New polymorphic psyllipsocids from Burmese amber (Psocodea: Psyllipsocidae)

Annulipsyllipsocus andreneli gen. et sp. nov. and A. inexspectatus gen. et sp. nov., the first records of the Psyllipsocidae from the Cretaceous amber of Myanmar are described, illustrated and their taxonomic position discussed. The new genus is characterized by antennae with 16 segments; flagellomeres with secondary annulations; radial cell closed, five-angled, bulged on its R1 border; triangular pterostigma. Species belonging to Annulipsyllipsocus gen. nov. are polymorphic. The described taxa represent the earliest evidence for polymorphism in Psocodea. A checklist of all known species of Psyllipsocidae is given.     

Extreme diversity of decapod crustaceans from the mid-Cretaceous (late Albian) of Spain: Implications for Cretaceous decapod paleoecology

Thirteen decapod crustacean species, eight of which are new, from the mid-Cretaceous (late Albian) limestones of the Koskobilo quarry in northern Spain are described, illustrated, and discussed. They include: Graptocarcinus texanus; Navarrara betsieae gen. nov., sp. nov.; Acareprosopon bouvieri n. comb.; Laeviprosopon hispanicum sp. nov.; L. planum sp. nov.; L. edoi sp. nov.; L. crassum sp. nov.; Viaia robusta; Cretamaja granulata gen. nov., sp. nov.; Koskobilius postangustus gen. nov., sp. nov.; Navarrahomola hispanica; Glytodynomene alsasuensis; and Albenizus minutus gen. nov., sp. nov. Cretamaja and Koskobilius represent the oldest known spider crabs. In total, 36 species are now known from the Koskobilo locality based on 1078 specimens. To discover the magnitude of the diversity of the decapod fauna of Koskobilo, a comprehensive overview of decapod-rich localities and formations from the Cretaceous worldwide was compiled. It appears that Koskobilo is the most diverse decapod fauna from a single locality currently known from the Cretaceous. A rarefaction analysis shows that the maximum number of species is nearly reached. The number of genera, 26, is also unsurpassed for the Cretaceous. Forty-two species are found from localities within the Eguino Formation to which sediments from the Koskobilo quarry are ascribed, which is also unprecedented for a single formation within the Cretaceous. Evidence suggests that the most diverse decapod faunas from the Cretaceous are found in coral-associated limestones. This is consistent with evidence from the Recent, where decapod diversity is high in coral reefs compared to other habitats. This also suggests that the decapod peak diversity in Koskobilo is largely ecological in nature and not caused by a preservational bias. This is one of the most comprehensive studies on Cretaceous decapod diversity so far. Field work in coral-associated strata is expected to yield more decapod-rich faunas.     

Diverse stigmaphronid wasps in Early Cretaceous amber from Spain (Hymenoptera: Ceraphronoidea: Stigmaphronidae)

A diverse fauna of wasps of the extinct parasitoid family Stigmaphronidae (Ceraphronoidea) are recorded in Early Cretaceous (Lower Albian) amber from Spain. Seven new species in five genera are described and figured based on 51 specimens, representing more material than in all the world’s other amber deposits combined. New species include: Elasmophron mari sp. nov., Libanophron sugaar sp. nov., Hippocoon basajauni sp. nov., Burmaphron jentilak sp. nov., B. sorginak sp. nov., B. iratxoak sp. nov., and Tagsmiphron olentzero sp. nov. The significance of the fauna is discussed and compared with that of other Cretaceous amber deposits, in particular the tremendous richness of the Spanish fauna is contrasted with the complete absence of stigmaphronids in the slightly younger and nearby French amber. Whether this stark difference represents particularly favorable conditions for these parasitoids, or their hosts, in the Cretaceous Spanish archipelago, or whether it is owing to taphonomic factors is discussed.     

Brochocolein beetles (Insecta: Coleoptera) from the Lower Cretaceous of northeast China and southern England

Lower Cretaceous beetles belonging to the extinct tribe Brochocoleini (Coleoptera: Cupedidae: Ommatinae) are revised and five new species proposed in the extinct Eurasian genus Brochocoleus: Brochocoleus yangshuwanziensis sp. nov. (Yixian Formation, China); Brochocoleus crowsonae sp. nov., Brochocoleus keenani sp. nov., Brochocoleus tobini sp. nov. (Weald Clay Group, UK); and Brochocoleus maximus sp. nov. (Purbeck Limestone Group, UK). New morphological data is provided for Brochocoleus impressus and Brochocoleus sulcatus stat. nov. of which Brochocoleus angustus is considered a synonym (Yixian Fm.). The hindwing venation is described for the first time supporting the basal position of this tribe. A simple key is provided to all Cretaceous species of this presumably xylophagous, warm climate genus.     

New taxa of unusual Dilaridae (Neuroptera) with siphonate mouthparts from the mid-Cretaceous Burmese amber

Cretadilar olei gen. et sp. nov. and Burmopsychops groehni sp. nov. (Neuroptera: Dilaridae) are described from the Late Cretaceous (earliest Cenomanian/late Albian) Burmese amber. Their dilarid affinities are supported by two autapomorphies of the family: the reduction of the ligula and the presence of pectinate antennae in males. They are considered to belong to two new subfamilies, Cretanallachiinae subfam. nov. and Cretadilarinae subfam. nov., respectively. These subfamilies are easily distinguished from other dilarids (Dilarinae, Nallachiinae, and Berothellinae) by possessing siphonate mouthparts; however they strongly differ from each other. In particular, the broad gonocoxites 9 of Cretanallachiinae females are not fused to form a distinct ovipositor, and bear long gonostyli, whereas a distinct saber-like ovipositor (short, lacking gonostyli) is present in Cretadilarinae. It is assumed that Cretanallachiinae primarily fed on pollen and Cretadilarinae on the nectar of flowers.     

New Stigmaphronidae and Megaspilidae (Hymenoptera: Ceraphronoidea) from Canadian Cretaceous amber

Three new species within the stigmaphronid genus Tagsmiphron Engel and Grimaldi, 2009, and one new species within the megaspilid genus Conostigmus Dahlbom, 1858 are described from Upper Cretaceous (Campanian) amber originating at the Grassy Lake locality in Alberta, Canada. New taxa include Tagsmiphron spiculum sp. nov., Tagsmiphron leucki sp. nov., Tagsmiphron exitorum sp. nov., and Conostigmus cavannus sp. nov. The new Conostigmus species is a rare discovery. It is the third megaspilid species to be found in Cretaceous amber, with the two specimens described herein effectively doubling the number of known Mesozoic exemplars for the family. We provide the first comprehensive report of known Ceraphronoidea within Canadian amber, and contrast this against other Cretaceous amber assemblages, discussing the potential palaeobiogeographic and palaeoenvironmental implications of the Canadian amber assemblage.     

Archegocimicidae (Insecta: Heteroptera) from the Purbeck Limestone Group (Lower Cretaceous: Berriasian) of southern England

Four new genera and five new species of Archegocimicidae are described from the Lower Cretaceous of England: namely Mortalia martini gen. et sp. nov., Tyrion lannister gen. et sp. nov., T. cersei sp. nov., Stannis baratheon gen et sp. nov., Daenerys khaleesi gen. et sp. nov. A transitional position of the complex between Asian Late Jurassic and Early Cretaceous entomofaunas is indicated by this new material.     

The earliest Attagenus species (Coleoptera: Dermestidae: Attageninae) from Upper Cretaceous Burmese amber

A new species belonging to the extant dermestid genus Attagenus, Attagenus burmiticus sp. nov., is described and illustrated based on a well-preserved specimen from the Upper Cretaceous Burmese amber. The discovery suggests that Attagenus is an ancient group, originating as early as in the mid-Cretaceous. Along with another species of Attagenus known from the Upper Cretaceous New Jersey amber, it implies that Attagenus were widespread in the Mesoozic.     

New Cretaceous Elcanidae from China and Myanmar (Insecta,Orthoptera)

A new species of Elcanidae (Orthoptera: Elcanoidea), Panorpidium yixianensis sp. nov., is described based on two new specimens from the Lower Cretaceous Yixian Formation of northeastern China. It differs from other species in forewing characters and spines on the hind tibiae. In addition, a new specimen Burmelcana sp., is described and figured based on an amber inclusion from the mid-Cretaceous Burmese (Myanmar) amber. P. yixianensis sp. nov. represents the first definite record of Elcanidae in the Early Cretaceous Jehol biota, and Panorpidium is the only genus of Ensifera to be found in the Early Cretaceous faunas of England, Russia and China.     

The first Mesozoic palmetto beetle (Coleoptera: Smicripidae) in Upper Cretaceous Burmese amber

Fossil smicripid beetles are very rare. Here we describe and figure the first Mesozoic representative of Smicripidae. Smicrips cretacea sp. nov., a new species belonging to the extant Smicrips LeConte is preserved in the Upper Cretaceous amber from northern Myanmar. Our discovery suggests that Smicrips is an ancient group, originating at least in the earliest Late Cretaceous. Together with other two fossil Smicrips species from the Eocene of Europe, it reinforces the hypothesis that the Recent distribution of Smicripidae is relict, and the family was more widespread in the Mesozoic and early Cenozoic than it is in the Recent.     

New fossil mesosciophilids (Diptera: Nematocera) from the Yixian Formation (Lower Cretaceous) of western Liaoning,China

A new genus with two new species, Orentalphila gravia gen. et sp. nov. and O. caloa sp. nov., and three new species in two known genera, Sinosciophila angustia sp. nov., Sinosciophila seboa sp. nov. and Similsciophila undulata sp. nov., are described from the Lower Cretaceous of Yixian Formation at Huangbanjigou, Beipiao City, western Liaoning, China. These new taxa, representing the first records of mesosciophilids from this locality, provide morphological information and diversity for Mesosciophilidae in the Early Cretaceous. An updated key to known genera of Mesosciophilidae is provided. In addition, according the generic diagnosis revised by Zhang, 2007, Mesoplecia antiqua Hao and Ren, 2009 should be transferred to Mesosciophila Rohdendorf, 1946.     

The first Mesozoic pleasing lacewing (Neuroptera: Dilaridae)

Cretanallachius magnificus gen. et sp. nov., first Mesozoic and earliest record of the Dilaridae (Neuroptera), is described from the Cretaceous Burmese amber. Its putative closest relative is the recent subfamily Nallachiinae known by the sole genus Nallachius.     

Biting midges of the extinct genus Protoculicoides Boesel from Lower Cretaceous amber of San Just,Spain and new synonymy in recently described fossil genera (Diptera: Ceratopogonidae)

We describe two new species of extinct biting midges in the fossil genus Protoculicoides: Protoculicoides hispanicus Szadziewski and Arillo, sp. nov. and Protoculicodes sanjusti Szadziewski and Arillo, sp. nov., from Albian (101–113 Ma) amber of San Just, Spain. Atriculicoides Remm, 1976 is recognized as a new junior synonym of Protoculicoides Boesel, 1937 (NEW SYNONYM). Devalquia Choufani et al., 2013 is regarded here as a new junior synonym of the extant genus Culicoides Latreille, 1809 (NEW SYNONYM). Metahelea roggeroi Choufani et al., 2013 from Upper Cretaceous French amber, is excluded from the tribe Heteromyiini and placed in the extant genus Stilobezzia Kieffer, 1911, tribe Ceratopogonini: Stilobezzia roggeroi (Choufani et al., 2013), comb. nov. The extinct tribe Atriculicoidini Szadziewski, 1996 is regarded as a new subfamily, Atriculicoidinae, stat. nov., that includes the fossil genus Protoculicoides, with 13 species reported from Albian, Turonian, Cenomanian, Coniacian, Santonian and Campanian ambers (78–113 Ma). Protoculicoides skalskii Szadziewski & Arillo, 1998 from Lower Cretaceous amber of Álava, Spain, Protoculicoides succineus Szadziewski, 1966 from Lower Cretaceous Lebanese amber and Protoculicoides burmiticus Szadziewski & Poinar, 2005 from Upper Cretaceous Burmese amber are transferred to the extinct genus Archiculicoides Szadziewski, 1996, comb. nov. Protoculicoides krzeminskii Choufani et al., 2014 from Lower Cretaceous Lebanese amber is transferred to the extinct genus Archiaustroconops Szadziewski, 1996, comb. nov. The Cretaceous subfamily Atriculicoidinae forms an unresolved trichotomy with the extant subfamilies Forcipomyiinae and Dasyheleinae, both of which date to the Eocene. The fossil record indicates that wings with macrotrichia in biting midges evolved during the mid-Cretaceous greenhouse climate. We suggest that this was an evolutionary adaptation to new atmospheric conditions with higher levels of CO₂ in order for the antennal Johnston's organs of males to receive the vibrational sex signals produced by females during flight.     

A new earwig (Dermaptera: Pygidicranidae) from the Upper Cretaceous Myanmar amber

A new genus and species, Gracilipygia canaliculata gen. et sp. nov., in the family Pygidicranidae of Dermaptera is described from the Upper Cretaceous Myanmar amber. The new species is assigned to the subfamily Pyragrinae mainly based on the following characters: broadly separated cercal forceps without segmentation and the posterior abdominal tergum with mediolongitudinal furrow and lateral ridges. The G. canaliculata gen. et sp. nov. is the earliest record hitherto for the subfamily Pyragrinae, and probably represents a stem taxa of Pyragrinae.     

Non-marine gastropods from the Cretaceous–Paleogene transition in the Pingyi Basin,eastern China

Our results present a taxonomic and palaeoecological study on non-marine gastropods from the latest Cretaceous to Paleocene deposits of the Pingyi Basin, Shandong Province, eastern China. These gastropods are systematically described: three species belonging to three genera including Physa dongtaiensis Gu, 1989, Hydrobia datangensis Yü, 1977, and the newly established species Cyathopoma pingyiensis sp. nov.; two indeterminata genera and species including Truncatelloidea gen. et sp. indet., and Pomatiopsidae gen. et sp. indet. Among them, Truncatelloidea gen. et sp. indet. and Cyathopoma pingyiensis sp. nov. are the dominant species with the longest record. Cyathopoma pingyiensis, sp. nov. is the earliest representative of this widely distributed Asian extant genus. Sedimentological facies analysis of the gastropod-bearing beds suggested that Truncatelloidea gen. et sp. indet. lived in a small pond with a river inlet and a shallow lake, while Ph. dongtaiensis, ?Pomatiopsidae gen. et sp. indet, and ?H. datangensis only thrived in the shallow lake. Cyathopoma pingyiensis sp. nov. inhabited the land area around the shallow lake. Our results showed that no significant species change of the gastropod fauna across the K/Pg (Cretaceous/Paleogene) boundary was observed in the Pingyi Basin.     

A golden age for ectoparasitoids of Embiodea: Cretaceous Sclerogibbidae (Hymenoptera,Chrysidoidea) from Kachin (Myanmar), Charentes (France) and Choshi (Japan) ambers

Sclerogibbid wasps are obligate parasitoids of webspinners (Embiodea). Both groups have a particularly scarce geological record and are known since the Cretaceous: there are only four species of webspinners known from Burmese amber, and only two sclerogibbids were described from Barremian Lebanese and Cenomanian Burmese ambers. Here we report transferred genus from Aptian Choshi (Japan) amber and new sclerogibbids from Cenomanian Burmese and Charentese (France) ambers. The taxa described from Burmese amber are: Burmasclerogibba aptera gen. et sp. nov., Cretosclerogibba gen. nov. (with C. antennalis sp. nov., C. contractocollis sp. nov., C. neli sp. nov. and C. rasnitsyni sp. nov.) and Edrossia vetusta gen. et sp. nov. The first European fossil sclerogibbid Gallosclerogibba alnensis gen. et sp. nov. is described from Charentese amber. The holotype of Chosia yamadai Fujiyama, from Choshi amber, is re-described; it appears to be the oldest Laurasian sclerogibbid. The significant abundance and variety of Burmese sclerogibbid wasps (60% of fossil species known worldwide), as proxy of their hosts, were probably caused by the protection granted to them by the silk webs and possibly by the limited predation from ornithuromorph birds or crown-group ants. While all three extant sclerogibbid genera have apterous females, genera with winged females (Cretosclerogibba and Edrossia) dominated in Burmese amber. Small silk galleries from hosts may have favored the preservation of wings in females of Cretaceous sclerogibbids. Most new species described in the present paper, in addition to C. yamadai, are characterized by a very slender neck and a very long frontal process concealing the antennal toruli. These characters disappeared in extant species. We suggest that this loss was caused by a change in the fauna of predators, penalizing species with long neck and rostrum.     

The first record of Ichneumonidae (Insecta: Hymenoptera) from the Upper Cretaceous of Myanmar

A new subfamily of Ichneumonidae, Novichneumoninae subfam. nov., is established based on two new genera with two new species: Novichneumon longus gen. et sp. nov. and Caloichneumon perrarus gen. et sp. nov. These two new species are the first ichneumonids described from the Upper Cretaceous Myanmar (Burmese) amber. A list of all described Mesozoic ichneumonid fossil species with their respective localities and ages is summarized. The distribution of Ichneumonoidea during the Cretaceous indicates that Cretaceous ichneumonids were documented from localities at high latitudes while braconids were distributed worldwide, a pattern consistent with the distribution of extant ichneumonids and braconids.     

New lacewings (Insecta: Neuroptera) from the Lower Cretaceous Wealden supergroup of Southern England

Eight new genera and thirteen new species of lacewings (Neuroptera) are described from the Lower Cretaceous Wealden Supergroup, Weald Sub-basin: Principiala rudgwickensis sp. nov. (Ithonidae), Stenomylina medialis gen. et sp. nov., Protosmylina bifasciata gen. et sp. nov. (Osmylidae), Cretapsychops corami gen. et sp. nov., Micropsychops parallelus gen. et sp. nov., Valdipsychops logunovi gen. et sp. nov., V. brigidae gen. et sp. nov., V. maculosus gen. et sp. nov., V. proudlovei gen. et sp. nov., V. minimus gen. et sp. nov., Psychopsites rolandi gen. et sp. nov. (Psychopsidae), Actinophlebioides valdensis gen. et sp. nov. and Protohemerobius perexiguus gen. et sp. nov. (Neuroptera familia incertae sedis). Psychopsidae are shown to dominate the neuropteran fauna, while the presence of Brongniartiellidae has not been confirmed in the Wealden, due to previously thought representatives of this family being in fact psychopsids. This is the first fossil record of Ithonidae and Osmylidae in Britain. The range of Kalligrammatidae has been extended in the Cretaceous of Britain from the mid-Valanginian to early Barremian.     

New pelecinid wasps (Hymenoptera: Pelecinidae) from Upper Cretaceous Myanmar amber

A new genus with a new species, Brachypelecinus euthyntus gen. et sp. nov., and two new species, Abropelecinus tytthus sp. nov. and Zoropelecinus periosus sp. nov., are described and figured from three exquisitely preserved pelecinid wasps in the Upper Cretaceous Myanmar (Burma) amber. These taxa, owing to their well-preserved characters in amber, provide a better understanding of morphological changes and relationships among the constituent groups, while further highlighting the diversity of Pelecinidae in the Mesozoic.     

Two new genera of belytine wasps (Hymenoptera: Diapriidae) from Cretaceous Burmese amber

Christophus myanmarensis gen. et sp. nov., Christophus ohmkuhnlei gen. et sp. nov., and Mintara parva gen. et sp. nov., three new species and two new genera of belytine wasps (Diapriidae: Belytinae) are described and figured from males preserved in mid-Cretaceous Kachin (Burmese) amber. These new fossils enrich the fossil record of Cretaceous diapriids, summarized here, demonstrating the diversity of the family before the K/Pg extinction and suggesting an older origin for the family.