A new genus and species of pygidicranid earwigs from the Upper Cretaceous of southern Asia (Dermaptera: Pygidicranidae)

A third genus and species of fossil pygidicranid earwigs from Burmese amber is described and figured as Stonychopygia leptocerca Engel, Huang, Thomas, and Cai, gen. et sp. nov. (type species of Stonychopygia Engel and Huang, gen. nov.; Stonychopygiinae Engel and Huang, subfam. nov.). Stonychopygia have features similar to the subfamily Echinosomatinae such as the combination of shorter, subequal second and third flagellomeres and femora that are neither compressed or keeled. However, the new fossil species differs from echinosomatines in many details, particularly the slender form, head longer than wide, slender scape, slender and elongate forceps, and absence of stout, short bristles over the integument. The new fossil is compared with its contemporaneous and modern relatives, and comments are provided regarding the classification of Pygidicranoidea, with the genus Haplodiplatys Hincks removed from Diplatyidae to Haplodiplatyidae Engel, fam. nov.     

A remarkable evanioid wasp in mid-Cretaceous amber from northern Myanmar (Hymenoptera: Evanioidea)

A remarkable new wasp of the superfamily Evanioidea (Apocrita: Evaniomorpha) is described and figured from a female preserved in mid-Cretaceous amber from the Hukawng Valley of northern Myanmar. Othniodellitha mantichora Engel and Huang, gen. et sp. nov., is characterized by its unique combination of primitive and derived features in mesosomal and metasomal construction and wing venation, along with its peculiarly blocky head with a prominent facial horn, clypeal projection, and massive, squared mandibles, among other traits. The genus is placed in the new family, Othniodellithidae Engel and Huang, fam. nov., and is among the lower Evanioidea and distinguished from the Praeaulacidae and other Mesozoic evanioid families.     

Two new species of mid-Cretaceous webspinners in amber from northern Myanmar (Embiodea: Clothodidae,Oligotomidae)

Two new genera and species of Cretaceous webspinners (Embiodea) are described and figured, both preserved in amber from northern Myanmar. Atmetoclothoda orthotenes Engel and Huang, gen. et sp. nov., is the first fossil representative of the putatively primitive family Clothodidae, and is segregated into its own subfamily, Atmetoclothodinae Engel and Huang, subfam. nov., owing to its primitive retention of a distinct mentum, a quadrate submentum with straight borders, a postocciptal suture that meets the hypostomal sulcus at the posterior tentorial pit, and subgenae that do not meet medially (thus a ventral bridge or gula is lacking), completely symmetrical terminalia, and unsegmented cerci that apically bear a small inner lamellar projection. Litoclostes delicatus Engel and Huang, gen. et sp. nov., is the first definitive fossil of Oligotomidae (not including a subfossil species in Pleistocene copal and which is likely synonymous with an extant taxon), and differs from modern genera in a combination of head, tarsal, and terminalic morphology. The new fossil species double the known Cretaceous fossils and add a further two families to the Mesozoic record for the order.     

A plesiomorphic gasteruptiid wasp in Cenomanian amber from Myanmar (Hymenoptera: Gasteruptiidae)

The first definitive Burmese amber fossil of the family Gasteruptiidae s.str. (Evanioidea) is described and figured from a male entombed in amber from the Hukawng Valley, Myanmar. The Cenomanian-aged fossil is plesiomorphic in many respects when compared to the modern subfamilies Hyptiogastrinae and Gasteruptiinae. The genus Hypselogastrion Engel, gen. nov. (type species: Hypselogastrion simplex Engel and Wang, sp. nov.), is segregated into the extinct subfamily Hypselogastriinae Engel, subfam. nov., owing the more enriched wing venation, aulcid-like mesoscutal sculpturing, non-clavate metatibia, and absence of U-shaped notauli. The affinities of H. simplex among other living and fossil Aulaciformes are discussed.     

An unusual new lineage of sawflies (Hymenoptera) in Upper Cretaceous amber from northern Myanmar

A peculiar new lineage of sawflies (‘Symphyta’) is described and figured from a female beautifully preserved in Upper Cretaceous (Cenomanian) amber from northern Myanmar. Syspastoxyela rhaphidia Engel and Huang, gen. et sp. nov., shares many plesiomorphic features with the primitive Xyelidae, 2Xyelotomidae, and 2Xyelydidae such as enlarged and thickened first flagellomere succeeded by a series of thinner and shorter flagellomeres, absence of a transverse mesoscutal sulcus, multiple preapical spurs, and two protibial spurs among other traits. However, the new lineage has an apomorphically contracted forewing venation, lacks a subcostal vein, has a single marginal cell, and lacks crossvein 1r-rs, and thus it is segregated into a new family, Syspastoxyelidae Engel and Huang, fam. nov. The phylogenetic affinities of the new family are discussed and a position near Pamphilioidea or Pamphilioidea + Unicalcarida is advocated.     

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.     

A new species of Glaesoconis Meinander (Neuroptera: Coniopterygidae) from the Santonian Taimyr amber

Glaesoconis popovi sp. nov. (Neuroptera: Coniopterygidae) is described from Upper Cretaceous (Santonian) Taimyr amber of northern Siberia (Yantardakh locality). The new species may be distinguished from others in the genus by much smaller eyes and the shape of the terminal segment of the maxillary palpus. The generic affinity of Glaesoconis baliopteryx Engel, 2004 from the mid-Cretaceous Burmese amber needs confirmation.     

A thorny, ‘anareolate’ stick-insect (Phasmatidae s.l.) in Upper Cretaceous amber from Myanmar,with remarks on diversification times among Phasmatodea

A new fossil stick-insect (Phasmatodea) is described and figured from a male preserved in Cretaceous amber from Myanmar. Echinosomiscus primoticus Engel and Wang, gen. et sp. nov., is a robust, somewhat-compressed stick-insect bearing abundant acanthae over the head and body, and remarkably lacks an area apicalis on the tibiae. The species is described and assigned to a new, extinct subfamily of Phasmatidae s.l., as Echinosomiscinae Engel, subfam. nov. Brief remarks are made regarding the dating of phasmatodean lineages, with E. primoticus providing the first reliable evidence for Euphasmatodea and even Neophasmatodea in the Cenomanian.     

New Prionoglarididae from Burmese amber (Psocodea: Trogiomorpha: Prionoglaridetae)

One new genus with three new species of the family Prionoglarididae, Palaeosiamoglaris leinhardi Azar, Huang et Nel gen. et sp. nov.; P. burmica Azar, Huang et Nel sp. nov. and P. inexpectata Azar, Huang et Nel sp. nov. are characterised, illustrated and described from Cretaceous Burmese amber. Palaeosiamoglaris gen. nov. share several characters with the recent genus Siamoglaris Lienhard, 2004, endemic to the Oriental (Sino-Indian) Region. A new tribe Siamoglaridini Azar, Huang et Nel trib. nov. is established to accommodate the group Siamoglaris + Palaeosiamoglaris; the other Prionoglaridinae being attributed to Prionoglaridini. This discovery implicates a palaeobiogeographic repartition for this particular tribe (Siamoglarini) that is still the same currently. Most diagnostic features show a nearly stasis state, supporting the notion that these characters appeared at least by the early Cenomanian. A checklist of all prionoglaridids is given.     

A new damselfly (Odonata: Zygoptera: Platystictidae) from mid-Cretaceous Burmese amber

The genus Mesosticta Huang, Azar, Cai et Nel, 2015 was established based on the wing bases of two damselflies from mid-Cretaceous Burmese amber. Here we describe a new well-preserved platystictid damselfly, Mesosticta electronica sp. nov., with complete forewings and hindwings. The diagnosis of Mesosticta is revised and augmented in this paper. Mesosticta electronica sp. nov. differs from Mesosticta burmatica Huang, Azar, Cai et Nel, 2015 in having the arculus slightly distal of Ax2, a free subdiscoidal cell, the hindwing AA ending on the middle area of the posterior side of the discoidal cell, and the base of RP2 being three or four cells distal of the subnodus. The new discovery adds to the diversity of damselflies in mid-Cretaceous Burmese amber and puts the origin of Platystictidae to at least the mid-Cretaceous.     

A second species of the genus Cretoleptochromus Cai & Huang (Coleoptera: Staphylinidae: Scydmaeninae) from mid-Cretaceous Burmese amber

A second species of the extinct scydmaenine genus Cretoleptochromus Cai & Huang, C. burmiticus sp. nov., is described and figured based on three exquisitely preserved specimens embedded in mid-Cretaceous Burmese amber. The new taxon possesses an extremely elongate body form and strongly extended legs and antennae that provide reliable means for species diagnosis and a ready separation from the only known congener, the type species C. archaicus Cai & Huang, also reported from Burmese amber. The morphology of C. burmiticus also suggests that this species was probably an active diurnal predator living in open environments.     

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

Three new species within the stigmaphronid genus TagsmiphronEngel and Grimaldi, 2009, and one new species within the megaspilid genus ConostigmusDahlbom, 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.     

A new earwigfly from mid-Cretaceous Burmese amber (Mecoptera: Meropeidae)

A new species of Meropeidae (earwigfly) is described and figured based on an exceptionally well-preserved individual in mid-Cretaceous amber from Myanmar. Burmomerope clara Zhao and Wang, sp. nov. is distinguished from the type species B. eureka Grimaldi and Engel, 2013 by presence of broader wings with six longitudinal veins in radial sector and seven in medial field, CuA with two terminal branches, and long setae on the anterior margin of the wing. A detailed comparison of the forewings venation in all fossil and extant species is given. The new find is the third fossil species of Meropeidae and also the first fossil female to be described. The female genital structure of B. clara sp. nov. is remarkably similar to that of extant species, revealing 100 million years of morphological conservatism, and thus highlighting the antiquity of the group.     

New Cretaceous caddisflies from Burmese amber (Insecta,Trichoptera)

Three new caddisflies species are described and illustrated from the mid-Cretaceous Burmese amber: Wormaldia cretacea sp. nov., W. resina sp. nov. (Philopotamidae) and Neureclipsis burmanica sp. nov. (Polycentropodidae). Palerasnitsynus ohlhoffi (Psychomyiidae) is re-described based on new fossils. Wormaldia are common in Burmese amber, and its diversity shows that this genus had very ancient origin and diversified at least during the mid-Cretaceous. N. burmanica sp. nov. is the oldest record of Neureclipsis, revealing this genus originated at least in the mid-Cretaceous. So far, six extinct species representing the small Order Trichoptera have been found in Burmese amber.     

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 Oriental protorhyssaline wasp (Hymenoptera: Braconidae): A new genus and species in Upper Cretaceous amber from Myanmar

The exclusively Cretaceous braconid wasp subfamily Protorhyssalinae is reported from the Upper Cretaceous deposits of northern Myanmar. Archaeorhyssalus subsolanus Engel, gen. et sp. nov., is represented by a well-preserved female in Burmese amber and is similar to other protorhyssalines, but differs in numerous details of forewing venation and flagellomere number. This species is the first definitive Oriental occurrence for protorhyssalines, expanding their palaeogeographic distribution from the Lower Cretaceous of Spain, and the Upper Cretaceous of France and New Jersey, USA. A key to the known genera of protorhyssalines is provided.     

New caddisflies species (Insecta: Trichoptera) from the Cretaceous Taymyr amber

Four new Trichoptera species: Kliganigadukia taymyrensis gen. et sp. nov. (Hydrobiosidae), Archaeopolycentra yantardakh sp. nov. (Polycentropodidae), Taymyrodipseudon protopegasus gen. et sp. nov. (Dipseudopsidae), and Siberoclea parapolaria gen. et sp. nov. (Leptoceridae) from Late Cretaceous amber (Santonian, Kheta Formation, 85 Ma) of Taymyr (Siberia, Russian Federation) are described and illustrated. Data on the Cretaceous amber Trichoptera (13 families, 20 genera, 29 species) are summarized and discussed.     

New species of fossil oribatid mites (Acariformes,Oribatida), from the Lower Cretaceous amber of Spain

Mites are relatively common and diverse in fossiliferous ambers, but remain essentially unstudied. Here, we report on five new oribatid fossil species from Lower Cretaceous Spanish amber, including representatives of three superfamilies, and five families of the Oribatida. Hypovertex hispanicus sp. nov. and Tenuelamellarea estefaniae sp. nov. are described from amber pieces discovered in the San Just outcrop (Teruel Province). This is the first time fossil oribatid mites have been discovered in the El Soplao outcrop (Cantabria Province) and, here, we describe the following new species: Afronothrus ornosae sp. nov., Nothrus vazquezae sp. nov., and Platyliodes sellnicki sp. nov. The taxa are discussed in relation to other fossil lineages of Oribatida as well as in relation to their modern counterparts. Some of the inclusions were imaged using confocal laser scanning microscopy, demonstrating the potential of this technique for studying fossil mites in amber. A table, including all the known Mesozoic oribatid mites preserved in amber, is included.     

A look with μCT technology into a treasure trove of fossils: The first two fossils of the millipede order Siphoniulida discovered in Cretaceous Burmese amber (Myriapoda,Diplopoda)

Millipedes (Diplopoda) are an important fossil group of land arthropods in the Palaeozoic. However, there is a gap in the Mesozoic, with only slightly more than a dozen fossils being known, until the much more recent fossil records mainly from Cenozoic Dominican and Baltic ambers become available. Burmese amber, precisely dated to the Late Cretaceous, is known for an apparently rich but still undescribed millipede fauna, and might represent the ideal bridge to close this gap in the millipede fossil record. In addition, modern micro-computed tomography (μCT) technology allows complex 3D reconstructions of objects fossilized in amber. Here we utilize μCT technology to describe the first millipedes from Burmese amber, which are also the first two fossil representatives of the order Siphoniulida. Siphoniulida, with two known species and just nine recorded specimens, are the rarest and least known of the 16 extant orders of the Diplopoda. Records are known from Sumatra, Guatemala and Mexico. The two new fossils described here represent two distinct species, Siphoniulus muelleri sp. nov. and S. preciosus sp. nov., and indicate a wider distribution of this order in the Cretaceous. The holotype of S. muelleri sp. nov. was well-enough preserved so that characters of the head, such as the incisura lateralis, not investigated before in extant representatives of the order, could be studied. This study highlights the possibilities and challenges provided by μCT technology in investigating Diplopoda fossilized in amber.     

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.