Snipe flies (Diptera: Rhagionidae) from the Daohugou Formation (Jurassic), Inner Mongolia,and the systematic position of related records in China

Abstract: Four new species referred to three new genera and one known genus of fossil snipe flies (Family Rhagionidae) from the Daohugou Formation of Chifeng City in Inner Mongolia, north‐eastern China, are described as Daohugorhagio elongatus gen. et sp. nov., Parachrysopilus jurassicus gen. et sp. nov., Sinorhagio sinuatus sp. nov. and Trichorhagio gregarius gen. et sp. nov. These taxa represent only a fraction of the rhagionid assemblage in the Daohugou biota. They contribute towards the high diversity and abundance of snipe flies during the latest Middle Jurassic – earliest Late Jurassic. The diagnosis of Sinorhagio K. Zhang, Yang and Ren, 2006 is revised based on the new results. Many genera and species from the Lower Cretaceous of China have been assigned to Rhagionidae. Most of these do not actually belong to this family: specifically Mesorhagiophryne incerta Hong and Wang, 1990, M. robusta Hong and Wang, 1990, Mesostratiomyia laiyangensis Hong and Wang, 1990, Stratiomyopsis robusta Hong and Wang, 1990, Longhuaia orientalis Hong, Wang and Sun, 1992, Basilorhagio venusius Ren, 1995. Pauromyia oresbia Ren, 1998 is transferred to an archisargid genus as Sharasargus oresbius (Ren, 1998) comb. nov.     

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 .     

Taxonomic review and phylogenetic inference elucidate the evolutionary history of Mesozoic Procercopidae,with new data from the Cretaceous Jehol Biota of NE China (Hemiptera,Cicadomorpha)

The Mesozoic family Procercopidae is widely treated as the ancient group of Cercopoidea and a transitional unit to recent lineages, but its evolution and diversity are vague due to fragmentary fossil record and confusing taxonomic history. Herein, an extensive taxonomic review of Procercopidae is presented and some new fossils are reported from the Lower Cretaceous Yixian Formation of NE China. As a result, Chengdecercopis Hong, 1983 is transferred from Procercopidae to Sinoalidae; Procercopis longipennis Becker-Migdisova, 1962 and P shawanensis Zhang, Wang and Zhang, 2003 are transferred to Procercopina Martynov, 1937, resulting in Procercopina longipennis (Becker-Migdisova, 1962), comb. n. and P shawanensis (Zhang, Wang and Zhang, 2003), comb. n.; Luanpingia senjituensis Hong, 1984 is transferred to Stellularis Chen, Yao and Ren, 2015, leading to Stellulari senjituensis (Hong, 1984), comb. n.; Anthoscytina macula Hu, Yao and Ren, 2014 is transferred to Sinocercopis Hong, 1982, and Sunoscytinopteris (Scytinopteridae) and Cathaycixius (Cixiidae) are treated as junior homonym names of Sinocercopis, leading to Sinocercopis macula (Hu, Yao and Ren, 2014), comb. n., S lushangfenensis (Hong, 1984), comb. n., S pustulosis (Ren, 1995), comb. n., and S trinervis (Ren, 1995), comb. n. Additionally, two new species are erected: Stellularis bineuris Chen and Wang, sp. n. and S minutus Chen and Wang, sp. n. Our cladistic analysis based on wing (tegmen and hind wing) characteristics recovers the high-level relationships within Cercopoidea: Sinoalidae + (Procercopidae + (Cercopionidae + modern cercopoids)). Within the family Procercopidae, the cladistic analysis reveals that the Middle to Late Jurassic Titanocercopis and Jurocercopis and the Cretaceous Cretocercopis occupy the basal position, and a gradual change in wing venation can be recognized from the Early Jurassic Procercopis and Procercopina to the Jurassic Anthoscytina, and then to the Cretaceous Stellularis and Sinocercopis. The two Cretaceous genera, sharing wing traits with extant cercopoids, likely represent transitional forms between Procercopidae and recent Cercopoidea; however, they are very similar to their Jurassic relatives in body structures, suggesting it is applicable to attribute them to Procercopidae. Furthermore, our analysis suggests that the extinction of Procercopidae and the origin and early diversification of modern Cercopoidea approximately coincided with the rise and explosive radiation of angiosperms in the late Early Cretaceous and onwards.     

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 .     

Jurassic artematopodid beetles and their implications for the early evolution of Artematopodidae (Coleoptera)

Fossil Artematopodidae are rarely collected and previously confined to middle Eocene Baltic amber. Here we report the first definitive artematopodid, Sinobrevipogon jurassicus gen. et sp.n. , from the Middle Jurassic Daohugou beds (c. 165 Ma) in Inner Mongolia, northeastern China. It exhibits a number of defining features of Artematopodidae, including paired carinae on prosternum and an internal apical interlocking tongue on the ventral side of each elytron. However, it differs from any modern Artematopodidae by having the mesocoxal cavitiy closed by the mesepimeron and the anterolateral edge of metanepisternum. The discovery of this new genus represents the earliest fossil record for Artematopodidae, highlighting the antiquity of the family. The systematic positions of Forticatinius Tan & Ren and Tarsomegamerus Zhang are discussed, and the latter is formally transferred to Artematopodidae. Phylogenetic relationships within Artematopodidae were investigated to elucidate the relationships between the two Jurassic genera and Recent genera. Eleven in‐group taxa and two out‐groups were included in a cladistic analysis based on 30 adult characters; the resulting tree recovered the family Artematopodidae in three clades: (i) Electribius authority, (ii) Ctesibius authority + Brevipogon authority + Sinobrevipogon + Tarsomegamerus and (iii) the remaining Recent genera, including Allopogonia authority.     

New genera and species of Jumping Ground Bugs (Hemiptera: Schizopteridae) in Dominican and Burmese amber,with a description of a meloid (Coleoptera: Meloidae) triungulin on a Burmese specimen

New genera and species of jumping ground bugs (Hemiptera: Schizopteridae) are described in Dominican and Burmese amber. In Dominican amber are described: Hypselosoma dominicana Poinar & Brown sp. n., Schizoptera dominicana Poinar sp. n. and Schizoptera hispaniolae Poinar sp. n. In Burmese amber are described: Lumatibialis burmitis Poinar gen. et sp. n. and Hexaphlebia burmanica Poinar gen. et sp. n. A triungulin attached to the dorsum of the abdomen of Lumatibialis burmitis is described as Microentomus epibatus Poinar gen. et sp. n. in the family Meloidae (Coleoptera). This specimen represents the oldest known triungulin and the first fossil phoretic association of a triungulin and hemipteran.     

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 .     

Fossil flowers and fruits of the Actinidiaceae from the Campanian (Late Cretaceous) of Georgia

A new genus and species of Actinidiaceae (Parasaurauia allonensis gen. et sp. nov.) are established for fossil flowers and fruits from the early Campanian (Late Cretaceous) Buffalo Creek Member of the Gaillard Formation in central Georgia, USA. The fossil flowers, which are exquisitely preserved as charcoal, have five imbricate, quincuncially arranged sepals and petals. The androecium consists of ten stamens with anthers that are deeply sagittate proximally. The gynoecium is tricarpellate, syncarpous, and has three free styles that emerge from an apical depression in the ovary. The fruit is trilocular and contains numerous ovules on intruded axile placentae. The structure of mature fruits is unknown. Comparisons with extant taxa clearly demonstrate that the affinities of Parasaurauia allonensis are with the Ericales, and particularly with the Actinidiaceae, which have been placed among the Ericales in recent cladistic analyses. Because Parasaurauia allonensis is not identical to any one genus of Actinidiaceae, or other member of the Ericales, phylogenetic relationships of the fossil were evaluated through a cladistic analysis using morphological and anatomical characters. Results of this analysis place Parasaurauia allonensis within the Actinidiaceae as sister to the extant genera Saurauia and Actinidia. Parasaurauia allonensis differs from extant Saurauia only in having ten rather than numerous stamens.     

LUTETIACADER, A PUZZLING NEW GENUS OF CANTACADERID LACE BUGS (HETEROPTERA: TINGIDAE) FROM THE MIDDLE EOCENE MESSEL MAAR, GERMANY

Abstract:  A new genus of cantacaderid lace bugs, Lutetiacader gen. et sp. nov., is described from the lower Middle Eocene of Messel Quarry, Germany. It clearly belongs to the cantacaderid lineage of Tingidae, and is the second known fossil genus of the tribe Cantacaderini (after Paleocader Froeschner). It appears to be most similar to extant species of the genera Nectocader Drake (Brazil) and Teratocader Drake (Malay Peninsula). Lutetiacader occupies a central position with respect to the geographical gap between the ranges of these two extant genera. Its biogeographical significance is discussed.     

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 .     

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 .     

A review of the morphologically diverse leafhopper subfamily Stegelytrinae (Hemiptera: Cicadellidae) with description of new taxa

Abstract. The morphologically diverse leafhopper subfamily Stegelytrinae is reviewed and its biology and biogeography are discussed as well as its classification, monophyly and relationship to Deltocephalinae s.l. In addition, the first stegelytrine is recorded east of Wallace's and Weber's lines, extending the distribution of the subfamily outside the Oriental and Palaearctic regions. A checklist and key to the genera of the subfamily are given. The following new genera and species are described: Honguchia rubrofascia Wei & Webb, gen.n & sp.n. from Sabah; H. superba Wei & Webb, sp.n. from Sumatra; Yaontogonia flavopicta Wei & Webb, gen.n & sp.n. from Brunei and Sabah; Y. darjeelingensis Wei & Webb, sp. n. from West Bengal; Sychentia breviata Wei & Webb, gen.n & sp.n. from Malaysia; Louangana stellata Wei & Webb, gen.n & sp.n. from Laos; Shangonia sarawakensis Wei & Webb, gen.n & sp.n. from Sarawak; Quiontugia fuscomaculata Wei & Zhang, gen.n. & sp.n. from Hainan; and Neophansia wallacei Wei & Webb, gen.n & sp.n. from New Guinea.     

Ichneumonids of the subfamily Tanychorinae (Insecta: Hymenoptera: Ichneumonidae) from the Lower Cretaceous of Transbaikalia and Mongolia

A new species, Tanychora loki, sp. nov., and a new genus and species, Megachora sibirica gen. et sp. nov., are described from the Lower Cretaceous of Transbaikalia and Mongolia. The genus Tanychora Townes, 1973 is divided into two genera: Tanychora s. str., composed of T. petiolata Townes, 1973 (type species), T. sinensis Zhang, 1991, and T. loki sp. nov., and Amplicella, gen. nov., composed of A. sessilis (Townes, 1973) (type species), A. exquisite (Zhang et Rasnitsyn, 2003), A. beipiaoensis (Zhang et Rasnitsyn, 2003), and A. spinata (Zhang et Rasnitsyn, 2003). The frequency of occurrence of ichneumonids in the principal Cretaceous localities of central and northern Asia is calculated.     

Diverse oxyporine rove beetles from the Early Cretaceous of China (Coleoptera: Staphylinidae)

Two new genera and species, Protoxyporus grandis gen. et sp.n. and Cretoxyporus extraneus gen. et sp.n. , are described and figured on the basis of two well‐preserved impression fossils from the Lower Cretaceous Yixian Formation of China. The two genera display a combination of traits that are diagnostic of the extant staphylinid subfamily Oxyporinae: large and prominent mandibles, distinctly separated mesocoxae, abdominal tergites III–VII each with a pair of basolateral ridges, and abdominal intersegmental membranes with ‘brick‐wall’ pattern. Protoxyporus differs from extant Oxyporus in retaining the following plesiomorphic features: an elongate intercoxal process (or longitudinal carina) between metacoxae, relatively narrowly separated mesocoxae, less developed metaventral anterior process, and long infraorbital ridges. Cretoxyporus is morphologically very similar to Oxyporus, but retains distinct elongate infraorbital ridges. The discovery of two new oxyporine genera suggests that the early oxyporines were more diverse and their evolutionary history more complicated than previously documented. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:11503976‐D577‐45D2‐ACCB‐8226782EF817 .     

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 .     

New and little known phalangopsinae (Orthoptera,Gryllidae): 7. Neotropical taxa of the tribes Paragryllini and Luzarini

Three new genera, one new subgenus, and 13 new species (Strogulomorpha separata sp. n., S. davidi sp. n., S. proxima sp. n., Luzarida lata sp. n., Luzara venado sp. n., L. sapani sp. n., Ucayacla pulchella gen. et sp. n., Peruzara atalaya gen. et sp. n., P. loreto sp. n., Amazonacla imitata gen. et sp. n., A. primitiva sp. n., Leptopsis (Leptopsis) ucayali sp. n., and L. (Aberracla subgen. n.) morona sp. n.) are described. The composition of the tribes Paragryllini and Luzarini and of the genera Luzarida Heb. and Luzara Walk is discussed. The new synonymies Luzarida Hebard, 1928 = Ecuazarida Nischk et Otte, 2000, syn. n.; Lerneca Walker, 1869 = Doposia Otte et Perez-Gelabert, 2009, syn. n.; Nemoricantor Desutter-Grandcolas et Hubbell, 1993 = Kumalorina Otte et Perez-Gelabert, 2009, syn. n. are given. The systematic position of some previously described species is corrected.     

Eocene tortoise beetles from the Green River Formation in Colorado,U.S.A. (Coleoptera: Chrysomelidae: Cassidinae)

Abstract The fossil history of leaf beetles (Chrysomelidae) is relatively poorly documented despite an abundance of available material. Of particular interest is the origin and radiation of the diverse tortoise beetles, a derived group within Cassidinae s.l. (=Cassidinae + Hispinae) defined by the exophagous life history and specialized morphology of the immature stages. Cassidinae is also a group with relatively few fossil records that can be assigned with any degree of certainty. Here we report two of the oldest definitive tortoise beetle fossils, Eosacantha delocranioides gen.n. et sp.n. and Denaeaspis chelonopsis gen.n. et sp.n. , from the Eocene Green River Formation (ca. 47 million years old) in northwestern Colorado, U.S.A. Owing to the fine level of preservation, many important features can be observed and are coded into the recent cladistic analysis for the subfamily. Phylogenetic analysis highlights that both genera have affinities with modern lineages, one restricted to the Old World and the other restricted to the Neotropics. Although Cassidinae as a whole extend into the Cretaceous, the available information suggests that the tortoise beetles perhaps originated and diversified during the Early Tertiary. As such, the morphological and biological transitions from the leaf‐mining hispiforms to the distinctive tortoise‐like cassidiforms, with their elaborate defensive larval shields and other unique behaviours, probably took place during the latest Paleocene or earliest Eocene. These Green River fossils are the oldest yet to document the specialized morphology associated with the transition in cassidine feeding and immature biology.     

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 .