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For over a century, there was a major gap in our understanding of the evolution of the flying Mesozoic reptiles, the pterosaurs, with a major morphological gap between the early forms and the derived pterodactyloids. 1 Recent discoveries have found a cluster of intermediate forms that have the head and neck of the pterodactyloids but the body of the early grade, 2 yet this still leaves fundamental gaps between these intermediates and both earlier and more derived pterosaurs. Here, we describe a new and large Jurassic pterosaur, Skiphosoura bavarica gen. et sp. nov., preserved in three dimensions, that helps bridge the gap between current intermediate pterosaurs and the pterodactyloids. A new phylogeny shows that there is a general progression of key characteristics of increasing head size, increasing length of neck and wing metacarpal, modification to the fifth toe that supports the rear wing membrane, and gradual reduction in tail length and complexity from earlier pterosaurs into the first pterodactyloids. This also shows a clear evolution of the increasing terrestrial competence of derived pterosaurs. Furthermore, this closes gaps between the intermediates and their ancestors and descendants, and it firmly marks the rhamphorhynchines and ctenochasmatid clades as, respectively, being the closest earliest and latest groups to this succession of transitional forms., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Pterosaurs, the first true flying vertebrates, played a crucial role in Mesozoic terrestrial ecosystems. However, our understanding of their ability to move around on the ground and, more broadly, their terrestrial paleoecology remains limited. Here, we demonstrate an unexpectedly high degree of variation in the hands and feet of pterosaurs, comparable with that observed in extant birds. This suggests that pterosaurs were adapted to a remarkably broad range of non-aerial locomotor ecologies. Small, early, long-tailed pterosaurs (non-pterodactyliforms) exhibit extreme modifications in their hand and foot proportions indicative of climbing lifestyles. By contrast, the hands and feet of later, short-tailed pterosaurs (pterodactyliforms) typically exhibit morphologies consistent with more ground-based locomotor ecologies. These changes in proportions correlate with other modifications to pterosaur anatomy, critically, the separation along the midline of the flight membrane (cruropatagium) that linked the hindlimbs, enabling a much more effective locomotory ability on the ground. Together, these changes map a significant event in tetrapod evolution: a mid-Mesozoic colonization of terrestrial environments by short-tailed pterosaurs. This transition to predominantly ground-based locomotor ecologies did not occur as a single event coinciding with the origin of short-tailed forms but evolved independently within each of the four principal radiations: euctenochasmatians, ornithocheiroids, dsungaripteroids, and azhdarchoids. Invasion of terrestrial environments by pterosaurs facilitated the evolution of a wide range of novel feeding ecologies, while the freedom from limitations imposed by climbing permitted an increase in body size, ultimately enabling the evolution of gigantism in multiple lineages., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

Pterosaurs evolved a broad range of body sizes, from small-bodied early forms with wingspans of mostly 1-2 m to the last-surviving giants with sizes of small airplanes. Since all pterosaurs began life as small hatchlings, giant forms must have attained large adult sizes through new growth strategies, which remain largely unknown. Here we assess wing ontogeny and performance in the giant Pteranodon and the smaller-bodied anurognathids Rhamphorhynchus , Pterodactylus and Sinopterus . We show that most smaller-bodied pterosaurs shared negative allometry or isometry in the proximal elements of the fore- and hindlimbs, which were critical elements for powering both flight and terrestrial locomotion, whereas these show positive allometry in Pteranodon . Such divergent growth allometry typically signals different strategies in the precocial-altricial spectrum, suggesting more altricial development in Pteranodon . Using a biophysical model of powered and gliding flight, we test and reject the hypothesis that an aerodynamically superior wing planform could have enabled Pteranodon to attain its larger body size. We therefore propose that a shift from a plesiomorphic precocial state towards a derived state of enhanced parental care may have relaxed the constraints of small body sizes and allowed the evolution of derived flight anatomies critical for the flying giants.

Exquisite discoveries and new interpretations regarding an enigmatic group of cursorial avemetatarsalians led to a new phylogenetic hypothesis regarding pterosaur affinities. Previously thought to be dinosaur precursors, lagerpetids are now considered to be the closest relatives to pterosaurs. This new hypothesis sheds light on a new explorable field, especially regarding the character acquisition and evolution within the pterosaur lineage. In the present study, the morphospace occupation of distinct skeletal regions of lagerpetids within the morphological spectrum of avemetatarsalians is investigated. This approach indicates which portions of the skeleton are more similar to the anatomy of pterosaurs and which portions present different homoplastic signals. The analyses demonstrate that the craniomandibular traits of lagerpetids are pterosaur-like, the pectoral girdle and forelimb are dinosauromorph-like and the axial skeleton and the pelvic girdle and hindlimb are unique and highly specialized among the analyzed sample. So, despite the close phylogenetic relationships, the postcranial skeleton of lagerpetids and pterosaurs are very different. The occurrence of two distinct and highly specialized groups of pterosauromorphs coexisting with a wide ecological range of dinosauromorphs during the Triassic suggests pressure for new niches occupation., (© 2022 American Association for Anatomy.)

Studies of soft tissue effects on joint mobility in extant animals can help to constrain hypotheses about joint mobility in extinct animals. However, joint mobility must be considered in three dimensions simultaneously, and applications of mobility data to extinct taxa require both a phylogenetically informed reconstruction of articular morphology and justifications for why specific structures' effects on mobility are inferred to be similar. We manipulated cadaveric hip joints of common quail and recorded biplanar fluoroscopic videos to measure a 'ligamentous' range of motion (ROM), which was then compared to an 'osteological' ROM on a ROM map. Nearly 95% of the joint poses predicted to be possible at the hip based on osteological manipulation were rendered impossible by ligamentous constraints. Because the hip joint capsule reliably includes a ventral ligamentous thickening in extant diapsids, the hip abduction of extinct ornithodirans with an offset femoral head and thin articular cartilage was probably similarly constrained by ligaments as that of birds. Consequently, in the absence of extraordinary evidence to the contrary, our analysis casts doubt on the 'batlike' hip pose traditionally inferred for pterosaurs and basal maniraptorans, and underscores that reconstructions of joint mobility based on manipulations of bones alone can be misleading., (© 2018 The Author(s).)

The Istiodactylidae is one of three ornithocheiroid pterosaur lineages, characterised by a large nasoantorbital fenestrae and labiolingually compressed teeth. Their remains have been reported from Early Cretaceous rocks of northeastern China and Western Europe. A new istiodactylid pterosaur from the upper part of the Yixian Formation (Jingangshan Member) of the Jehol Group, distributed in western Liaoning Province and neighbouring areas, is described on the basis of an incomplete specimen including the partial skull, forelimb and hindlimb bones, and pelvis. This individual, here designated as a referred specimen of Nurhachius (Nurhachius sp.) is a skeletally almost mature pterosaur with an estimated wingspan of 1.6 m. This specimen is not only the geologically oldest known occurrence of Nurhachius, but also a new component of the pterosaur assemblage of the Jingangshan Member, and provides new information on the pelvis and hindlimb of istiodactylids and the palaeobiology of this clade. Considering previous reports of the istiodactylid remains and the geological ages of the strata that yielded them, it is suggested that different istiodactylids existed in Europe and China during the late Barremian. [ABSTRACT FROM AUTHOR]

Pterosaurs, flying reptiles that lived during the Mesozoic era, were the first vertebrates to develop powered flight. Despite being patchy, the pterosaur fossil record extends worldwide, being quite diverse at the Araripe Basin, Brazil. The Romualdo Formation (Aptian-Albian), a Konservat Lagerstätten rich in calcareous concretions, has yielded several of these exceptionally well-preserved, three-dimensional fossils. MN 4727-V, the focus of the present study, consists exclusively of three-dimensionally preserved post-cranial axial and appendicular elements. The anatomy suggests that MN 4727-V is a pteranodontoid since it exhibits an enlarged, warped deltopectoral crest and a subtriangular distal articulation of the humerus. The phylogenetic analysis recovered MN 4727-V as an anhanguerid by the combination of the following characters: scapula substantially shorter than the coracoid (sca/co < 0.80), and posterior expansion of the sternal articulation of the coracoid. The presence of elliptical-shaped, dorsomedially inclined depressions on the dorsal surface of the prezygapophyses of the preserved cervicals has never been reported for any other anhanguerid, although they might not be diagnostic to MN 4727-V. The tail bears unique middle caudals that exhibit ventral projections, a character previously reported only for Anhanguera piscator. The wingspan of MN 4727-V, a juvenile individual, was estimated to be about 3.9 metres. [ABSTRACT FROM AUTHOR]

The pterosaur fossil record of Argentina is increasing in recent times, both in the number of localities and their temporal range. The new materials are found in levels that span from the Late Triassic to the Late Cretaceous in age. In this contribution we describe twenty isolated pterosaur teeth from the Cerro de los Leones locality (Picún Leufú, Neuquén Province), where the Lohan Cura Formation (Albian) widely crops out. A new, detailed analysis of these remains allowed us to detect a set of morphological features, such as the enamel texture, cross-section shape, crown curvature, and absence of sharp carinae, which suggests close affinities with Ornithocheiriformes. Moreover, with the large number of tooth specimens, a discrimination of three morphotypes based on the crown curvature, the basal cross-section shape, and the presence of carinae was carried out. We performed extensive morphological comparisons between different members of Ornithocheiriformes with the aim to test the hypothesis that the presence of these different morphotypes reflects distinct anatomical positions of the teeth in the jaws . Thus, considering the heterodont dentition of ornithocheiriforms, the tooth morphological classification proposed here represents a new methodology, not only to identify isolated ornithocheiriform teeth, but also to assign them to the rostral, medial, or the posterior portion of the tooth row. Finally, the evidence of ornithocheiriform pterosaurs in Cerro de los Leones has paleobiogeographical implications, allowing us to correlate it with other Albian faunas from South America, expanding our knowledge of the distribution of Early Cretaceous pterosaurs across southwestern Gondwana. [ABSTRACT FROM AUTHOR]

This article discusses the discovery of pterosaur teeth in the Albian deposits of Kaniv, Ukraine. The teeth belong to the clade Anhangueria, but due to their heterodont dentition and poor preservation, it is difficult to assign them to a specific genus or species. This finding is significant as it represents the first known occurrence of pterosaurs in Ukraine and contributes to the understanding of pterosaur diversity and biogeography in Eastern Europe. Further exploration of the area may yield more pterosaur and dinosaur material. [Extracted from the article]

Fragmentary but scientifically important pterosaur fossils have been reported in Lower Cretaceous strata such as the Jinju and Hasandong formations of South Korea. Recently, several large teeth and a second wing phalanx found in these strata have been suggested to have affinities with Boreopteridae, a clade in which has so far been reported only in the Yixian Formation of China. Here, the phylogenetic affinities of these proposed boreopterid specimens are reassessed, using two-dimensional geometric morphometric shape analyses. The morphometric analyses found the South Korean pterosaur teeth in different morphospaces from the Yixian Fm. boreopterids, and plot them closer to anhanguerians. An isolated second wing phalanx does plot close to a boreopterid Zhenyuanopterus longirostris in a morphospace defined by first two principal component axes, but taxa from different lineages are also plotted as closely, or even closer. Thus, there seems to be no clear evidence that these specimens belong to Boreopteridae, and the tooth fossils appear to be very different from those of established members of this clade. Therefore, the hypothesis that some South Korean pterosaur fossils can be referred to boreopterids, is here tentatively rejected [ABSTRACT FROM AUTHOR]

Three ichnospecies of the Late Jurassic trace fossil genus Rhamphichnus were first described and interpreted as the walking tracks of non-pterodactyloid pterosaurs like Rhamphorhynchus. This despite not resembling any previously identified pterosaur tracks and having a morphology unmatched by pterosaur skeletal elements. To fit the pterosaurian model, elongated imprints of R. pereiraensis and R. lafaurii , displaying four or five digit impressions were interpreted as manus prints, while shorter, five-digit prints were interpreted as imprints made by pedes with dislocated metatarsals and disarticulated phalanges. Comparison of published images and descriptions of Rhamphichnus ispp. to skeletal elements of six contemporaneous non-pterodactyloid taxa, skeletons and tracks from pterodactyloid pterosaurs, and modern and ancient lepidosaurs and crocodilians suggests an alternative interpretation. R. crayssacensis closely resembles crocodilian (Crocodylopodus (Sustenodactylus) isp.) and crocodylomorph (Batrachopus isp.) tracks; its holotype trackway was found 50 cm away from and parallel to a crocodilian swimming trackway with the same sized feet, possibly made by the same individual. R. pereiraensis and R. lafaurii strongly resemble lepidosaurian tracks and match the size and morphology of contemporary rhynchocephalians. Prints originally interpreted as manual and pedal in R. pereiraensis and R. lafaurii are reinterpreted and transposed in light of new data from pterosaurian, lepidosaurian, and crocodilian tracks and anatomy. Identification of fossil trackmakers depends on comparisons of a wide range of candidate taxa and elimination of those that cannot be considered further. With this reinterpretation of Rhamphichnus , there are no non-pterodactyloid pterosaurian tracks yet identified in the fossil record. [ABSTRACT FROM AUTHOR]

Studies of soft tissue effects on joint mobility in extant animals can help to constrain hypotheses about joint mobility in extinct animals. However, joint mobility must be considered in three dimensions simultaneously, and applications of mobility data to extinct taxa require both a phylogenetically informed reconstruction of articular morphology and justifications for why specific structures' effects on mobility are inferred to be similar. We manipulated cadaveric hip joints of common quail and recorded biplanar fluoroscopic videos to measure a 'ligamentous' range of motion (ROM), which was then compared to an 'osteological' ROM on a ROM map. Nearly 95% of the joint poses predicted to be possible at the hip based on osteological manipulation were rendered impossible by ligamentous constraints. Because the hip joint capsule reliably includes a ventral ligamentous thickening in extant diapsids, the hip abduction of extinct ornithodirans with an offset femoral head and thin articular cartilage was probably similarly constrained by ligaments as that of birds. Consequently, in the absence of extraordinary evidence to the contrary, our analysis casts doubt on the 'batlike' hip pose traditionally inferred for pterosaurs and basal maniraptorans, and underscores that reconstructions of joint mobility based on manipulations of bones alone can be misleading.

In the palaeogeographical context of the opening of the South Atlantic Ocean at the end of the Early Cretaceous, we document here the first microvertebrate fauna recorded from the Aptian deposits of Gadoufaoua, Niger. A systematic study of the fauna has resulted in a significant expansion of the existing faunal list and increase our knowledge of the palaeobiodiversity of the Gadoufaoua fossil site. Some taxa were previously recorded (lungfishes, crocodilians, chelonians, etc.), but several new taxa are described here for the first time, including the first occurrence of a stem-boreosphenidan mammal in Africa for this time-interval. In addition, chondrichthyans, pterosaurs and lissamphibians are documented. The analysis of taxonomic diversity and preservation of the fossils confirms a floodplain depositional environment (more precisely channel-type), in a higher velocity flow regime than previously thought based on studies of the macrofauna. To confirm the depositional environment inferred by the state of preservation of the fossils, a preliminary comparison with the fauna of similar age from the Santana Formation (Brazil) has been undertaken. The hypothesis of a communication between Africa and South America during the Aptian, as had already been demonstrated based on studies of the macrofauna, is strengthened. [ABSTRACT FROM AUTHOR]

Here, we describe the first pterosaur remains from Angola, an assemblage of fourteen bones from the Lower Maastrichtian marine deposits of Bentiaba, Namibe Province. One new species is introduced, Epapatelo otyikokolo, gen. et sp. nov., which comprises an articulated partial left humerus and ulna as well as an articulated left ulna and radius (from a second individual). Phylogenetic analysis confirms a non-nyctosaurid pteranodontian attribution for this new taxon and supports a new apomorphy-based clade, Aponyctosauria, which is here defined. Late Cretaceous pteranodontians are rare in Sub-Saharan Africa and throughout the Southern Hemisphere. Preliminary histological analysis also reveals a likely sub-adult age for one of the specimens. This fossil assemblage provides a first glimpse of Angolan pterosaur paleobiodiversity providing further insight into the Gondwanan ecosystems of the Upper Cretaceous. [ABSTRACT FROM AUTHOR]

Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus, which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals. [ABSTRACT FROM AUTHOR]

Here we describe a new pterosaur footprint assemblage from the Hwasun Seoyuri tracksite in the Upper Cretaceous Jangdong Formation of the Neungju Basin in Korea. The assemblage consists of many randomly oriented prints in remarkably high densities but represents a single ichnotaxon, Pteraichnus. Individuals exhibit a large but continuous size range, some of which, with a wingspan estimated at 0.5 m, are among the smallest pterosaurs yet reported from the Upper Cretaceous, adding to other recent finds which contradict the idea that large and giant forms entirely dominated this interval. Unusual features of the tracks, including relatively long, slender pedal digit impressions, do not match the pes of any known Cretaceous pterosaur, suggesting that the trackmakers are as yet unknown from the body fossil record. The Hwasun pterosaur footprints appear to record gregarious behavior at the exact location by individuals of different ages, hinting at the possibility that pterosaurs gathered in mixed-age groups. [ABSTRACT FROM AUTHOR]

We describe fossil liverworts, mosses, and thallose forms of unknown affinity from the Barreal and Cortaderita formations (Sorocayense Group, Middle Triassic) in the Barreal depocenter, southwestern San Juan Province. The specimens are impressions-compressions of gametophytes and sporophytes, preserved in claystones, siltstones, and tuffaceous and bentonitic silty sandstones. The thallose forms consist of dichotomous simple thalli lacking evidence of a midrib and are assigned to the genus Thallites. Within the liverworts, we identified a new genus and two new species. Ricciellites unsaltoensis gen. et sp. nov. is characterized by thalli forming partial rosettes with distinct midrib, polygonal cells, air pores surrounded by a ring of cells, enclosed sporophytes, and ventral scales arranged in two rows. Ricciopsis cortaderitaensis sp. nov. has thalli forming rosettes, with distinct midrib, polygonal cells, and rhizoids. We also describe two types of acrocarpous mosses assignable to the genus Muscites. They are characterized by delicate unbranched axes with lanceolate unicostate leaves. Muscites sp. 1 is represented by several specimens arranged in a loose cushion, while Muscites sp. 2 corresponds to isolated specimens. Together, these fossils represent the first unequivocal evidence of mosses and liverworts for the Middle Triassic of Argentina. Given the scarcity of bryophyte megafossils of Triassic age worldwide (31 worldwide, three confirmed records for Argentina), these findings add significantly to the bryophyte fossil record. [ABSTRACT FROM AUTHOR]

Knowledge about the pterosaur diet and digestive system is limited, and there is little direct evidence in the fossil record. Here, we report two specimens of the wukongopterid Kunpengopterus sinensis, a juvenile and an adult, from the Late Jurassic Yanliao Biota of China with associated bromalites. Both of these concentrations are identified as emetolites, fossilized gastric pellets. These pellets contain scales of an unnamed palaeonisciform fish, confirming the pterosaur was a piscivore. It probably vomited the pellets, indicating the presence of two-part stomachs and efficient antiperistalsis in both juveniles and adults. Comparing the ganoid scales found in the pellets with those of complete fishes, it was possible to determine that the prey of the smaller pellet is an average-sized individual, while the prey of the larger pellet represents a large specimen. Kunpengopterus sinensis might have preyed on the same fish during ontogeny, with adults being able to feed on larger individuals. [ABSTRACT FROM AUTHOR]

The Khok Pha Suam locality in the province of Ubon Ratchathani, northeastern, Thailand, is known as "the last home of Thai dinosaurs", because it belongs to the Lower Cretaceous Khok Kruat Formation (Aptian-Albian) which is currently the youngest Mesozoic vertebrate fossil producing formation in the Khorat Group. Here, we describe a diverse vertebrate assemblage, including hybodonts, ray-finned fishes, turtles, crocodyliforms, pterosaurs, and dinosaurs from the Khok Pha Suam locality. The updated data on the Khok Kruat fauna provides a better understanding of the variety and distribution of Early Cretaceous continental ecosystems, which are useful for palaeoenvironmental reconstruction. In addition to consolidating unincorporated data on fauna, this study also provides the palaeontological data necessary to illustrate the palaeoecosystem to the general public, as well as improving the academic value of the Pha Chan-Sam Phan Bok Geopark. [ABSTRACT FROM AUTHOR]

The pterosaur fossil record in Portugal is scarce, comprising mainly isolated teeth and rare postcranial material. Here, we describe a well-preserved right proximal femur of a pterodactyloid pterosaur from the Kimmeridgian, Upper Jurassic Praia da Amoreira-Porto Novo Formation of Peniche, Portugal. It is noteworthy for its relatively large size, compared to other Jurassic pterosaurs. It shows affinities with dsungaripteroids based on a combination of features including the bowing of the shaft, the mushroom-like cap of the femoral head, and the distinctly elevated greater trochanter. The femur has a relatively thinner bone wall compared to dsungaripterids, and is more similar to basal dsungaripteroids. A histological analysis of the bone cortex shows it had reached skeletal maturity. The preserved last growth period indicates fast, uninterrupted growth continued until the final asymptotic size was reached, a growth pattern which could best be compared to pterodactyloid femora from the Early Cretaceous. The specimen is the second confirmed report of a dsungaripteroid from the Jurassic, and it is the first record of this group from the Iberian Peninsula. [ABSTRACT FROM AUTHOR]

The record of Cretaceous pterosaur remains from Germany is sparse. The material recovered to date includes the fragmentary holotypes of Targaryendraco wiedenrothi and Ctenochasma roemeri, as well as a few isolated pterodactyloid teeth and some indeterminate skeletal elements, together with a plaster cast of a large Purbeckopus manus imprint. Here, we report the discovery of a pterodactyloid pterosaur mandible from lower Valanginian strata of the Stadthagen Formation in the Lower Saxony Basin of Northern Germany. Based on the size and spacing of its alveoli, this fossil is attributable to the cosmopolitan Early Cretaceous pteranodontoid clade Anhangueria. Moreover, it represents the first and only known pterosaur from the Valanginian of Germany and is one of only a handful Valanganian pterosaur occurrences presently recognized worldwide. In addition to the approximately coeval Coloborhynchus clavirostris from the Hastings Bed Group of southern England, the Stadthagen Formation pterosaur mandible is among the stratigraphically oldest identifiable anhanguerians. [ABSTRACT FROM AUTHOR]

Pterosaur fossils from Australia are exceptionally rare. Since the discovery of the continent's first pterosaur some 40 years ago, fewer than 20 specimens have been described. The Lower Cretaceous (upper Albian) Toolebuc Formation of North West Queensland is the most productive horizon for Australian pterosaurs. Herein, we describe a new species of pterosaur, Thapunngaka shawi gen. et sp. nov., from the Toolebuc Formation, near Richmond, North West Queensland. The specimen (KKF494) comprises the rostral portion of a crested mandible and represents the largest pterosaur yet described from Australia. The new species presents features that indicate an affinity with Anhangueridae, which is consistent with their reported cosmopolitan distribution during this period. Thapunngaka shawi can be distinguished from other anhanguerids through the possession of a mandible with a smooth dorsal surface medially and uniquely sized alveoli that are positioned laterally along the jaw. Phylogenetic analysis reveals a close relationship among all Australian anhanguerids and points to an endemic Australian radiation within Anhangueridae. Thapunngaka shawi has the largest mandibular crest of any anhanguerian worldwide, and provides further evidence for the existence of an increasingly diverse range of large crested pterosaurs in the Australian part of eastern Gondwana during the Cretaceous. [ABSTRACT FROM AUTHOR]

The pterosaurs first appear in the fossil record in the middle of the Late Triassic. Their earliest representatives are known from Northern Hemisphere localities but, by the end of the Jurassic Period, this clade of flying reptiles achieved a global distribution, as well as high levels of diversity and disparity. Our understanding of early pterosaur evolution and the fundamental interrelationships within Pterosauria has improved dramatically in recent decades. However, there is still debate about how the various pterosaur subgroups relate to one another and about which taxa comprise these. Many recent phylogenetic analyses, while sampling well from among the known Triassic and Early Jurassic pterosaurs, have not included many non-pterosaurian ornithodirans or other avemetatarsalians. Given the close relationship between these groups of archosaurs, the omission of other ornithodirans and avemetatarsalians has the potential to adversely affect the results of phylogenetic analyses, in terms of character optimisation and ingroup relationships recovered. This study has addressed this issue and tests the relationships between the early diverging pterosaur taxa following the addition of avemetatarsalian taxa and anatomical characters to an existing early pterosaur dataset. This study has, for the first time, included taxa that represent the aphanosaurs, lagerpetids, silesaurids and dinosaurs, in addition to early pterosaurs. Anatomical characters used in other recent studies of archosaurs and early dinosaurs have also been incorporated. By expanding the outgroup taxa and anatomical character coverage in this pterosaur dataset, better resolution between the taxa within certain early pterosaur subclades has been achieved and stronger support for some existing clades has been found; other purported clades of early pterosaurs have not been found in this analysis--for example there is no support for a monophyletic Eopterosauria or Eudimorphodontidae. Further support has been found for a sister-taxon relationship between Peteinosaurus zambelli and Macronychoptera, a clade here named Zambellisauria (clade nov.), as well as for a monophyletic and early diverging Preondactylia. Some analyses also support the existence of a clade that falls as sister-taxon to the zambellisaurs, here named Caviramidae (clade nov.). Furthermore, some support has been found for a monophyletic Austriadraconidae at the base of Pterosauria. Somewhat surprisingly, Lagerpetidae is recovered outside of Ornithodira sensu stricto, meaning that, based upon current definitions at least, pterosaurs fall within Dinosauromorpha in this analysis. However, fundamental ornithodiran interrelationships were not the focus of this study and this particular result should be treated with caution for now. However, these results do further highlight the need for broader taxon and character sampling in phylogenetic analyses, and the effects of outgroup choice on determining ingroup relationships. [ABSTRACT FROM AUTHOR]

The current understanding of UK Middle Jurassic pterosaur taxonomy is under-developed, leading to it being previously considered a time of low diversity. This is despite the presence of a productive but under-studied pterosaur-bearing horizon extending over parts of Oxfordshire and Gloucestershire. This unit, informally called the Stonesfield Slate, is part of the Great Oolite Group and it produces the largest number of Middle Jurassic pterosaurs. There are over 200 specimens distributed across museums in the United Kingdom, America, and Australia, almost all of which are accessioned under the genus Rhamphocephalus and referred to three species: the type species Rhamphocephalus prestwichi, Rhamphocephalus bucklandi, and Rhamphocephalus depressirostris. This study reviews the British Middle Jurassic Pterosauria assemblage, evaluating both their systematics and taxonomic diversity. The holotype of Rhamphocephalus, an isolated skull table, is found to be a misidentified crocodylomorph skull and the genus is considered a nomen dubium. The holotype of Rhamphocephalus bucklandi is identified as missing and that of Rhamphocephalus depressirostris has characters diagnostic at a family level, not a generic or specific one. Both species are considered dubious. Detailed examination of the entire assemblage shows that rather than being monogeneric, the assemblage contains at least five pterosaur taxa, representing three families. This diversity includes the potential earliest occurrences of both Monofenestrata and Pterodactyloidea. A new genus, Klobiodon rochei gen. et sp. nov. is described based on a well-preserved mandible. The English Bathonian pterosaur assemblage is shown to be diverse and indicates that, as has been suggested in other studies, the low-diversity signal in the Middle Jurassic is at least partially artificial. [ABSTRACT FROM AUTHOR]

Pterosaurs are the second most abundant fossil vertebrates in the Araripe Basin, Northeastern Brazil. As a matter of fact, this sedimentary unit has produced some of the best-preserved pterosaur fossils worldwide. In spite of that, most Araripe pterosaur fossils lack collection data, including precise geographic coordinates and fine-scale stratigraphic procedence. Here we describe the first pterosaur fossils from the Sobradinho site (Ceará state, Northeast Brazil). The new specimens have full collection data, being composed by exceptionally-preserved, uncompressed bones. Preserved materials include one humerus, two scapulae, two coracoids, one vertebra, one rib, and a fragment of the pelvic girdle with diagnostic features of the Pteranodontoidea. Aside from the morphological description of the new specimen, we also discuss its ontogeny, associated fauna, and the importance of geographic and stratigraphically controlled pterosaur collections, especially within the context of the Araripe Basin. [Display omitted] • Morphological description of new pterosaurs from the Romualdo Formation. • Specimens with geographic and stratigraphic procedence. • First pterosaurs described for the Sobradinho site. [ABSTRACT FROM AUTHOR]

Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeo-diversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group's evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non-significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large-bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic-Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution. [ABSTRACT FROM AUTHOR]

An apparently nearly complete and fairly well-articulated specimen of a large toothed pterosaur from the Lower Cretaceous of Brazil is exhibited at the CosmoCaixa - Museo de la Ciencia, Barcelona, Spain. The museum label refers it to Anhanguera piscator and reports a provenance from the Crato Formation. Its study shows that it is from the Romualdo Formation and is an assembled and composite specimen over 45% reconstructed. The original skeletal elements show no diagnostic feature of Anhanguera piscator, belong to non-tapejarid pterosaurs, and do not furnish further significative scientific information. Several mistakes were made in the reconstruction and assemblage of the skeleton and some elements do not follow the proportions of Anhanguera piscator. Care is suggested when describing pterosaur material from the Romualdo Formation prepared and sold by private fossil dealers in the absence of an adequate documentation on the original material and its preparation. [ABSTRACT FROM AUTHOR]

A pterosaur wing phalange from the Kimmeridgian of Eathie (Ross and Cromarty) is identified as the first pterosaur to have been collected in Scotland. The specimen was acquired by the British Museum (Natural History) in 1888, but an old collector's label gives the date of collection as 1850. The handwriting matches that of Charles W. Peach (1800-1886), who later became better known for his fossil fish and plant collections, which still exist in the Natural History Museum, London, and other museums. Peach collected this pterosaur specimen during his first year of working in Scotland, and it was probably one of his earliest finds of north of the border. Pterosaur remains are exceptionally rare in Scotland and otherwise unknown from this particular site, so this newly recognised specimen is a significant addition to the Scottish fossil record. It is also among the earliest UK pterosaur finds, post-dating Mary Anning's discovery of Dimorphodon by just over two decades. [ABSTRACT FROM AUTHOR]

The biogeographical history of pterosaurs has received very little treatment. Here, we present the first quantitative analysis of pterosaurian biogeography based on an event-based parsimony method (Treefitter). This approach was applied to a phylogenetic tree comprising the relationships of 108 in-group pterosaurian taxa, spanning the full range of this clade's stratigraphical and geographical extent. The results indicate that there is no support for the impact of vicariance or coherent dispersal on pterosaurian distributions. However, this group does display greatly elevated levels of sympatry. Although sampling biases and taxonomic problems might have artificially elevated the occurrence of sympatry, we argue that our results probably reflect a genuine biogeographical signal. We propose a novel model to explain pterosaurian distributions: pterosaurs underwent a series of 'sweep-stakes' dispersal events (across oceanic barriers in most cases), resulting in the founding of sympatric clusters of taxa. Examination of the spatiotemporal distributions of pterosaurian occurrences indicates that their fossil record is extremely patchy. Thus, while there is likely to be genuine information on pterosaurian diversity and biogeographical patterns in the current data-set, caution is required in its interpretation. [ABSTRACT FROM AUTHOR]

The pterosaur fossil record from Africa is exceedingly scarce and one of the least known for any continental land mass. The specimens here described are housed at the Naturkundemuseum of the Humboldt University and consist of two cervical vertebrae, a coracoid and a wing metacarpal recovered from the Upper Jurassic Tendaguru Formation, Tanzania. Due to the general morphology and the absence of a lateral pneumatic foramen in both vertebrae, as well as the presence of a longitudinal depression, not previously reported in pterosaurs, we consider these specimens as representatives of a new species of Azhdarchidae. Moreover, because the coracoid, which bears three well-developed pneumatic foramina, has a well-excavated depression that is medially positioned at the posterior face of the acrocoracoid process, we regard this as a new basal pterodactyloid species. The wing metacarpal is greatly elongated and clearly belongs to Pterodactyloidea. Its elongation and slender aspect, as well as the sub-triangular shape of its proximal articular end, likely place it within the Tapejaroidea. The material here described shows the potential of these deposits to provide more informative pterosaur material and provisionally extends the oldest record of azhdarchids to the Kimmeridgian-Tithonian of Africa. [ABSTRACT FROM AUTHOR]

Five pterosaur localities are currently known from the Late Cretaceous in the northeastern Aral Sea region of Kazakhstan. Of these, one is Turonian-Coniacian in age, the Zhirkindek Formation (Tyulkili), and four are Santonian in age, all from the early Campanian Bostobe Formation (Baibishe, Akkurgan, Buroinak, and Shakh Shakh). All so far collected and identifiable Late Cretaceous pterosaur bones from Kazakhstan likely belong to Azhdarchidae: Azhdarcho sp. (Tyulkili); Aralazhdarcho bostobensis (Shakh Shakh); and Samrukia nessovi (Akkurgan). These latter two taxa, both from the Bostobe Formation might be synonyms. Azhdarcho sp. from the Zhirkindek Formation lived in a tropical-to-subtropical relatively humid climate on the shore of an estuarine basin connected to the Turgai Sea. Known fossils were collected in association with brackish-water bivalves and so the overall paleoenvironment of this pterosaur was likely an estuarine marsh as indicated by the dominance of conifers and low relative counts of ferns and angiosperms. Aralazhdarcho bostobensis, from the Bostobe Formation, lived on a coastal fluvial plain along the Turgai Sea. This paleoenvironment was either floodplain (Akkurgan, Buroinak, and Shakh Shakh) or estuarine (Baibishe). In the Santonian -- early Campanian, shallow waters near this coastal plain were sites for the intensive accumulation of phosphates under upwelling conditions caused by strong winds from the ancient Asian landmass. These winds also caused significant aridization of the climate during this time. We speculate that pterosaurs may have been attracted to this area by the abundant resources in the bioproductive estuaries and nearshore upwelling waters. [ABSTRACT FROM AUTHOR]

The Early Cretaceous Jehol Biota of northeastern China has become famous over the last two decades as a source of feathered avialan and non-avialan theropods, preserved alongside an array of other fossil vertebrates, invertebrates, and plants. Still more recently, a rich assemblage referred to in this paper as the Daohugou Biota has begun to emerge from Jurassic strata in the same region. Like their counterparts from the Jehol Biota, Daohugou Biota vertebrate specimens are typically preserved in fine-grained lacustrine beds and often retain feathers and other soft-tissue features. At present, 30 vertebrate taxa (five salamanders, one anuran, two lizards, 13 pterosaurs, five dinosaurs, and four mammals) are known from the Daohugou Biota, which was first recognized at the Daohugou locality in Inner Mongolia. The presence of the salamanderChunerpeton tianyiensis, proposed in this paper as an index fossil for the Daohugou Biota, links the Daohugou locality to five other fossil-producing areas in the provinces of Hebei and Liaoning. The strata containing the Daohugou Biota are close to the Middle–Upper Jurassic boundary and belong at least partly to the regionally widespread Tiaojishan Formation. In general, the vertebrate fauna of the Daohugou Biota is strikingly different from that of the Jehol Biota, although paravian dinosaurs, anurognathid pterosaurs, and salamanders with cryptobranchid and hynobiid affinities occur in both. Nevertheless, the Daohugou Biota and the Jehol Biota are two successive Lagerstätte assemblages that collectively offer a taphonomically consistent window into the Mesozoic life of northeast Asia over a significant span of geologic time. [ABSTRACT FROM PUBLISHER]

The article offers a study of a Brazilian fossil that suggests that pterosaurs were also clad in tiny feathers of varying colors. The feathers may have evolved more than 150 million years. The colorful feathers were presumed for signaling. It reinforces the idea that pterosaurs were fluffy, and some of them probably had complex, colorful patterns.

Discoveries of Jurassic pterosaurs in the Southern Hemisphere are extremely unusual. In Argentina, pterosaurs from the Upper Jurassic (Tithonian) have only been found in the Northwest of Patagonia (Neuquén Basin). These come from marine deposits and three specimens have been discovered up to the present. In this paper, we report a new finding from the Neuquén Basin. This material is identified as a tibiotarsus, which probably belonged to an osteologically adult individual and represents a new species of a pterodactyloid pterosaur of medium size. This discovery provides new evidence that at least two different species of pterodactyloid pterosaurs may have coexisted in Los Catutos Member, Vaca Muerta Formation, from the shallow marine deposits of the Neuquén Basin. [ABSTRACT FROM AUTHOR]

Extinct animal behavior has often been inferred from qualitative assessments of relative brain region size in fossil endocranial casts. For instance, flight capability in pterosaurs and early birds has been inferred from the relative size of the cerebellar flocculus, which in life protrudes from the lateral surface of the cerebellum. A primary role of the flocculus is to integrate sensory information about head rotation and translation to stabilize visual gaze via the vestibulo-occular reflex (VOR). Because gaze stabilization is a critical aspect of flight, some authors have suggested that the flocculus is enlarged in flying species. Whether this can be further extended to a floccular expansion in highly maneuverable flying species or floccular reduction in flightless species is unknown. Here, we used micro computed-tomography to reconstruct “virtual” endocranial casts of 60 extant bird species, to extract the same level of anatomical information offered by fossils. Volumes of the floccular fossa and entire brain cavity were measured and these values correlated with four indices of flying behavior. Although a weak positive relationship was found between floccular fossa size and brachial index, no significant relationship was found between floccular fossa size and any other flight mode classification. These findings could be the result of the bony endocranium inaccurately reflecting the size of the neural flocculus, but might also reflect the importance of the flocculus for all modes of locomotion in birds. We therefore conclude that the relative size of the flocculus of endocranial casts is an unreliable predictor of locomotor behavior in extinct birds, and probably also pterosaurs and non-avian dinosaurs. [ABSTRACT FROM AUTHOR]

Abstract: Stomach contents preserved in fossil specimens provide direct evidence for the diet of extinct animals. Such exceptional fossils remain rare for predatory non-avian dinosaurs and each can add significantly to our understanding of trophic interactions between various taxa. Here we present evidence for the dromaeosaurid theropod Velociraptor scavenging on the carcass of an azhdarchid pterosaur, with a long bone of the pterosaur being found as gut contents of the dinosaur. Despite previous inferences of dromaeosaurs as hyper-predators, scavenging appears to have been an important part of their ecology. [Copyright &y& Elsevier]

Hone, D.W.E., Naish, D. & Cuthill, I.C. 2011: Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs? Lethaia, Vol. 45, pp. 139-156. Cranial ornamentation is widespread throughout the extinct non-avialian Ornithodira, being present throughout Pterosauria, Ornithischia and Saurischia. Ornaments take many forms, and can be composed of at least a dozen different skull bones, indicating multiple origins. Many of these crests serve no clear survival function and it has been suggested that their primary use was for species recognition or sexual display. The distribution within Ornithodira and the form and position of these crests suggest sexual selection as a key factor, although the role of the latter has often been rejected on the grounds of an apparent lack of sexual dimorphism in many species. Surprisingly, the phenomenon of mutual sexual selection - where both males and females are ornamented and both select mates - has been ignored in research on fossil ornithodirans, despite a rich history of research and frequent expression in modern birds. Here, we review the available evidence for the functions of ornithodiran cranial crests and conclude that mutual sexual selection presents a valid hypothesis for their presence and distribution. The integration of mutual sexual selection into future studies is critical to our understanding of ornithodiran ecology, evolution and particularly questions regarding sexual dimorphism. □ Behaviour, Dinosauria, ornaments, Pterosauria, sexual selection. [ABSTRACT FROM AUTHOR]

Associations of large vertebrates are exceedingly rare in the Late Jurassic Solnhofen Limestone of Bavaria, Southern Germany. However, there are five specimens of medium-sized pterosaur Rhamphorhynchus that lie adjacent to the rostrum of a large individual of the ganoid fish Aspidorhynchus. In one of these, a small leptolepidid fish is still sticking in the esophagus of the pterosaur and its stomach is full of fish debris. This suggests that the Rhamphorhynchus was seized during or immediately after a successful hunt. According to the fossil record, Rhamphorhynchus frequently were accidentally seized by large Aspidorhnychus. In some cases the fibrous tissue of the wing membrane got entangled with the rostral teeth such that the fish was unable to get rid of the pterosaur. Such encounters ended fatally for both. Intestinal contents of Aspidorhynchus-type fishes are known and mostly comprise fishes and in one single case a Homoeosaurus. Obviously Rhamphorhynchus did not belong to the prey spectrum of Aspidorhynchus. [ABSTRACT FROM AUTHOR]

Variation in daily activity patterns facilitates temporal partitioning of habitat and resources among species. Knowledge of temporal niche partitioning in paleobiological systems has been limited by the difficulty of obtaining reliable information about activity patterns from fossils. On the basis of an analysis of scleral ring and orbit morphology in 33 archosaurs, including dinosaurs and pterosaurs, we show that the eyes of Mesozoic archosaurs were adapted to all major types of diel activity (that is, nocturnal, diurnal, and cathemeral) and provide concrete evidence of temporal niche partitioning in the Mesozoic. Similar to extant amniotes, flyers were predominantly diurnal; terrestrial predators, at least partially, nocturnal; and large herbivores, cathemeral. These similarities suggest that ecology drives the evolution of diel activity patterns. [ABSTRACT FROM AUTHOR]

The article discusses a study on the possible presence of postcranial skeletal pneumatization (PSP) and air-sacs in the ancestor of pterosaurs in the Late Triassic and Jurassic era and ornithodirans. The study revealed the occurrence of evidence of the pattern of PSP in Dimorphodon macronyx which was first stated in the research by S. Sangster. Also explored are other research that showed the associations between air-sacs and axial pneumatization elements in extant birds.