Abstract The cliffs and foreshore at Lyme Regis on the Dorset coast expose a sequence of Early Jurassic marine limestones and mudstones of the Blue Lias Formation and the overlying Charmouth Mudstone Formation, the lowest units of the Hettangian–Aalenian Lias Group. Known for its fossils since at least the mid-seventeenth century, this coastal section became famous in the early nineteenth century for the bones of ‘fossil crocodiles’. Many of the most significant discoveries were made by a family of fossil dealers whose best-known member was Mary Anning (1799–1847). Anning and her family recovered the first scientifically described ichthyosaurs, the first complete plesiosaur and the first British pterosaur to be identified. Anning's finds from Lyme Regis formed the basis of the pioneering palaeoenvironmental reconstruction Duria antiquior . Her geologist friends, some with close personal associations to the town, did much to publicize her discoveries and contribute to both her fame and that of the locality. This famous coastal section, with its important historical associations with a key period in the development of geology and the source of so many significant discoveries in the early nineteenth century, now lies within the Jurassic Coast UNESCO World Heritage Site.

Abstract Ctenochasmatid pterosaurs flourished and diversified in the Early Cretaceous Jehol Biota. Here, a partial mandible of Forfexopterus is described based on a three-dimensional reconstruction using high-resolution X-ray computed tomography (CT) data. The first nine pairs of functional teeth of the rostral dentition revealed along with their replacements. The functional teeth are evenly arranged with a tooth density of 2.2 teeth/cm. The tooth crown is distinctly reduced from its base to the tip, and framed by two weak ridges, possibly a pair of vestigial carinae. The replacement teeth are sharp and pointed, and have erupted slightly against the medial surface of the functional teeth. Surprisingly, tooth wear is observed in this specimen, the first record of tooth–tooth occlusion in ctenochasmatids. The wear facets exhibit high-angled lingual and lower-angled labial facets, implying a unique tooth–tooth occlusion in pterosaur clade. This discovery indicates that the Jehol ctenochasmatids possibly employed a more active feeding strategy than other filter-feeding pterosaurs (e.g. Ctenochasma , Pterodaustro , Gnathosaurus ).

Abstract Starting in the early 1950s, palaeontologists began to discover a wide range of Cretaceous terrestrial fossils in the Laiyang, Qingshan and Wangshi groups of the Jiaolai Basin, which resides in the eastern part of the Shandong Peninsula of northern China. Significant specimens from these deposits include various dinosaur eggs, footprints and fossils, including hadrosauroids, tyrannosaurids and ankylosaurids. These expanded the understanding of evolution, biodiversity and palaeoecology in East Asia. While many examples of the Jehol Biota from this area are not well constrained in terms of their stratigraphy and geochronology, previous studies have generally suggested that fossils from this region represent the second or third phase of Jehol Biota development. This paper reviews fossils, stratigraphic correlations, tectonic history and age estimates for the less well-studied outcrops of the Shandong Peninsula that host the Jehol Biota. We report three new 40 Ar/ 39 Ar ages for the Qingshang Group and discuss how these somewhat imprecise ages still constrain chronostratigraphic interpretation for the fossil-rich units.

ABSTRACT Professor of paleontology at the University of Basel, Switzerland, Manfred Reichel was as much an accomplished scientist as a talented artist. Skilled in mental 3-D-visualization, aided by a sharp memory, and with a fine hand for illustration, he introduced comparative anatomy to the study of foraminifera, which is masterly illustrated in his analysis of Alveolinids, and used his knowledge of locomotion in modern birds as an actualistic method for portraying flying reptiles. Teaching was his motivation, and to aid his classes, beyond multiple drawings, he created a large number of scaled structural models of foraminifera and a life-size wooden replica of the Pteranodon with mobile articulations. Manfred Reichel was a perfectionist who left a large part of his studies and drawings unpublished.

Abstract: Pterosaurs were the first vertebrates to evolve powered flight and occupied the skies of the Mesozoic for 160 million years. They occurred on every continent, evolved their incredible proportions and anatomy into well over 100 species, and included the largest flying animals of all time among their ranks. Pterosaurs are undergoing a long-running scientific renaissance that has seen elevated interest from a new generation of palaeontologists, contributions from scientists working all over the world and major advances in our understanding of their palaeobiology. They have especially benefited from the application of new investigative techniques applied to historical specimens and the discovery of new material, including detailed insights into their fragile skeletons and their soft tissue anatomy. Many aspects of pterosaur science remain controversial, mainly due to the investigative challenges presented by their fragmentary, fragile fossils and notoriously patchy fossil record. With perseverance, these controversies are being resolved and our understanding of flying reptiles is increasing. This volume brings together a diverse set of papers on numerous aspects of the biology of these fascinating reptiles, including discussions of pterosaur ecology, flight, ontogeny, bony and soft tissue anatomy, distribution and evolution, as well as revisions of their taxonomy and relationships.

Abstract: Understanding the ecological roles of pterosaurs is a challenging pursuit, but one aided by a growing body of fossil evidence for their dietary preferences and roles as food sources for other species. Pterosaur foraging behaviour is represented by preserved gut content, stomach regurgitates, coprolites and feeding traces. Pterosaurs being eaten by other species are recorded by tooth marks and teeth embedded in their fossil bones, consumer gut content and regurgitate, and their preservation entangled with predatory animals. This palaeoecological record has improved in recent years, but remains highly selective. The Jurassic rhamphorhynchid Rhamphorhynchus , Cretaceous ornithocheiroid Pteranodon and azhdarchid pterosaurs currently have the most substantial palaeoecological records. The food species and consumers of these taxa conform to lifestyle predictions for these groups. Rhamphorhynchus and Pteranodon ate and were eaten by aquatic species, matching expectations of these animals as sea-going, perhaps partly aquatic species. Possible azhdarchid pterosaur foraging traces alongside pterosaur tracks, and evidence that these animals were eaten by dinosaurs and Crocodyliformes, are consistent with hypotheses that azhdarchids foraged and lived in terrestrial settings. Fossil evidence of pterosaur palaeoecology remains rare: researchers are strongly encouraged to put specimens showing details of dietary preferences, foraging strategies or interactions with other animals on record.

Abstract: Simple, three-dimensional, digital models of the crania and mandibles of 22 pterosaurs – 13 pterodactyloids and nine non-pterodactyloids (‘rhamphorhynchoids’) – were generated to investigate gross-level mechanical aspects of the skulls as they would related to feeding behaviour such as bite force and speed of jaw motions. The key parameter was the determination of second moments of area of the mid-muzzle region and the computation of the bending moment relative to the occiput. The shorter, stockier skulls of basal ‘rhamphorhynchoids’ were the strongest for their size in terms of potential resistance to dorso-ventral bending, and this finding correlates with their robust dentitions. More derived ‘rhamphorhynchoids’ showed the start of a trend towards weaker skulls, but faster jaw adduction was interpreted to be an adaptation for the snatching of small prey. Pterodactyloids continued the trend to lengthen the skull and to reduce its cross-sectional area, resulting in less stiff skulls, but more rapid opening and closing of the jaws. Changes in the rear of the skulls and the development of coronoid eminences on the mandibles of all the pterodactyloids are correlated with the reduction in bite force and a concomitant increase in jaw closing speed.

Abstract: The unique morphology of pterosaurs makes them a compelling group to study, but the lack of ready analogues or descendant clades presents problems when inferring their biology and ecology. In this paper, the extant phylogenetic bracket is used to reconstruct the musculature of the pterosaur Vectidraco daisymorrisae and a detailed comparison is made between the musculature of this taxon and other reconstructions in the literature. M. iliofibularis is reconstructed as originating upon the angular process of the posterior iliac process, putting it into a mechanically advantageous position as an abductor. M. flexor tibialis internus is reconstructed as greatly enlarged over the ancestral state, probably in response to the reduction of M. caudofemoralis brevis and the tail. This enlargement is considered to correspond to the increasing role of M. flexor tibialis internus as a hip retractor.

Abstract: The preservation of the wing membrane of pterosaurs is very poor and the available fossil evidence does not allow its properties to be reconstructed. In contrast, the fossil record for the wing bones is relatively good and the advent of CT scanning has made it possible to build high-fidelity structural models of the wing spar. The bending strength of the wing spar of a 6 m wingspan ornithocheirid pterosaur is used to infer the likely membrane tension. The tensions required to suppress aeroelastic flutter and to minimize ballooning of the membrane under flight loads are also estimated. All three estimates are of similar magnitude and imply that the membrane must have contained high-modulus material, supporting the view that the reinforcing aktinofibrils were keratinous.

Abstract: Four specimens of the pterosaur Pteranodon exhibit patterns of irregular alternating light and dark bands on the lateral surfaces of the upper jaw anterior to the nasoantorbital fenestra. Examinations reveal that the maxilla and premaxilla of Pteranodon consisted of two thin sheets of bone interconnected by regularly spaced septa with the spaces contained within presumably pneumatized, resulting in a structure analogous to modern honeycomb sandwich panels. The alternating light and dark bands resulted from waves of bone deposition moving anteriorly along the external surface of the lateral sheet of bone and laying down thin laminae of new bone while bone was simultaneously resorbed from the internal surface of the lateral sheet to maintain its thickness. The specimens that exhibit the bands were immature males and no banding was found in mature specimens or immature females. Therefore, the presence of the bands in immature males is interpreted as correlated with the enlargement and reshaping of the rostrum as males approached and attained sexual maturity.

Abstract: We report on a diminutive pterosaur specimen (MIC-V246), from the Lower Cretaceous Lagarcito Formation, which has anatomical features and general proportions that agree with those of other juvenile specimens of the filter-feeding pterosaur Pterodaustro guinazui . MIC-V246 is nearly complete, with the majority of its bones in natural articulation. The specimen is preserved within a small oval surface inferred to demarcate the outline of an egg. It includes remains of the skull and mandible, some cervical, dorsal, sacral and caudal vertebrae, ribs, gastralia, both shoulder girdles, part of the ilium, and the forelimbs and hindlimbs. The skeleton is arranged in a position similar to that of avian embryos, with the wings folded sideways, the hindlimbs flexed, and the skull tucked beneath a wing. The bones are partially covered by an irregularly distributed substance differing in texture and colour from the bony elements, and identified as remnants of the eggshell. The diminutive size of the skeleton, along with its arrangement and morphology, support the identification of this specimen as one of the very few pterosaur embryos known worldwide. Its similarity in size to other Pterodaustro specimens interpreted as hatchlings suggests that MIC-V246 was near hatching when it died.

Abstract: A new anurognathid pterosaur, Vesperopterylus lamadongensis gen. et sp. nov., is erected based on a complete skeleton with a skull preserved. It is characterized by two short distinct ridges present on the ventral surface of the cervical vertebrae; coracoids slightly longer than scapula; humerus, wing phalanx 3 and tibia nearly the same in length; grooves clearly present on the posterior surface of the wing phalanges 1–3; and the first toe reversed. It is the first anurognathid pterosaur from China with a definitively short tail, and the first pterosaur with a reversed first toe. The reversed first toe of Vesperopterylus indicates that it had arboreal habitats. The discovery of Vesperopterylus lamadongensis from the Jiufotang Formation strongly expands the geological age range for anurognathid pterosaurs. Supplementary material: The character list and data matrix for phylogenetic analysis are available at https://doi.org/10.6084/m9.figshare.c.3873394

Abstract: Pterosaur fossils are rare in the Morrison Formation, and most are poorly preserved. The Breakfast Bench Facies (BBF) at Como Bluff produces incomplete but uncrushed limbs. One proximal and two distal femora match a complete femur (BYU 17214) referred to Mesadactylus . Unexpectedly, both of the BBF distal femora possess a large intercondylar pneumatopore. BYU 17214 also possesses an intercondylar pneumatopore, but it is smaller than in the BBF femora. Distal femoral pnuematicity is previously recognized only in Cretaceous azhdarchoids and pteranodontids. A peculiar BBF jaw fragment shows strongly labiolingually compressed, incurved crowns with their upper half bent backwards; associated are anterior fangs. We suspect this specimen is a previously undiagnosed pterosaur. Additional BBF material documents a diverse pterosaur fauna including a humerus with a greatly expanded ectepicondyle possibly from a non-pterodactyloid monofenestratan.

Abstract: The Solnhofen pterosaurs Pterodactylus antiquus , Aerodactylus scolopaciceps , Diopecephalus kochi , Germanodactylus cristatus and Germanodactylus rhamphastinus all have complicated taxonomic histories. Species originally placed in the genus Pterodactylus , such as Aerodactylus scolopaciceps , Ardeadactylus longicollum , Cycnorhamphus suevicus and Germanodactylus cristatus possess apomorphies not observed in the type species of Pterodactylus , and consequently have been placed in new genera. The affinities of another Solnhofen pterosaur previously placed in Pterodactylus , Diopecephalus kochi , are less clear. It has been proposed that D. kochi is a juvenile specimen of Pterodactylus antiquus , or perhaps ‘ Germanodactylus rhamphastinus ’ specimens are mature examples of D. kochi . Furthermore, studies have suggested that ‘ Germanodactylus rhamphastinus ’ is not congeneric with the type species of Germanodactylus. Geometric morphometric analysis of prepubes and a cladistic analysis of the Pterosauria elucidate plesiomorphic and apomorphic conditions for basal Jurassic pterodactyloids. Germanodactylus is found to be a monotypic genus and Pterodactylus , Diopecephalus , and ‘ G. rhamphastinus ’ are found as distinct taxa belonging in individual genera, diagnosable using a combination of characters. Thus, Diopecephalus kochi is not demonstrated to be congeneric with Germanodactylus or Pterodactylus and is maintained as a valid taxon. ‘ G. rhamphastinus ’ is readily distinguishable from other Solnhofen pterosaur taxa, and a new genus is erected for its reception. Supplementary material: Technical details of the mechanical theory behind this article are available at https://doi.org/10.6084/m9.figshare.c.3809401.v1

Abstract: After being inaccessible for a number of years, the holotype and other specimens of the dsungaripterid pterodactyloid pterosaur Noripterus complicidens are again available for study. Numerous taxa assigned to the Dsungaripteridae have been described since the erection of Noripterus , but with limited comparisons to this genus. Based on the information from Young’s original material here we revise the taxonomic identity of N. complicidens and that of other Asian dsungaripterids. We conclude that N. complicidens is likely to be distinct from the material recovered from Mongolia and this latter material should be placed in a separate genus.