New aposematic and presumably repellent bark cockroach from Lebanese amber

Balatronis cretacea Šmídová et Lei, 2017 – ancestor of the repellent harlequin cockroach (Insecta: Blattaria) is reported in this issue as the earliest representative of the type cockroach family Blattidae in the Cenomanian Myanmar amber (98 Ma). Balatronis libanensis sp.n. from the Lebanese amber (130 Ma) extends the palaeogeographic (to Gondwana) and chronostratigraphic (to Early Cretaceous) range of this advanced Mesozoic bark cockroach and shifts the expected origin of the Blattidae to the J/K boundary. In relation also to the living harlequin Neostylopyga rhombidifolia, B. libanensis is primitive in being extremely small, less coloured and with well-developed wings. It has the standard cockroach head and rudimentary, but clearly visible plesiomorphic macula-like central ocellus (as the only representative of the modern Mesoblattinidae–Blattidae–Ectobiidae–Blaberidae lineage) and represents the earliest significant record of an aposematic and thus very probably also a repellent and/or poisonous insect.     

The first fossil species of subfamily Piestinae (Coleoptera: Staphylinidae) from the Lower Cretaceous of China

Two new Early Cretaceous fossil rove beetle species, Paleosiagonium brevelytratum n. gen. n. sp. and Paleosiagonium adaequatum n. gen. n. sp. are described from the Lower Cretaceous Yixian Formation of Liaoning Province, China. They are clearly placed in the extant Piestinae according to the following characters: body elongate and flat; anterior coxae small and globose; abdomen long and parallel-sided, with six visible sterna and one pair of paratergites on segments III–VII. This is the first report on the occurrence of the coleopteran subfamily Piestinae in the fossil record. The new discoveries indicate that Piestinae have originated at least by the Early Cretaceous, at about 125–120 Ma.     

Early Cretaceous protist flagellates (Parabasalia: Hypermastigia: Oxymonada) of cockroaches (Insecta: Blattaria) in Burmese amber

Two Early Cretaceous Burmese amber cockroaches contained protists related to mutualistic flagellates occurring in extant Cryptocercus cockroaches and lower termites. The fossil protists are described as Devescovites proteus Poinar n. gen., n. sp. (Parabasalia: Trichomonadida: Devescovinidae), Paleotrichomones burmanicus Poinar n. gen., n. sp. (Parabasalia: Trichomonida), Burmanymphus cretacea Poinar n. gen., n. sp. (Hypermastigia: Trichonymphida: Burmanymphidae n. fam.) and Oxymonas gigantea Poinar, n. sp. Additional putative protists are also illustrated. Evolutionary implications of this discovery are discussed.     

Charophytes from the Upper Berriasian of the Western Interior Basin of the United States

A charophyte flora from the Upper Berriasian is described from the Lakota (Black Hills, South Dakota) and Cedar Mountain formations (San Rafael Swell, Utah) of the Western Interior Basin, United States of America. Whereas the latter is dominated by monotonous assemblages of the clavatoracean Nodosoclavator bradleyi (Harris, 1939), found in temporary lakes within palustrine facies, the flora of the Lakota Formation consists of more varied assemblages of the clavatoraceans N. bradleyi (Harris, 1939), Clavator grovesii grovesii Harris 1939, C. bilateralis Peck 1957 and early characeans (Mesochara sp. or Tolypella sp.). This flora was found in deposits related to permanent lakes in fluvial floodplains, i.e. lacustrine marls and limestones that do not show any evidence of subaerial exposure. To date, little is known about C. bilateralis, we provide a new definition on the basis of its particular structure, which shows lateral bract-cell units with a pinnate arrangement. This species appears to be endemic to North America and stratigraphically limited to the earliest Cretaceous. C. grovesii grovesii is part of a long-lasting charophyte lineage, which until now was considered to have originated in the Central Tethyan Archipelago (Europe) during the Early Berriasian, about 145 Ma, and limited to Eurasia until most of the Early Cretaceous. The new data presented here suggest that during the Berriasian this species was very broadly distributed, comprising North America, Europe and China.     

The first water measurers from the Lower Cretaceous amber of Spain (Heteroptera,Hydrometridae, Heterocleptinae)

Two fossils belonging to a new genus and species of water measurer (Gerromorpha, Hydrometridae), Alavametra popovi Sánchez-García and Nel gen. n., sp. n., are described as first definitive record of the family in Lower Cretaceous (upper Albian) amber from the Utrillas Group (Peñacerrada I site) in Spain. Although several parts of the specimens are obscured due to preservation, a sufficient number of taxonomical characters are visible to consider adequate placement within Heterocleptinae, including the very long posterior pair of cephalic trichobothria inserted on tubercles and the preapical articulation between the first and second antennal segments. The new fossil taxon is included into a cladistic analysis with extinct and extant hydrometrids, and it results putatively basal among the subfamily Heterocleptinae, suggesting that this clade was already present 105 Ma ago.     

The earliest amber-recorded type cockroach family was aposematic (Blattaria: Blattidae)

Only seven adult cockroaches have been described from Mesozoic amber. Balatronis cretacea gen. et sp. nov. from the Cenomanian burmite is the most ancient member of the type cockroach family Blattidae and the only one recorded from Cretaceous strata. Individuals represented by immature and adults (n = 14, 4) constitute 3.2 per cents of all Myanmar amber cockroaches surveyed by us – indicating habits close to source trees supported with bark body pattern (absent in sedimentary fossils). Preservation in small-sized pieces suggests longer post-depositional transportation. Its characteristic aposematic coloration suggests direct ancestry to the living repellent harlequin cockroach Neostylopyga rhombifolia, which lost its wings. Along with the bizarre Manipulatoridae, Eadiidae and Alienopteridae species, it exemplifies the high disparity of cockroaches in the ancient amber–producing tropical forest, but also documents the appearance of advanced modern taxa.     

The first fossil of dasycerine rove beetle (Coleoptera: Staphylinidae) from Upper Cretaceous Burmese amber: Phylogenetic implications for the omaliine group subfamilies

The enigmatic staphylinid subfamily Dasycerinae consists of only 17 species within a single extant genus Dasycerus, but it is easily distinguished from other rove beetles by overall, latridiid beetle-like body and extremely slender, verticillate antennae. Direct fossil evidence is lacking for this group. Here, I describe the first fossil of this peculiar subfamily, Protodasycerus aenigmaticus gen. and sp. n., from the Upper Cretaceous Burmese amber. Despite considerable external similarities to extant dasycerine species, this new genus possesses several important morphological differences: a much smaller body size (ca. 1.1 mm), narrowly elongate antennomeres 1 and 2, not strongly transverse metaventrite, elytra striate but not tricostate, and truncate posterior margin of the elytra exposing four abdominal tergites dorsally. The new finding of a Cretaceous representative of the Dasycerinae helps elucidate the origin and early evolution of the omaliine group subfamilies, implying rather close similarities between the Dasycerinae and the monobasic subfamily Neophoninae. Discovery of P. aenigmaticus suggests a rather ancient origin of the subfamily by the Upper Cretaceous.     

The oldest tachyporine rove beetle in amber (Coleoptera,Staphylinidae): A new genus and species from Upper Cretaceous Burmese amber

A new genus and species of extinct rove beetle, Procileoporus burmiticus gen. and sp. n., is described from Upper Cretaceous Burmese amber (Cenomanian; ca. 99 Ma). Due to the condition of the fossil, parts of the body could not be examined, but sufficient characteristics were visible to allow the specimen to be placed within the subfamily Tachyporinae (tribe Tachyporini). Procileoporus is most similar to the extant genus Cileoporus Campbell in having a slender abdomen, but it differs in having much shorter abdominal setae, prominent striated microreticulation on the elytra, non-lobed tergite VIII, and in other details of the male genital segments. This is the first tachyporine rove beetle from Burmese amber and the oldest record of the subfamily in amber. The evolution of sexual dimorphism in the genital segments is briefly discussed.     

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.     

Foraminifera in elevated Bermudian caves provide further evidence for +21 m eustatic sea level during Marine Isotope Stage 11

Two hypotheses have been proposed to explain the origin of marine isotope stage (MIS) 11 deposits in small Bermudian caves at +21 m above modern sea level: (1) a +21 m MIS 11 eustatic sea-level highstand, and (2) a MIS 11 mega-tsunami event. Importantly, the foraminifera reported in these caves have yet to be critically evaluated within a framework of coastal cave environments. After statistically comparing foraminifera in modern Bermudian littoral caves and the MIS 11 Calonectris Pocket A (+21 m cave) to the largest available database of Bermudian coastal foraminifera, the assemblages found in modern littoral caves – and Calonectris Pocket A – cannot be statistically differentiated from lagoons. This observation is expected considering littoral caves are simply sheltered extensions of a lagoon environment in the littoral zone, where typical coastal processes (waves, storms) homogenize and rework lagoonal, reefal, and occasional planktic taxa. Fossil protoconchs of the Bermudian cave stygobite Caecum caverna were also associated with the foraminifera. These results indicate that the MIS 11 Bermudian caves are fossil littoral caves (breached flank margin caves), where the total MIS 11 microfossil assemblage is preserving a signature of coeval sea level at +21 m. Brackish foraminifera (Polysaccammina, Pseudothurammina) and anchialine gastropods (95%, >300 individuals) indicate a brackish anchialine habitat developed in the elevated caves after the prolonged littoral environmental phase. The onset of sea-level regression following the +21 m highstand would first lower the ancient brackish Ghyben-Herzberg lens (<0.5 m) and flood the cave with brackish water, followed by drainage of the cave to create a permanent vadose environment. These interpretations of the MIS 11 microfossils (considering both taphonomy and paleoecology) are congruent with the micropaleontological, hydrogeological and physical mechanisms influencing modern Bermudian coastal cave environments. In conclusion, we reject the mega-tsunami hypothesis, concur with the +21 m MIS 11 eustatic sea-level hypothesis, and reiterate the need to resolve the disparity between global marine isotopic records and the physical geologic evidence for sea level during MIS 11.     

Late Cretaceous (Campanian/Maastrichtian) freshwater to restricted marine mollusc fauna from the Loncoche Formation,Neuquén Basin,west-central Argentina

This study is the first detailed account of freshwater to restricted marine molluscs from the Upper Cretaceous sedimentary rocks in the northern sector of the Neuquén Basin. The fossils are from the Campanian–Maastrichtian Loncoche Formation in southern Mendoza, west-central Argentina, which records the initial connection of the Neuquén Basin to the Atlantic Ocean. Six species of bivalves (Diplodon bodenbenderi, Pleiodon? sp., Isognomon? mechanquilensis, Mactridae? indet., Panopea? sp., and Laternula sp.) and three of gastropods (Paleoanculosa macrochilinoides, Paleoanculosa ameghiniana, and a possible cerithioidean) are described. Specimens were collected from fine to coarse sandstones, which may be massive or with planar stratification, planar-cross stratification or trough-cross stratification, and a few from bioclastic limestones and mudstones. Although the sections are from 50 to 300 m thick, the specimens are found only in the lower 120 m. Molluscs represent autochthonous/parautochthonous assemblages composed of mostly non-broken gastropods and articulated bivalve specimens, some of which show signs of postmortem transport; however, they were not removed far from their original habitat. Review of the habitats of living genera supports the inference of dominantly freshwater palaeoenvironments in the lower and middle part of the Loncoche Formation, with restricted marine influence in the southernmost localities studied where there are a few samples that contain specimens belonging to predominantly marine groups (e.g., Laternula, Panopea).     

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.     

Cretoleptochromus archaicus gen. et sp. nov., a new genus of ant-like stone beetles in Upper Cretaceous Burmese amber (Coleoptera,Staphylinidae, Scydmaeninae)

A new genus and species of the ant-like stone beetle supertribe Mastigitae, Cretoleptochromus archaicus gen. et sp. nov., is described and figured based on an exceptionally well-preserved individual in the Upper Cretaceous (lowermost Cenomanian, ca. 99 Ma) amber from Myanmar. Cretoleptochromus is undoubtedly placed in Mastigitae based on its distinctly geniculate antennae with strongly elongate antennal scape, and pointed labial palpomere III. Cretoleptochromus is most likely the sister taxon to the extinct Palaeoleptochromus O'Keefe from the Late Cretaceous Canadian amber. Cretoleptochromus shares with Clidicini the distinct posterior collar of pronotum; with Leptomastacini the presence of mesocoxal lateroventral bristles. The new discovery may help to elucidate the relationships among the Recent three tribes, particularly for Leptomastacini and Clidicini.     

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.     

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.     

New Early Cretaceous sharks (Chondrichthyes,Elasmobranchii) from deep-water deposits of Austria

Early Cretaceous elasmobranchs still are very insufficiently known despite all progress that has been accomplished in recent years. Here, a small elasmobranch assemblage is presented from the Valanginian of Austria that contributes significantly to a better understanding of early Cretaceous elasmobranch diversity. The new assemblage comprises two new carcharhiniform sharks, Altusmirus triquetrus gen. et sp. nov. and Fornicatus austriacus gen. et sp. nov., a new orectolobiform shark, Similiteroscyllium iniquus gen. et sp. nov., and a galeomorph shark tooth of uncertain affinities. The recent identification of Similiteroscyllium gen. nov. has shown that it has strong similarities with Ornatoscyllium rugasimulatum from the Lower Cretaceous of France. Significant differences of the tooth morphology of O. rugasimulatum and the type specimen O. freemani justify full reconsideration of the systematic position of O. rugasimulatum and require it to be reassigned to Similiteroscyllium gen. nov. described in this paper. The new assemblage described here, and those from the Valanginian of France and Poland comprising 30 additional taxa, indicates that Early Cretaceous elasmobranch diversity was significantly higher than previously assumed. Consequently, the supposed diversity decline of elasmobranchs across the Jurassic-Cretaceous boundary represents a collecting bias rather than a real pattern.     

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.     

First U–Pb SHRIMP age for the Pilmatué Member (Agrio Formation) of the Neuquén Basin,Argentina: Implications for the Hauterivian lower boundary

Ammonite-based biostratigraphic schemes for the Lower Cretaceous are fairly well refined across the world, from the standard zonation in the West Mediterranean province to the Boreal and Austral provinces in the northern and southern hemispheres, respectively. However, the lack of radioisotopic ages associated to the fossil-rich, Lower Cretaceous marine successions has hindered the accurate establishment of the numerical ages for the lower boundaries of its several stages (from Berriasian to Albian). Geochronological dating by U–Pb SHRIMP of a tuff layer that occurs within beds belonging to the Holcoptychites neuquensis Zone in the Pilmatué Member of the Agrio Formation in the Austral province (Neuquén Basin, Argentina) has resulted in an absolute age of 130.0 ± 0.6 Ma (2 sigma internal errors only) or 130.0 ± 0.8 Ma (including calibration and decay constant uncertainties). This age is interpreted to represent the time of eruption and thus the timing of the pyroclastic deposit. The H. neuquensis Zone is the equivalent of the A. radiatus Zone in the West Mediterranean province. Therefore, the obtained age is the first numerical data that could help constrain the Hauterivian lower boundary. Indeed, there is reasonable agreement with the latest proposed lower boundary of the Hauterivian at ∼132.9 Ma. On the other hand, the duration recently established for this stage would be hard to reconcile with the stratigraphic record of the entire Hauterivian in the study region (northeastern Neuquén Basin). Therefore, the results of this contribution could also help to assess the extent of the Hauterivian and associated stages.     

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.