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: 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: Six specimens accessioned to the Bavarian State Collection for Palaeontology and Geology in Munich, Germany, in 1966 are identified as coming from a gigantic pterodactyloid pterosaur. The previously undescribed material was obtained in 1955 by Jean Otto Haas and compares favourably in size with the type specimen of the Late Cretaceous (Maastrichtian) azhdarchid pterosaur Arambourgiania philadelphiae (Arambourg 1959) from the same locality/region. The material represents fragments of two cervical vertebrae, a neural arch, a left femur, a ?radius, and a metacarpal IV and bones of problematic identity, and does not duplicate the type material of Arambourgiania . The timing of its collection and its locality of Ruseifa, Jordan suggest it might pertain to the same individual as the holotype.

Abstract: A pterosaur humerus (PRC 64) from the Upper Jurassic of Thailand was initially assigned to the Azhdarchoidea, an important clade of edentulous Cretaceous pterodactyloid pterosaurs. Here it is reassigned, on the basis of morphological comparisons and a phylogenetic analysis, to the Rhamphorhynchidae, a widely distributed clade of small to medium-sized seemingly piscivorous basal pterosaurs. This record extends the geographical range of rhamphorhynchids and confirms recent findings that these pterosaurs inhabited continental as well as marginal marine ecosystems.

Abstract: A specimen of a pterodactyloid pterosaur from the Upper Tunbridge Wells Sand Formation (Early Cretaceous, Valanginian) of Bexhill, East Sussex, southern England is described. It comprises a small fragment of jaw with teeth, a partial vertebral column and associated incomplete wing bones. The juxtaposition of the bones suggests that the specimen was originally more complete and articulated. Its precise phylogenetic relationships are uncertain but it represents an indeterminate lonchodectid with affinities to Lonchodectes sagittirostris (Owen 1874) which is reviewed here, and may belong in Lonchodraco Rodrigues & Kellner 2013 . This specimen is only the third record of pterosaurs from this formation.

Abstract Pterodactyls or pterosaurs, well-known flying reptiles of the Mesozoic, were already compared with dragons and vampires well before the discovery of the spectacularly large species from North America with wing spans of over 6 m. First described in 1784, they were not recognized as flying reptiles until 1801, when Baron Cuvier described a specimen that a few years later he called Ptero Dactyle which later became Pterodactylus . The name Pterodactylus is technically invalid – it is a junior synonym of Ornithocephalus Soemmerring 1812 – but it has stuck in the psyche of both palaeontologists and public alike. By the end of the nineteenth century numerous workers had compared pterosaurs with demons, dragons and vampires and life restorations had appeared in books, magazines and as gargoyles on the external architecture of the Natural History Museum, London. Sir Arthur Conan Doyle, famously the creator of Sherlock Holmes, was a polymath, with interests in science, sport, politics, travel, the occult and of course writing. He trained as, and became, a physician, with an eventually thriving general practice in Southsea, Hampshire from 1882 to 1890. In 1912, first as a series in Sunday magazines in the USA and in Strand Magazine in the UK, and shortly after as a hardback, he published The Lost World , an adventure story about the exploration of a South American tableland with prehistoric creatures that had persisted to the present. Although dinosaurs existed in this anachronistic fictional ecosystem, the ‘star’ animals were pterodactyls. Here we discuss the notoriety of pterodactyls generated by The Lost World , and hold Conan Doyle responsible for the widespread popularity of these iconic prehistoric reptiles right up to the present day.

Abstract The discovery of dinosaurs and other large extinct ‘saurians’, a term under which the Victorians commonly lumped ichthyosaurs, plesiosaurs, pterosaurs and their kin, makes exciting reading. The story of how early ‘fossilists’ first found the remains of these ‘primeval monsters’ has been told again and again in popular and semi-popular books about the history of palaeontology. Mary Anning making a living by collecting extinct reptiles along the Dorset coast, William Buckland and Gideon Mantell finding the ‘terrible lizards’ for which Richard Owen was to coin the word ‘Dinosauria’, O. C. Marsh and E. D. Cope fighting over new fossil vertebrates in the American West – all of these well-known stories have almost achieved the status of legends, and have often been retold with little regard for historical or scientific accuracy. The purpose of the present volume is not to retell these tales. The papers in this collection focus on relatively little-known episodes in the discovery and interpretation (from both a scientific and an artistic point of view) of dinosaurs and other Mesozoic animals. They cover a long time span, from the beginnings of scientific palaeontology to the present, and deal with many parts of the world, from the Yorkshire coast to central India, from Bavaria to the Sahara. The characters in these stories include professional palaeontologists and geologists (some of them well known, others more obscure), explorers, amateur fossil collectors and artists, linked together by their interest in Mesozoic creatures.

Abstract The first pterosaur fossil was described by Cosimo Alessandro Collini in 1784, but the epithet ptero dactyle was not applied until Georges Cuvier recognized the fossil as that of a volant animal in 1801. In eighteenth-century Britain, pterosaur bones had been discovered in Jurassic strata at Stonesfield, Oxfordshire but were considered to be bird bones, and largely went unnoticed. Bones of pterosaurs considerably larger than those of the first pterosaurs were discovered in the early nineteenth century by Gideon Mantell, but because of their comparatively large size were considered by Cuvier to also be the bones of birds. This perception by early nineteenth-century palaeontologists, including William Buckland and Gideon Mantell, that pterosaurs were relatively small animals was probably the reason their remains went unrecognized in British Jurassic and Cretaceous strata for several decades. Furthermore, the eighteenth- and early nineteenth-century dogmatic acceptance that fossil birds were present in the Jurassic Stonesfield ‘slate’ of Oxfordshire delayed the identification of medium-sized pterosaurs until the late 1820s, when Dean William Buckland described the Liassic Pterodactylus (= Dimorphodon ) macronyx in 1829. Even after that date many fragmentary, but large, pterosaur bones were misidentified as avian, despite there being no convincing evidence for Mesozoic birds until the discovery of Archaeopteryx in the 1860s. Truly gigantic pterosaurs were first discovered in Great Britain some 20 years before Pteranodon was found in the Late Cretaceous of Kansas. However, the British material was so fragmentary that it was easily eclipsed by the spectacular, near-complete skeletons of Pteranodon found by O. C. Marsh and others from the 1870s onwards.

Abstract An exceptionally well-preserved cranium and mandible of a new species of pterodacty-loid pterosaur from the Nova Olinda Member of the Crato Formation (Aptian, Early Cretaceious) of the Araripe Basin, northeastern Brazil, is described. The new taxon is characterized by the presence of a caudally directed parietal crest similar to that seen in pteranodontids, but is referred to the Ornithocheiridae of the Ornithocheiroidea. The specimen is referred to a new genus within the Ornithocheiridae, as it lacks the diagnostic rostral crest and instead possesses this parietal crest oriented. A lanceolate leaf with frayed distal end wedged between the mandibular rami suggests the cause of death for the specimen.

Abstract Two specimens of a tapejarid pterosaur (Pterodactyloidea, Tapejaridae) are described as representing a new species. Both specimens show evidence for soft tissues preserved in association with a sagittal bony cranial crest. Both specimens are from the Nova Olinda Member Konservat Lagerstätte of the Crato Formation of the Araripe Basin, northeastern Brazil. They represent the second tapejarid species from this formation. Comparisons are made with other crested pterosaurs and comments on the utility and aerodynamics of pterosaurian head crests are made.

Abstract New specimens of pterosaurs with soft-part preservation from the Solnhofen Lithographic Limestone (S Germany) and the Crato Formation (northeastern Brazil) yield hitherto unknown and unexpected details of pterosaur anatomy: the presence and internal anatomy of soft-tissue crests, the internal anatomy of the brachiopatagium, including a blood vessel system and structural details of foot and hand. Some consequences for pterosaurian flight, thermoregulation and aspects of evolution are discussed.

Abstract Among Pterosauria there are three types of scapulocoracoid construction. In the ornitho-cheirid scapulocoracoid the scapula is oriented almost horizontally; it is shorter than the coracoid and the glenoid fossa is level with the ventral margin of the vertebral column. In the azhdarchid scapulocoracoid the scapula is curved ventrally and is as long as the coracoid. In this construction the glenoid fossa lies approximately in the mid-horizontal plane of the chest. In the tapejaroid construction, the scapula is about one-third longer than the coracoid, which is oriented subhorizontally, and the glenoid fossa is level with the dorsal rim of the sternal plate. Both latter conditions are hitherto unknown among flying vertebrates and result in an unstable, but manoeuvrable flight, probably powered with wing beats.