Alexei S. Bashkuev

A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies

Long-Lost Pollinators The rise of angiosperms in the Early Cretaceous (∼140 million years ago) was accompanied by coevolution of a variety of insects, including flies, bees, and wasps required for pollination. Ren et al. (p. 840; see the Perspective by Ollerton and Coulthard) show that three families of scorpionflies had already evolved specialized mouth parts for feeding on the nectar of gymnosperms, as early as the Middle Jurassic (∼170 million years ago). The diversity and specialization of these insects and related plant structures suggests that they were also involved in pollination. These families died out later in the Cretaceous as angiosperms began to dominate. Prior to the coevolution of angiosperms and pollinating insects, scorpionflies may have been pollinating gymnosperms. The head and mouthpart structures of 11 species of Eurasian scorpionflies represent three extinct and closely related families during a 62-million-year interval from the late Middle Jurassic to the late Early Cretaceous. These taxa had elongate, siphonate (tubular) proboscides and fed on ovular secretions of extinct gymnosperms. Five potential ovulate host-plant taxa co-occur with these insects: a seed fern, conifer, ginkgoopsid, pentoxylalean, and gnetalean. The presence of scorpionfly taxa suggests that siphonate proboscides fed on gymnosperm pollination drops and likely engaged in pollination mutualisms with gymnosperms during the mid-Mesozoic, long before the similar and independent coevolution of nectar-feeding flies, moths, and beetles on angiosperms. All three scorpionfly families became extinct during the later Early Cretaceous, coincident with global gymnosperm-to-angiosperm turnover.

Fossil Insects of the Middle and Upper Permian of European Russia

Fossil insects of European Russia from the Urzhumian to Vyatkian stages are reviewed, new taxa are described, and dynamics of insect taxonomic diversity around the Permian-Triassic boundary in light of the Paleozoic-Mesozoic boundary global extinction problem is analyzed. Traces of interactions between arthropods and plants are analyzed. Insect-bearing deposits of the Late Paleozoic found in the northern and eastern areas of the East European Platform are unique on the global scale in their completeness and continuity, allowing us to trace especially comprehensively the biotic processes that occurred around the boundary described as the time of the greatest biotic catastrophe of the Phanerozoic. A total of 28 genera and 111 species are newly described. Within the range from the Urzhumian to the Permo-Triassic boundary, 15 representative successive assemblages, including 112 families, are recognized (seven in the area in question and eight in other regions of Asia, Australia, and Africa). New tools are developed for the analysis of the dynamics of diversity. These tools show an approximately equilibrium (slightly positive) dynamics in the Urzhumian and Severodvinian and a drop in diversity during the Vyatkian Age. It is shown that Permian insect assemblages acquired a substantially post-Paleozoic pattern much earlier than the end of the Paleozoic. The character of changes that took place in the Induan and Olenekian remains uncertain, but a large-scale extinction event did not occur here: most families that have not been recorded at the beginning of the Triassic are recorded again in the Middle and Upper Triassic. Nevertheless, a biotic crisis probably actually took place, but was reduced to reorganization of the biota’s structure, which provided enormous growth of biodiversity over subsequent hundreds of millions of years, rather than resulted in catastrophic extinction. This study is intended for entomologists, stratigraphers, and all readers interested in the biotic events that took place around the Permian-Triassic boundary.

Insects from the Buntsandstein of Lower Franconia and Thuringia

Upper Buntsandstein deposits (mainly Myophoria beds, Röt Formation, Early Anisian) in Lower Franconia and Thuringia have yielded a rather rich insect fauna comprising ca. 300 insect specimens assigned to ten orders: Archaeognatha (Dasyleptidae), Ephemeroptera, Blattodea, Grylloblattida, Orthoptera, Hemiptera, Glosselytrodea, Coleoptera, Mecoptera and Diptera. The systematic list of recorded insects is provided. Two species are identified as Triassodotes vogesiacus Sinitshenkova, Marchal-Papier, Grauvogel-Stamm et Gall, 2005 (Ephemeroptera: Misthodotidae) and Pseudopolycentropus triasicus Papier, Nel et Grauvogel-Stamm, 1996 (Mecoptera), which were previously described from the “Grès à Voltzia” Formation of the Vosges, the stratigraphically closest insect fauna. All grylloblattid specimens are identified as Chauliodites picteti Heer, 1864 (Chaulioditidae), known previously from the Middle Buntsandstein of Gödewitz, Saxony-Anhalt. The new genus and species Hammephemera pulchra Sinitshenkova, gen. et sp. n. (Ephemeroptera: Sharephemeridae) is described.KurzfassungDer Oberen Buntsandstein (hauptsächlich aus Myophorien-Schichten, Röt Formation, Unter-Anisium) in Unterfranken und Thüringen bringt ziemlich reiche Insektenfauna, in der ca. 300 Exemplare festgestellt wurden, die zu zehn Ordnungen gehören: Archaeognatha (Dasyleptidae), Ephemeroptera, Blattodea, Grylloblattida, Orthoptera, Hemiptera, Glosselytrodea, Coleoptera, Mecoptera und Diptera. Zwei Arten, Triassodotes vogesiacus Sinitshenkova, Marchal-Papier, Grauvogel-Stamm et Gall, 2005 (Ephemeroptera: Misthodotidae) und Pseudopolycentropus triasicus Papier, Nel et Grauvogel-Stamm, 1996 (Mecoptera) sind auch in der stratigraphisch nächsten Insektenfauna der “Grès à Voltzia” Formation der Vogesen verbreitet. Alle Grylloblattiden-Reste werden zu Chauliodites picteti Heer, 1864 (Chaulioditidae) gestellt. Neu aufgestellt werden eine neue Gattung und Art der Sharephemeridae (Ephemeroptera): Hammephemera pulchra Sinitshenkova, gen. et sp. n.

Upper Jurassic Lagerstätte Shar Teg, southwestern Mongolia

One of the most interesting Mesozoic Lagerstätten, Shar Teg in southwestern Mongolia, is reviewed. The geological structure and oryctocoenoses of Shar Teg are described. Shar Teg is one of the most diverse Jurassic Lagerstätten in terms of fossils represented. Fossils from Shar Teg include aquatic and terrestrial plants, mollusks, crustaceans, insects, fishes, amphibians, reptiles, and mammals. Insects are the most diverse group. To date, a total of 297 species of 161 families and 22 orders have been described or recorded in Shar Teg, making it possible to reconstruct comprehensively the Jurassic biota of the locality. The oryctocoenosis composition is peculiar; it includes only six species described from other localities. The oryctocoenosis appears to link the faunas of eastern Asia, Central Asia, and Europe. A total of 31 new species are described.

New Jurassic Hangingflies (Insecta: Mecoptera: Bittacidae) from Inner Mongolia, China

A new bittacid genus, Composibittacus gen. nov., with two new species, C. bipunctatus gen. et sp. nov. and C. reticulatus sp. nov., and a new species of Orthobittacus Willmann 1989, O. maculosus sp. nov., are described from the latest Middle Jurassic Jiulongshan Formation of Daohugou, Inner Mongolia, China. Composibittacus gen. nov. has unique wing characters, such as five pterostigmal crossveins between R1 and R2 and R1 and R2+3 and an elongated pterostigma area, which distinguishes it from all other known genera in Bittacidae. Orthobittacus maculosus sp. nov. differs from other species of Orthobittacus by a combination of the following wing characters: M with six branches in forewings and hind wings, two crossveins between C and Sc, and two pterostigmal crossveins in the forewing. In addition, O. maculosus sp. nov. has light-colored or white spots on the fore- and hind wings. These new venational characters of Composibittacus gen. nov. and O. maculosus sp. nov. enhance our understanding of the diverse morphological characters of early hangingflies. Furthermore, based on the striking similarity of the wings of O. maculosus sp. nov. and Juracimbrophlebia ginkgofolia Wang, Labandeira, Shih & Ren 2012 (Cimbrophlebiidae), we propose that leaf mimesis and mutualism with ginkgo plants might have been present in the Bittacidae, as has been proposed in the Cimbrophlebiidae.

Coptoclavid beetles (Insecta: Coleoptera: Adephaga) from the Triassic of Lower Franconia, Germany

Many fossils of water beetles of the family Coptoclavidae have been found unexpectedly among insect fossils from the Middle Triassic (Anisian) and Upper Triassic of Franconia, Germany. Very small (about 3 mm long) beetles with longitudinal dark stripes on the elytra, similar in coloration pattern to beetles of the genus Holcoptera Handlirsch, 1906 from the terminal Triassic and Lower Jurassic of England and Lower Jurassic of the United States, have been found in Anisian deposits. Similar but larger beetles (about 5 mm long) have been found in Keuper (Carnian) deposits, in which they occur much more frequently than in Anisian localities, even dominating some of these localities. Unfolded or fragmentary hindwings that also belong to beetles of the same species and match the elytra in size have also been found. There are no known localities in which so many beetle hindwings occur, especially hindwings of the same species. They are described here as wings of Coptoclavidae. A small incomplete larva matching these beetles in size has also been found. The beetles are described as representatives of a new genus of the subfamily Coptoclaviscinae. In addition to them, the larvae of a rather large coptoclavid Protonectes Prokin et Ponomarenko, 2013 was described earlier from the same deposits; it is assigned here to the subfamily Timarchopsinae. Only two elytra with numerous fine grooves not bearing any punctures match this larva in the available collection. Similar elytra were described earlier from the Jurassic of Europe and Siberia as Dinoharpalus Handlirsch, 1906. A small elytron with many grooves was described in the same genus from the terminal Permian of Eastern Europe. Similar elytra are known in coptoclavids described from the Lower Cretaceous of Spain as Hispaniclavina. The presence of such advanced beetles as coptoclavids as early as the Lower Anisian shown that the famous Permian–Triassic crisis was not so deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic. Keywords: insects, beetles, Triassic

First Triassic record of the beetle family Permocupedidae (Insecta: Coleoptera): a peculiar example of a Lazarus taxon

The first Mesozoic representative of the extinct archostematan beetle family Permocupedidae, Frankencupes ultimus, gen. et sp. nov., is described based on two isolated elytra from the Lower Anisian (Middle Triassic) Röt Formation of Lower Franconia, Germany. The new fossil occurrence extends the range of the family from the Lower Wuchiapingian (Upper Permian) up to the Anisian, and represents a fine example of a Lazarus taxon in the fossil record of beetles.