The structure of wing in the earliest Permopsocida.

Permopsocids are small acercarian insects with mouthparts specialized for sucking. They are closely related to Hemiptera and Thysanoptera. The earliest known representatives are from the Early Permian. Here evidence is presented that the Permopsocida occurred even earlier in Pennsylvanian (Moscovian) deposits in the Piesberg quarry near Osnabrück (Lower Saxony, Germany). This material is assigned to the Permian family Psocidiidae; Carbonopsocus mercuryi gen. et sp. nov., based on the wing venation diagnosed by the unique branching pattern of the main veins, the shape of the areola postica being longer than wide, the angular shape of the pterostigma, the ir crossvein directed proximally mid of pterostigma (apomorphy) and the vannus formed by the three veins of PCu, A1 and A2. The shape of the veins, with a Y-vein formed by the distal fusion of PCu with A1, could be a putative symplesiomorphy of the Psocodea with Permopsocida and Hemiptera. C. mercuryi gen. et sp. nov. is the first appearance date for Permopsocida and roots the Acercaria tree. In addition, another specimen of Dichentomum cf. arroyo (Psocidiidae) from Carrizo Arroyo is presented and figured, confirming the presence of the genus Dichentomum near the Carboniferous-Permian boundary and linking it to the Artinskian species from Elmo in Kansas, USA.

Thoracic and abdominal outgrowths in early pterygotes: a clue to the common ancestor of winged insects?

One of the fundamental questions in insect evolution is the origin of their wings and primary function of ancestral wing precursors. Recent phylogenomic and comparative morphological studies broadly support a terrestrial ancestor of pterygotes, but an aquatic or semiaquatic ancestor cannot be ruled out. Here new features of the branchial system of palaeodictyopteran larvae of several different instars of Katosaxoniapteron brauneri gen. et sp. nov. (Eugereonoidea) from the late Carboniferous collected at Piesberg (Germany) are described, which consist of delicate dorsolateral and lamellate caudal abdominal gills that support an aquatic or at least semiaquatic lifestyle for these insects. Moreover, the similar form and surface microstructures on the lateral abdominal outgrowths and thoracic wing pads indicate that paired serial outgrowths on segments of both tagmata presumably functioned as ancestral type of gills resembling a protopterygote model. This is consistent with the hypothesis that the wing sheaths of later stage damselfly larvae in hypoxic conditions have a respiratory role similar to abdominal tracheal gills. Hence, the primary function and driving force for the evolution of the precursors of wing pads and their abdominal homologues could be respiration.

A holomorph approach to xiphosuran evolution--a case study on the ontogeny of Euproops.

Specimens of Euproops sp. (Xiphosura, Chelicerata) from the Carboniferous Piesberg quarry near Osnabrück, Germany, represent a relatively complete growth series of 10 stages. Based on this growth sequence, morphological changes throughout the ontogeny can be identified. The major change affects the shape of the epimera of the opisthosoma. In earlier stages, they appear very spine-like, whereas in later stages the bases of these spine-like structures become broader; the broadened bases are then successively drawn out distally. In the most mature stage known, the epimera are of trapezoidal shape and approach each other closely to form a complete flange around the thoracetron (=fused tergites of the opisthosoma). These ontogenetic changes question the taxonomic status of different species of Euproops, as the latter appear to correspond to different stages of the ontogenetic series reconstructed from the Piesberg specimens. This means that supposed separate species could, in fact, represent different growth stages of a single species. It could alternatively indicate that heterochrony (=evolutionary change of developmental timing) plays an important role in the evolution of Xiphosura. We propose a holomorph approach, i.e., reconstructing ontogenetic sequences for fossil and extant species as a sound basis for a taxonomic, phylogenetic, and evolutionary discussion of Xiphosura.