistate coding) from 72 terminal taxa and four outgroups was developed in order to analyse and determine the relationships amongst the genera and subgenera of the Saprininae subfamily. Cladograms were rooted with exemplars of Dendrophilinae (genus Dendrophilus), Bacaniini (genus Bacanius), Abraeinae (genus Chaetabraeus), and Anapleini (genus Anapleus). Parsimony‐based phylogenetic analyses were performed based on the type species of each genus and subgenus of the Saprininae occurring around the world, with the exception of three taxa: Paramyrmetes foveipennis (type species of the genus Paramyrmetes), Satrapister nitens (type species of the genus Satrapister) and Xerosaprinus (Auchmosaprinus) laciniatus (type species of the subgenus Auchmosaprinus) that were not available. In addition, in order to test the monophyly of several questionable genera, multiple exemplars were added in a few cases. The analysis also included an exemplar of an apparently undescribed genus. The results of the analysis confirm the monophyly of the subfamily supported by two unique synapomorphies: (1) presence of sensory structures of the antenna; and (2) presence of the antennal cavity, as well as several other weaker synapomorphies. However, the phylogeny inferred here shows mostly low support for the deeper branches and consequently no major changes in the Saprininae classification are proposed. The presented cladogram is discussed together with its implications for the evolution of the subfamily. The most informative characters and their respective states are outlined. Multiple shifts in lifestyles have evolved during the evolutionary history of the group. Taxa found near the root of the cladogram are mostly nidicolous or myrmecophilous, and inquiliny is presumed to be the plesiomorphic lifestyle of the subfamily. The nidicolous lifestyle has undergone several transformations to other lifestyles and myrmecophily has evolved three times independently during the evolution of the subfamily. Termitoxeny has evolved two times independently in the group whereas ecological adaptation for life in caves has likewise evolved two times independently. The analyses yielded a large clade of predominantly psammophilous taxa; psammophily is thought to have evolved once and has been subsequently lost several times. © 2014 The Linnean Society of London     

Phylogeny and the fossil record of the Helophoridae reveal Jurassic origin of extant hydrophiloid lineages (Coleoptera: Polyphaga)

We performed a phylogenetic analysis focused on the hydrophiloid family Helophoridae (Coleoptera: Polyphaga) in order to test the phylogenetic position of selected Mesozoic fossils assigned to the Hydrophiloidea. The analysis is based on 92 characters of larvae and adults, and includes all extant subgenera of Helophorus and representatives of all other extant hydrophiloid families. Based on this analysis, we provide additional evidence for the monophyly of the helophorid lineage containing the families Helophoridae, Georissidae and Epimetopidae, as well as the first hypothesis on the phylogenetic relationships within Helophorus, revealing three main clades: Lihelophorus, Rhopalohelophorus and the clade of sculptured small subgenera; the subgenera Helophorus s.str., Gephelophorus, Trichohelophorus and Transithelophorus are recognized as paraphyletic or polyphyletic. Inclusion of fossil species in the analysis reveals the Mesozoic genera Hydrophilopsia Ponomarenko, Laetopsia Fiká?ek et al. (adult forms) and Cretotaenia Ponomarenko (larval form) as basal extinct clades of the helophorid lineage, the former genus Mesosperchus Ponomarenko as containing probable stem taxa of Helophorus and the former genus Mesohelophorus Ponomarenko as a member of the Helophorus clade containing extant sculptured subgenera. The extant subgenus Thaumhelophorus syn.nov. is synonymized with Rhopalohelophorus. Our results show that the family Helophoridae may be dated back to the late Jurassic (c. 150 Ma) and the extant clades of Helophorus back to the Early Cretaceous (c. 136 Ma). The basal groups of Helophorus and the supposed basal extinct lineages of the helophorid lineages are shown to be aquatic as adults. A single origin of trichobothria and ventral hydrophobic pubescence in the common ancestor of the Hydrophiloidea is hypothesized, indicating ancestral aquatic habits in the adult stage for the whole Hydrophiloidea.     

Phylogeny of the subfamily Chauliodinae (Megaloptera: Corydalidae), with description of a new genus from the Oriental Realm

Abstract.  A new Oriental fishfly genus, Sinochauliodes gen.n. , is described, including four species: S. fujianensis ( Yang & Yang, 1999 ) comb.n. , S. griseus ( Yang & Yang, 1999 ) comb.n. , S. maculosus sp.n. and S. squalidus sp.n. A cladistic analysis based on adult morphological characters clarified the phylogenetic status of the new genus and allowed the reconstruction of the intergeneric relationships of the subfamily Chauliodinae. Two main clades within Chauliodinae were recognized from the cladistic analysis. The Asian fishflies, together with the two Nearctic genera, Chauliodes and Nigronia , formed a monophyletic lineage, and the new genus was assigned as the sister group to the genus Parachauliodes . The biogeography of the Asian fishflies is discussed.     

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

The phylogeny of the basal hymenopteran lineages, including representatives of all ‘symphytan’ families, is anal; In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelic tenthredinoid, five pamphilioid, three cephoid, five ‘siricoid’, one orussid, and six apocritan taxa. The datas analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenopte strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relations of the ‘symphytan’ lineages are Xyeloidea +(Tenthredinoidea+ (Pamphilioidea + (Cephoidea + (Ariaxyelic (Siricidae + (Xiphydriidae +(Orussoidea+Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is supported, and all may be monophyletic except the ‘Siricoidea’, which is clearly paraphyletic. It is difficult to di whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae + (Orussoidea + Apocrita). support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative apomorphies being provided by most character systems. There is also good evidence in favour of the monophj the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier anal The Blasticotomidae are the sistergroup of the Tenthredinoidea s.s. Relationships at the base of the Tenthredini s.s. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a 1 paraphyletic grade within the Tenthredinoidea s.s. The Diprionidae may be the sistergroup to Cimbicidae +(Argidae+ Pergidae). Most relationships within the Cimbicidae + (Argidae + Pergidae) clade are corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae the Xiphydriida both to superfamily status. The family‐level classification of the Tenthredinoidea will probably have to be changed, but this must await further clarification of the phylogeny of this superfamily.     

Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera

Phytophagous ladybird beetles of the tribe Epilachnini are a cosmopolitan, species‐rich group of significant economic importance as pests of agricultural crops. The tribe is well characterized morphologically and clearly monophyletic, but very little is known about its internal phylogenetic relationships and their genus‐level taxonomy. In order to infer the evolutionary history of Epilachnini, test its monophyly and provide a phylogeny‐based classification, we assembled a comprehensive dataset, consisting of four DNA markers (18S and 28S rRNA and 16S, COI mtDNA) and a matrix of 104 morphological characters for 153 species of Epilachnini representing all previously recognised genera, ～11% of the known species, and 14 outgroup taxa. Molecular, morphological and combined datasets were analysed using maximum likelihood, parsimony and Bayesian inference. Bayes factors and Approximately Unbiased tests (AU) were used to compare alternative phylogenetic hypotheses of unconstrained and backbone‐constrained analysis. Only 14 of the 25 included genera were recovered monophyletic, as originally defined. Afidentula Kapur, Afidenta Dieke, Afissula Kapur, Epilachna Chevrolat, Henosepilachna Li Toxotoma Weise and Mada Mulsant are shown to be poly‐ or paraphyletic; Chnootriba Chevrolat, Subafissa Bielawski, Lalokia Szawaryn & Tomaszewska and Papuaepilachna Szawaryn & Tomaszewska form monophyletic groups within larger clades of genus level. All of these genera are redefined here. The two largest genera of Epilachnini, Epilachna Chevrolat and Henosepilachna Li were represented by multiple monophyletic clades, which we described as new genera: Chazeauiana Tomaszewska & Szawaryn gen.n. ; Diekeana Tomaszewska & Szawaryn gen.n .; Fuerschia Tomaszewska & Szawaryn gen.n. and Ryszardia Tomaszewska & Szawaryn gen.n . The following new synonyms are proposed: Afissa Dieke (=Afissula Kapur); Henosepilachna Li in Li & Cook (=Subafissa Bielawski); Papuaepilachna Szawaryn & Tomaszewska (=Lalokia Szawaryn & Tomaszewska). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:440E7FA4‐C859‐47E0‐8335‐30D478CBA8FA .     

A new genus of the subfamily Anaidinae (Coleoptera, Scarabaeoidea, Hybosoridae) from the Mesozoic of Transbaikalia

A new genus and species, Protanaides sibiricus gen et sp. nov., of the subfamily Anaidinae (family Hybosoridae) from the Lower Cretaceous Baisa locality in Transbaikalia is described. This find adds considerably to knowledge of the diversity of this subfamily in the Mesozoic of Asia and gives evidence of the relict status of the Recent Anaidinae.     

Phylogeny of Evanioidea (Hymenoptera,Apocrita), with descriptions of new Mesozoic species from China and Myanmar

The phylogeny of the superfamily Evanioidea is presented using morphology and DNA sequence data of selected extant and fossil genera by employing two phylogenetic methods, maximum parsimony and Bayesian inference. Based on our new results, the monophyly of Evanioidea is corroborated. Evanioidea, Anomopterellidae, Othniodellithidae, Andreneliidae and Evaniidae are monophyletic families, while Praeaulacidae, Aulacidae, Baissidae and Gasteruptiidae are paraphyletic families. In addition, four new genera (Sinuevania gen.n. , Curtevania gen.n. , Exilaulacus gen.n. , Heterobaissa gen.n. ) with five new species (Sinuevania mira sp.n. , Curtevania enervia sp.n. , Exilaulacus loculatus sp.n. , Exilaulacus latus sp.n. , Heterobaissa apetiola sp.n. ), and five additionally new species (Newjersevania longa sp.n. , Newjersevania brevis sp.n. , Cretevania tenuis sp.n. , Cretevania venae sp.n. , Praeaulacus rectus sp.n. ) and one new combination [Cretevania mitis (Li, Shih & Ren, 2014a) comb.n. ] are described based on well‐preserved fossils from the Middle Jurassic Jiulongshan Formation in Inner Mongolia, China, the Early Cretaceous Yixian Formation in Liaoning, and mid‐Cretaceous amber from Myanmar. This study documents the diversification of one major lineage of the mid‐Mesozoic parasitoid revolution that dramatically changed food‐web relationships in terrestrial ecosystems. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CBC04ADA‐0176‐402D‐9B43‐E1B3CDA080E1 .     

Mesozoic leaf beetles of the tribe Mesolpinini trib. nov. (Coleoptera,Chrysomelidae) from the Lower Cretaceous

This publication is based on recent studies of Lower Cretaceous leaf beetles from the Yixian Formation (Liaoning, China), which are represented by five new species of one new genus Mesolpinus gen. nov. (M. antenattus sp. nov. [type species], M. adapertilis sp. nov., M. angusticollis sp. nov., M. basicollis sp. nov., and M. trapezicollis sp. nov.) assigned to a new tribe, Mesolpinini trib. nov. of the subfamily Chrysomelinae. This tribe, which includes only species from the Jehol biota, is the oldest known group of the family in the fossil record. A key to species of the genus Mesolpinus gen. nov. is provided and the position of the new tribe is discussed. A brief overview of the Mesozoic data on the subfamily Chrysomeloidea is given.     

A new genus of the subfamily Apioninae (Coleoptera: Brentidae) from the Lower Eocene Oise amber

A new genus and species, Palaeotanaos oisensis, is the first described representative of Apioninae from the Lower Eocene Oise amber. The new genus belongs to the tribe Tanaini and is similar to the genus Tanaos Schoenherr, 1826 and differs from it in the convex, not flattened, and slightly elongate body, the antennae inserted in basal one-third of rostrum, comparatively long flagellum, and elongate club, large and strongly convex eyes, narrow and long tibiae without mucro, short precoxal part of prothorax, and narrower and long tarsi.     

Wing morphology of a new Cretaceous praying mantis solves the phylogenetic jigsaw of early‐diverging extant lineages

The extremely derived morphology and behaviour of extant praying mantises combined with a scarce record of fossil relatives introduce significant challenges to tracing their evolution from Palaeozoic stem‐dictyopterans. Extant members of Chaeteessidae, Mantoididae and Metallyticidae could be invaluable to resolving the mantodean tree, yet their inclusion in phylogenetic analyses led to conflicting hypotheses due to their highly disparate respective morphologies. In this contribution, we present Labradormantis guilbaulti gen. et sp.n. , a new fossil species described from both fore‐ and hind‐wing imprints discovered in the Redmond Mine locality (Late Cretaceous, Cenomanian, Redmond Formation; Labrador, Canada). The examination of its hind‐wing AA2* supports the hypothesis that this structure, unique to Chaeteessidae among extant mantises, is a true vein and that its occurrence represents a plesiomorphy for Mantodea. A parsimony analysis including newly coded wing‐related characters further established that L. guilbaulti gen. et sp.n. displays a unique combination of plesiomorphic and apomorphic character states that situates it within the extinct family Baissomantidae. This dataset resolved the phylogenetic relationships of early‐diverging extant lineages as (Chaeteessidae (Mantoididae (Metallyticidae, Artimantodea))), and suggested that the Eocene Lithophotina floccosa Cockerell might be a close relative of extant metallyticids. It also indicated a trend towards increased modularity within mantis fore‐wings, in contrast with a trend towards increased morphological integration in their hind‐wings, both of which are potentially associated with improved flight performance for modern mantises. This study emphasizes the importance of fossils for resolving phylogenetic relationships and for introducing transitional phenotypes to infer ancient evolutionary trends of extant derived clades.     

A new subfamily of ichneumonids from the Lower Cretaceous of Transbaikalia and Mongolia (Insecta: Hymenoptera: Ichneumonidae)

A new subfamily of ichneumonids, Palaeoichneumoninae, is described from the Lower Cretaceous of Transbaikalia (Baisa locality) and Mongolia (Bon Tsagan and Kholbotu Gol localities). The new subfamily is intermediate between the archaic subfamily Tanychorinae and the Recent Ichneumonidae. It includes 12 new species, described in three new genera: Palaeoichneumon freja gen. et sp. nov., P. ornatus sp. nov., P. danu sp. nov., P. micron sp. nov., P. mirabilis sp. nov., P. tenebrosus sp. nov., P. townesi sp. nov., Rudimentifera mora gen. et sp. nov., R. suspecta sp. nov., Dischysma maculata gen. et sp. nov., D. similis sp. nov., and D. ramulata sp. nov.     

First Mesozoic Scutigeromorph Centipede, from the Lower Cretaceous of Brazil

The first Mesozoic scutigeromorph centipede (Myriapoda: Chilopoda), Fulmenocursor tenax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. Previously described fossil Scutigeromorpha are known from Dominican and Baltic amber, the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois, the Silurian and Devonian of Britain, and the Devonian of New York State.     

Phylogeny and host-specificity of European seed beetles (Coleoptera, Bruchidae), new insights from molecular and ecological data

We used partial sequences of three mitochondrial genes (12S rRNA, cytochrome b, and cytochrome c oxidase subunit I) to reconstruct the phylogeny of European seed beetles (Bruchidae) belonging to the genera Bruchus Linnaeus and Bruchidius Schilsky. Adult beetles examined in this study were obtained from larvae bred from seeds directly collected in the field. Parsimony, maximum likelihood, and Bayesian inference were used to infer phylogenetic relationships among species. Both genera, Bruchidius and Bruchus, formed monophyletic groups in all analyses. Our results were partially in discrepancy with existing taxonomic groups (Borowiec, 1987). Critical analysis of relationships among taxa, and exhaustive review of host-plants data highlight the very high level of specialization of these seed beetles. Phylogenetically related insects were associated with host-plants belonging to the same botanical tribes.     

Description of a new species of Sternocoelis from Morocco with proposal of the Sternocoelis marseulii species group (Coleoptera, Histeridae)

TheSternocoelis marseulii species group is proposed based on antennal and prosternal characters. Five species are included in the group:Sternocoelis marseulii (Brisout de Barneville, 1866)(Spain), Sternocoelis viaticus Lewis, 1892 (Algeria), Sternocoelis vaucheri Lewis, 1896 (Morocco), Sternocoelis berberus Lackner & Yélamos, 2001 (Morocco)and Sternocoelis yelamosisp. n. (Morocco). The external morphology of Sternocoelis yelamosisp. n. is described and illustrated, the illustrations of genitalia of all species of the group (except for Sternocoelis vaucheri) are provided and a key to the species of the group is given.     

Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera)

Ensifera present an appropriate and interesting model for the study of acoustic communication, because of their diverse signal and communication modalities, and due to their accessibility for field and laboratory studies. Several hypotheses have been proposed to explain the acoustic evolution of Ensifera, but they were elaborated without any reference to a falsifiable phylogeny, and were consequently highly speculative. Similarly, phylogenetic relationships between ensiferan clades have not hitherto been studied using modern standard methodology, and the sole cladistic analysis by Gwynne in 1995 was methodologically flawed. No sound hypothesis therefore currently exists for ensiferan phylogeny, which precludes historical analysis of their communication modalities. In the present paper, the phylogeny is established on the basis of morpho‐anatomical characters and used to analyse the evolution of acoustic communication in this clade by mapping the characters related to auditory and stridulatory structures onto the resultant trees. Cladistic analyses resulted in two equi‐parsimonious cladograms (length 154, C 64, CI 58, RI 61) with the following topologies: (1) [(Grylloidea–Gryllotalpidae) (Rhaphidophoridae (Schizodactylidae (Gryllacrididae ((Stenopelmatidae–Cooloola) (Anostostomatidae (Prophalangopsis (Cyphoderris (Tettigoniidae–Lezina))))))))] (2) [(Grylloidea–Gryllotalpidae)(Rhaphidophoridae (Schizodactylidae (Gryllacrididae–Cooloola–(Stenopelmatidae (Anostostomatidae (Prophalangopsis (Cyphoderris (Tettigoniidae–Lezina))))))))]. According to these topologies, Ensifera were ancestrally devoid of acoustic and hearing systems. An acoustic (tegminal or femoro‐abdominal) apparatus appeared a number of times independently with convergent structures. Similarly, tibial tympana developed several times independently. Moreover, four hypotheses (each according to a definite pattern of character transformation) can be proposed to explain the evolution of acoustic communication in the different ensiferan clades and relate it to a definite communicatory context. These hypotheses do not apply equally to ensiferan subclades. Grylloidea and Gryllotalpoidea could have experienced convergently a direct development of an intraspecific acoustic communication. Acoustic communication in Tettigoniidea has evolved more ambiguously, and may either have resulted from a direct evolution analogous to that having occurred in Gryllidea, or have developed in a completely different behavioural context. Future studies of acoustic communication in the different ensiferan clades will have to take into account the fact that the involved structures most often are not homologous and that their evolution may not have taken place in similar conditions. Different hypotheses of acoustic communication evolution may apply to different clades, and there may be no single explanation for acoustic communication in Ensifera.     

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.