Laurasian ancestors and “Gondwanan” descendants of Rotoitidae (Hymenoptera: Chalcidoidea): What a review of Late Cretaceous Baeomorpha revealed

Baeomorphinae Yoshimoto, 1975, based on Baeomorpha Brues, 1937, is transferred from Tetracampidae Förster, 1856 and recognized as a junior synonym of Rotoitidae Bouček and Noyes, 1987 (Hymenoptera: Chalcidoidea) syn. nov. following ICZN (1999) article 35.5. Inclusions in Taimyr amber (84–100 Ma) assigned to Baeomorpha are reviewed and the following eleven new species, all of Gumovsky, are described: B. avamica sp. nov., B. baikurenis sp. nov., B. bianellus sp. nov., B. caeleps sp. nov., B. gracilis sp. nov., B. ingens sp. nov., B. quattorduo sp. nov., B. quattoruno sp. nov., B. popovi sp. nov., B. yantardakh sp. nov., and B. zherikhini sp. nov. The recognized species are differentiated in separate keys to females and males and illustrated through microphotography. Two of four previously described Baeomorpha species from Campanian Canadian amber are synonymized: B. distincta Yoshimoto and B. elongata Yoshimoto under B. ovatata Yoshimoto (syn. nov.). One enigmatic rotoitid inclusion, which differs from Baeomorpha species in the possession of very short stigmal vein, is described as Taimyromorpha pusilla Gumovsky gen. et sp. nov. Inclusions containing specimens identified as Baeomorpha and Taimyromorpha are found in amber from Taimyr and Canada that originated from Laurasia, not Gondwana. Two Realms are newly proposed to recognize different Cretaceous faunal elements, a more northern Baeomorpha Realm that is characterized by a temperate or warm temperate climate and very abundant aphid fossils, and the Isoptera Realm, an opposing southward territory with a warmer climate and common termite but rare aphid fossils. The newly described fossils indicate the southern hemisphere distribution of extant Rotoitidae is relictual with the pattern observed being formed at least in part by extinction events, though distributions of the only two extant rotoitid genera, Rotoita Bouček and Noyes, 1987 (New Zealand) and Chiloe Gibson and Huber, 2000 (small area in the southern Chile) may have been more extensive in the past. Both of known regions of extant Rotoitidae have highly suppressed ant faunas, which may suggest that their survival there depended on low biocenotic pressure by ants, perhaps as low as is hypothesized for the Late Cretaceous. The Canadian amber genera Distylopus Yoshimoto, 1975 (Distylopinae) and Bouceklytus Yoshimoto, 1975 (Bouceklytinae) are excluded from Tetracampidae and regarded as Chalcidoidea incertae sedis.     

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.     

First record of the beetle family Lasiosynidae (Insecta: Coleoptera) from the Lower Cretaceous of China

A new genus, Cretasyne gen. nov., in the family Lasiosynidae, with two species C. lata sp. nov. and C. longa sp. nov., is described from the Lower Cretaceous Yixian Formation of Inner Mongolia, China. These fossils are the first record of the family from the Yixian Formation and provide significant information on lasiosynid morphology and diversity. The fossil record of Lasiosynidae is also reviewed.     

A new Late Cretaceous genus and species of polypore fungus beetles (Coleoptera,Tetratomidae) from northern Myanmar

A new polypore fungus beetle is described and illustrated, under the name of Thescelostrophus cretaceus gen. et sp. nov., representing the first documented occurrence of the tribe Eustrophini. The well-preserved specimen was collected from the Upper Cretaceous (Cenomanian, ca. 99 Ma) amber near the Hukawng Valley of northern Myanmar. This fossil species can be assigned to the extant subfamily Eustrophinae based on its elongate oval and strongly narrowed posteriorly body, simple and narrow tarsi, and somewhat clubbed antennomeres. The comparison among the extinct and extant eustrophines supports the previous hypothesis that antennal morphology of early eustrophines was very diverse. Additionally, an overall similarity between Thescelostrophus and extant eustrophines suggests a potential fungivory of this fossil species. Morphological characters preserved in the Burmese amber highlight the diversity of tetratomids during the Late Mesozoic and provide data for future phylogenetic studies of Tetratomidae.     

The most mysterious beetles: Jurassic Jurodidae (Insecta: Coleoptera) from China

Jurodidae are thought to be some of the world's most mysterious beetles and display an intriguing mixture of characters occurring in three different suborders. Hitherto, all known fossil and extant Jurodidae were extremely rare and restricted to Russian Siberia and the Far East. Here we describe two new species, Jurodes daohugouensis sp. nov. and Jurodes pygmaeus sp. nov. from the Middle Jurassic of Daohugou, China. A key to all species of Jurodidae and their research history are given, and the detailed morphology of fossil Jurodidae is also presented. Our results confirm the presence of three ocelli in fossil Jurodidae, and reveal that hind wings of J. pygmaeus sp. nov. possess well developed radial and oblong cells, and are very similar to those of extant Jurodidae (Sikhotealinia zhiltzovae). Male genitalia of fossil Jurodidae were described herein for the first time, showing a basal trilobate structure. The presence of exposed propleuron, three pairs of external trochantins, metanepisterna with ridges, primitive characters of wing venation further support the probable placement of Jurodidae in the suborder Archostemata. J. daohugouensis and J. pygmaeus are very similar to S. zhiltzovae in having the big protruding eyes, elevated median portion of frons bearing three ocelli, pubescent body, contiguous procoxae, exposed trachantion of all coxae, elytra with rows of punctures, and wing venation. Our results show that these key characters of extant Jurodidae can be traced back to the Middle Jurassic, revealing that the family has been in a period of evolutionary stasis for at least 160 million years. Furthermore, our discoveries widen the paleogeographic distribution of fossil Jurodidae from Russian Siberia to northern China.     

A new family of scorpionflies (Insecta; Mecoptera) from the Lower Cretaceous of England

Seven specimens of fossil scorpionflies (Mecoptera) not assignable to any known family were discovered in the Wealden Supergroup (Lower Cretaceous) of southern England. They were found at Rudgwick Brickworks, West Sussex and Smokejacks Brickworks, Surrey and came from the Upper Weald Clay Formation, dated as Barremian (∼129.4–125 Ma). A new family – Englathaumatidae fam. nov., new genus – Englathauma gen. nov. and two new species E. crabbi sp. nov. and E. mellishae sp. nov. are described. A discussion of systematic position of these new taxa within the order Mecoptera is given. Englathaumatidae fam. nov. has been a nomen nudum since the year 2002, due to the first author's untimely death.     

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.     

The oldest representatives of the family Throscidae (Coleoptera: Elateriformia) from the Lower Cretaceous Lebanese amber

Two new genera and two new species of fossil Throscidae: Potergosoma gratiosa gen. et sp. nov. and Rhomboaspis laticollis gen. et sp. nov. are described from the Lower Cretaceous Lebanese amber and are compared with extant and extinct genera. The described amber inclusions are the oldest known representatives of the family Throscidae. Some hypotheses on the phylogeny of the family Throscidae and the position of it in the superfamily Elateroidea are discussed.     

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 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 earliest pygidicranid (Insecta: Dermaptera) from the Lower Cretaceous of China

A new earwig genus with a new species, Cylindopygia falcata gen. et sp. nov., is described and illustrated based on two well-preserved, nearly complete female specimens from the Lower Cretaceous, Yixian Formation in Huangbanjigou, Liaoning Province, China. C. falcata is assigned to Pygidicranidae mainly due to the following characters: head obtuse-triangular with posterior margin straight, abdomen robust, subcylindrical and densely setose, and femora compressed and carinate. The new finding represents the earliest fossil record of Pygidicranidae and the first record of Pygidicranidae in China.     

New genera and species of bark-gnawing beetles (Coleoptera: Trogossitidae) from the Yixian Formation (Lower Cretaceous) of Western Liaoning,China

Two new genera from the Lower Cretaceous Yixian Formation of Beipiao City, Liaoning Province, northeastern China are described and illustrated. Paracretocateres gen. nov. (type species P. bellus, sp. nov.) and Yixianteres gen. nov. (type species Y. beipiaoensis sp. nov.) are assigned to the subfamily Lophocaterinae based on the state of the procoxae which are not projecting, the transverse procoxal cavities, and the exposed protochantins. The new taxa broaden the diversity of this family and provide further morphological characters for phylogenetic studies of Trogossitidae. They also provide evidence that lophocaterines well-diversified prior to the trogossitines and their biologies may have been similar to extant forms.     

Myamalycocerus vitalii: A new genus and species of soldier beetle in Burmese amber (Coleoptera Cantharidae)

A new fossil soldier beetle Myamalycocerus vitalii gen. et sp. nov., is described and illustrated from an inclusion in Upper Cretaceous Burmese amber (Myanmar). It differs from all extant genera that have elytra adorned with small striae by possessing raised punctation, very probably an ancient character as it is absent among current species. It differs from the fossil Ornatomalthinus Poinar et Fanti, by the long elytra and relief points which are less raised and more numerous. This new genus appears vaguely related (not necessarily phylogenetically) to the current genus Lycocerus Gorham.     

New fossil flower bugs (Heteroptera: Cimicomorpha: Cimicoidea: Vetanthocoridae) with uniquely long ovipositor from the Yixian Formation (Lower Cretaceous), China

A new genus and new species, Longilanceolatus tenellus gen. et sp. nov., is described and assigned to the tribe Vetanthocorini of the fossil family Vetanthocoridae. In addition, a new species, Curvicaudus spinosus sp. nov., is also described. All specimens were collected from the Lower Cretaceous of the Yixian Formation at Huangbanjigou, Chaomidian Village, Beipiao City, Liaoning Province, northeastern China. Body size, ovipositor length, abdomen length, and other characters of eleven genera from the Middle Jurassic to Eocene are summarized. Comparison of body size of all specimens in the Vetanthocoridae and Anthocoridae (sensu lato) indicate a general trend toward smaller body size through their evolution. Moreover, Longilanceolatus, with uniquely long ovipositor, is related to a different type of oviposition compared with extant flower bugs.     

A new parasitoid wasp (Hymenoptera: Chalcidoidea) from the Lower Cretaceous Crato Formation of Brazil: The first Mesozoic Pteromalidae

A new genus and species of small (3.5 mm excluding ovipositor) parisitoid wasp is described from the Lower Cretaceous (Aptian) Crato Formation Lagerstätte of Brazil. Parviformosus wohlrabeae gen. et sp. nov. is known from a single female imago and is assigned to Pteromalidae. It is diagnosed by the robustness of the scutellum, the structure, size and positioning of the mesopleuron, the complexity of the propodeum–petiole junction and a posteriorly curved dorsal ‘lip’ on metasomal segment 4. At only 3.5 mm in length, P. wohlrabeae is the smallest fossil wasp from the Cretaceous of South America and the first Mesozoic representative of Pteromalidae.     

A new genus and species of soldier beetle from Upper Cretaceous Burmese amber (Coleoptera,Cantharidae, Malthininae)

A new Upper Cretaceous genus and species of soldier beetles, Archaeomalthodes rosetta gen. et sp. nov., is described and illustrated from an individual preserved in Upper Cretaceous (Cenomanian, ca. 99 Ma) amber from northern Myanmar. It is undoubtedly placed in extant subfamily Malthininae based on its small-sized body, somewhat abbreviated elytra and fusiform terminal maxillary palpomere, representing the oldest documented occurrence of Malthininae. It suggests that this subfamily is an ancient group, which originated at least in the earliest Late Cretaceous. Our discovery sheds light on the palaeodiversity of Cantharidae in the Late Mesozoic. Together with other previously reported fossil cantharids, it is likely that Malthininae has been fairly diverse during the early evolution of Cantharidae. On the other hand, a morphological similarity between Archaeomalthodes and Recent malthinines and the occurrence of flowering plants in the Burmese amber implies a potential flower-visiting behaviour of this fossil species.