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 .     

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 .     

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 .     

Emergence of a superradiation: pselaphine rove beetles in mid‐Cretaceous amber from Myanmar and their evolutionary implications

Pselaphinae is an exceptionally species‐rich, globally distributed subfamily of minute rove beetles (Staphylinidae), many of which are inquilines of social insects. Deducing the factors that drove pselaphine diversification and their evolutionary predisposition to inquilinism requires a reliable timescale of pselaphine cladogenesis. Pselaphinae is split into a small and highly plesiomorphic supertribe, Faronitae, and its sister group, the ‘higher Pselaphinae’ – a vast multi‐tribe clade with a more derived morphological ground plan, and which includes all known instances of inquilinism. The higher Pselaphinae is dominated by tribes with a Gondwanan taxonomic bias. However, a minority of tribes are limited to the Nearctic and Palearctic ecozones, implying a potentially older, Pangaean origin of the higher Pselaphinae as a whole. Here, I describe fossils from mid‐Cretaceous (～99 million years old) Burmese amber that confirm the existence of crown‐group higher pselaphines on the Eurasian supercontinent prior to contact with Gondwanan landmasses. Protrichonyx rafifrons gen. et sp.n. is placed incertae sedis within the higher Pselaphinae. Boreotethys gen.n. , erected for B. grimaldii sp.n . and B. arctopteryx sp.n. , represents an extinct sister taxon and putative stem group of Bythinini, a Recent tribe with a primarily Holarctic distribution. The Laurasian palaeolocality of the newly described taxa implies that higher pselaphines are indeed probably of Jurassic, Pangaean extraction and that the Laurasian‐Gondwanan tribal dichotomy of this clade may have developed vicariantly following Pangaean rifting. Higher pselaphines probably predate the earliest ants. Their physically protective morphological ground plan may have been a preadaptation for myrmecophily when ants became diverse and ecologically ubiquitous, much later in the Cenozoic. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:36E3FE2A-B947-422D-89CA-0EF43B99C382 .     

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 .     

A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure

An unusual Cretaceous trap jaw ant is described from Burmese amber dated to the Late Cretaceous. Linguamyrmex vladi gen.n. sp.n. is distinguished by an unusual suite of morphological characters indicating specialized predatory behaviour and an adaptive strategy no longer found among modern ant lineages. The clypeus, highly modified as in other closely related haidomyrmecine hell ants, is equipped with a paddle‐like projection similar to Ceratomyrmex. X‐ray imaging reveals that this clypeal paddle is reinforced, most probably with sequestered metals. Presumably this fortified clypeal structure was utilized in tandem with scythe‐like mandibles to pin and potentially puncture soft‐bodied prey. This unique taxon, which stresses the diversity of stem‐group ants, is discussed in the context of modern and other Cretaceous trap jaw ant species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:40D636A3‐4D88‐470A‐BC5B‐85ABFD1A49E2 .     

First discovery of fossil Coloninae in Cretaceous Burmese amber (Coleoptera,Staphylinoidea, Leiodidae)

Coloninae is a tiny and distinctive subfamily of the highly diverse family Leiodidae (round fungus and small carrion beetles). Although only one leiodid fossil is known from the Mesozoic, its taxonomic position and classification are doubtful. We describe the first definitive Coloninae fossil, Colon burmiticum Yamamoto sp. nov., from Upper Cretaceous amber of Myanmar. Our finding sheds light on the origin and early evolutionary history of this peculiar genus and its subfamily.     

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 .     

The first Mesozoic Leptopodidae (Hemiptera: Heteroptera: Leptopodomorpha), from Canadian Late Cretaceous amber

A new genus and species of leptopodid bug, Cretaceomira phalanx McKellar and Engel, is described from Canadian Late Cretaceous (Campanian) amber originating near Grassy Lake, in southern Alberta, Canada. This new record is the first described for the family within the Mesozoic, extending their fossil range by at least 26 Ma. The discovery adds further support to the idea that the subfamily was once much more widespread than its modern, relict distribution in the tropics – adding an occurrence in warm temperate conditions, on the western side of Laurentia (in the modern Palearctic). Beyond confirming the presence of the lineage in the Cretaceous, their expanded distribution suggests that the group is likely to be found in other Cretaceous amber deposits. Furthermore, the distinctive disk-shaped amber nodule that contains the C. phalanx holotype provides limited support for the interpretation of Leptosaldinae as subcortical inhabitants of resin-producing trees as early as the Cretaceous.http://www.zoobank.org/urn:lsid:zoobank.org:pub:E324DF2B-8D99-42B3-BBAC-8F9DC3603490     

The earliest fossil record of the extant rove beetle genus Phloeocharis from mid-Cretaceous Kachin amber of northern Myanmar and its biogeographic implications (Coleoptera: Staphylinidae: Phloeocharinae)

Phloeocharis Mannerheim is the largest genus within the problematic rove beetle subfamily Phloeocharinae, with a single extinct and 44 recent species recorded from the Holarctic Region. Until now, the oldest fossil record of Phloeocharis was known from Late Cretaceous (Turonian) amber from New Jersey, USA. Here we describe 2Phloeocharis burmana n. sp. from mid-Cretaceous (Albian–Cenomanian) Kachin amber from northern Myanmar, as the earliest extinct species of this genus. Our finding also sheds light on the biogeography of Phloeocharis, since no recent or extinct species have so far been recorded from the Oriental Region. Furthermore, the discovery of 2P. burmana n. sp. extends the Mesozoic diversity of the phloeocharine rove beetles both taxonomically and morphologically, particularly from Kachin amber.     

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 .     

100 million years of morphological conservation in bark beetles (Coleoptera: Curculionidae: Scolytinae)

Abstract Scolytine weevils (bark and ambrosia beetles) have a unique ecological significance in forest ecosystems, which equates to major effects on landscape ecology and to monetary losses. Fossilized galleries of scolytines have been reported in Late Mesozoic wood, but here we describe a well‐preserved body fossil from the Cretaceous, c. 100 Ma, preserved in amber from northern Myanmar. Moreover, the specimen is remarkably similar to Recent species of the genus Microborus, revealing stasis unexpected within scolytines and thus highlighting the antiquity of the group. Stratigraphic dating and comparison of insect palaeofaunas included in other well‐dated ambers from multiple sites support the age estimate of the Burmese amber. A minimum age for one clade of scolytines is thus established, indicating an early divergence of scolytines from other weevils in the Late Jurassic or Early Cretaceous and challenging the current perspective of weevil evolution.     

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 .     

The past and the present through phylogenetic analysis: the rove beetle tribe Othiini now and 99 Ma

In order to classify and taxonomically describe the first two fossil Othiini (Coleoptera: Staphylinidae: Staphylininae) species from three well‐preserved specimens in Cretaceous Burmese amber, a phylogenetic analysis was conducted, combining extant and extinct taxa. A dataset of 76 morphological characters scored for 33 recent species across the subfamilies Staphylininae and Paederinae was analysed using maximum parsimony and Bayesian inference methods. The many differing phylogenetic hypotheses for higher‐level relationships in the large rove beetle subfamilies Staphylininae and Paederinae were summarized and their hitherto known fossil record was reviewed. Based on the analyses, the new extinct genus Vetatrecus gen.n. is described with two new species: V. adelfiae sp.n. and V. secretum sp.n. Both species share character states that easily distinguish them from all recent Othiini and demonstrate a missing morphological link between subfamilies Staphylininae and Paederinae. This is the first morphology‐based evidence for the paraphyly of Staphylininae with respect to Paederinae, suggested earlier by two independent molecular‐based phylogenies of recent taxa. Our newly discovered stem lineage of Othiini stresses the importance of fossils in phylogenetic analyses conducted with the aim of improving the natural classification of extant species. It also suggests that the definitions of Staphylininae and Paederinae, long‐established family‐group taxa, may have to be reconsidered. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:817F39C4-F36B-4FD9-96CD-5F8FB064C39E .     

†Cormopsocidae: A new family of the suborder Trogiomorpha (Insecta: Psocodea) from Burmese amber

Cormopsocidae n. fam . of the psocid suborder Trogiomorpha was proposed for a fossil species from mid‐Cretaceous Burmese amber, Cormopsocus groehni n. gen . & n. sp . This family was estimated to be the sister group of all other trogiomorphan taxa, but the possibility of much deeper divergence (i.e. placement as a sister group of all Psocodea) could not be excluded. Cormopsocus groehni retains many plesiomorphic features, which will contribute greatly to elucidating the ancestral state of Psocodea.     

Burmaphlebia reifi gen. et sp. nov., the first anisozygopteran damsel-dragonfly (Odonata: Epiophlebioptera: Burmaphlebiidae fam. nov.) from Early Cretaceous Burmese amber

A new family, genus and species of damselfly, Burmaphlebia reifi gen. et sp. nov. (Burmaphlebiidae fam. nov.), is described as the second fossil odonate from Early Cretaceous Burmese amber. Its phylogenetic position is discussed and the fossil is attributed to a new family at the base of the anisozygopteran grade, probably closely related to the Recent relict group Epiophlebiidae. It is the first record of the ‘anisozygopteran’ grade from amber and the smallest known representative of this group.http://zoobank.org/6EFE7288-BD89-42F9-BFA5-804CE6B904A6     

Fossil water striders in Cretaceous French amber (Heteroptera: Gerromorpha: Mesoveliidae and Veliidae)

Fossil gerromorphan bugs from the Cretaceous (Albian–Cenomanian boundary) amber of Charentes, SW France, are reviewed. A larva described by Perrichot et al. (2005) as incertae familiae within the Gerromorpha is now placed in the Mesoveliidae. Three new genera and species are also described and illustrated: Emilianovelia audax Solórzano Kraemer & Perrichot, gen. et sp.n. , and Malenavelia videris Solórzano Kraemer & Perrichot, gen. et sp.n. , which are placed in the Mesoveliidae: Mesoveliinae; and Arcantivelia petraudi Solórzano Kraemer & Perrichot, gen. et sp.n. , which is the first Mesozoic member of the Veliinae. The relationships between these fossils and their palaeoecology are discussed. The new discoveries confirm the antiquity of the semi‐aquatic gerromorphans, particularly the clade (Veliidae + Gerridae). The habitat is described as aquatic and likely marine‐influenced, yet adaptation to a fully marine habitat in these gerromorphans remains difficult to establish. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:F8C034B8-7BD0-4C03-B8FC-CD81D75C7001 .     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.