Pterosaur Species Research Articles

Comments on the Pteranodontidae (Pterosauria, Pterodactyloidea) with the description of two new species

Considered one of the best known flying reptiles, Pteranodon has been subject to several reviews in the last century. Found exclusively in the Upper Cretaceous Niobrara Formation and Pierre Shale Group 11 species have been attributed to this genus (excluding the ones presently regarded as representing Nyctosaurus). While reviewers agree that this number is inflated, there is disagreement on how many species can be identified. The last review recognized only two species (Pteranodon longiceps and Pteranodon sternbergi) both being sexually dimorphic. Based on several cranial features, some specimens previously referred to the genus Pteranodon are re-evaluated leading to the recognition of the following species, two of which new that are described here: Pteranodon longiceps, Geosternbergia sternbergi, Geosternbergia maiseyi sp. nov., and Dawndraco kanzai gen. et sp. nov. They differ mainly by features such as the direction and extension of the frontal crest, the angle and extension of the posterior process of the premaxillae, the shape and extension of the lower temporal fenestra and the length and proportion of the rostrum. The procedures to recognize a pterosaur species are also discussed here, and must take into account primarily morphology, in conjunction with stratigraphic and geographic data. Although well aware that changes in morphology not always reflect taxonomy, the lack of stratigraphic data and the limited number of specimens that can be confidently assigned to one species hampers our understanding on the morphological variations as a function of ontogeny, individual variation and sexual dimorphism. Although the present study has not eliminated the possibility to recognize such differences, caution is needed before models are generalized for pterosaurs.

Pterosaur body mass estimates from three-dimensional mathematical slicing

ABSTRACT Body masses for 14 species of pterosaur spanning four orders of magnitude were estimated using three-dimensional, digital models. The modeled taxa comprised seven paraphyletic ‘rhamphorhynchoids’: Anurognathus ammoni, Dimorphodon macronyx, Eudimorphodon ranzii, Jeholopterus ningchengensis, Preondactylus buffarinii, Rhamphorhynchus muensteri, and Sordes pilosus; and seven pterodactyloids: Anhanguera santanae, Dsungaripterus weii, Pteranodon longiceps, Pterodaustro guinazui, Pterodactylus sp., Quetzalcoatlus northropi, Tupuxuara longicristatus. The reliability of the mass estimation methods were tested with equivalent models of six extant species of bird with masses that spanned three orders of magnitude. The close agreement between model bird mass estimates and those of the living forms provides a level of confidence in the results obtained for the extinct pterosaurs. The masses of the axial body regions (tail, trunk, neck, head), limbs, and patagia of the pterosaurs were individually estimated and distinct differences in relative body proportions were found between species. Allometric relationships between body length and wingspan and body mass were derived for ‘rhamphorhynchoids’ and pterodactyloids to facilitate the estimation of body masses for other pterosaurs known from incomplete material, and these relationships also highlight differences in phyletic shape change between the two groups. The estimated mass for the largest pterosaur known, Quetzalcoatlus northropi, exceeds the previous highest estimates by more than 100%, and it is argued that this extremely large pterosaur is better interpreted as a secondarily flightless form.

A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs

ABSTRACT A new rhamphorhynchid pterosaur species, Sericipterus wucaiwanensis, gen. et sp. nov., is described from the Upper Jurassic part of the Shishugou Formation in the Xinjiang Autonomous Region of northwest China. Pterosaurs from this unit are the earliest and only records of pterosaurs in the Jurassic of northwest China. The individual specimen is one of the largest known among ‘rhamphorhynchoids,’ or non-pterodactyloid pterosaurs. The holotype comprises an associated skeleton of mostly disarticulated, largely three-dimensional material. Although partly crushed, the preservation in this specimen reveals morphology rarely seen in non-pterodactyloid pterosaurs. This includes a distinct cervical intervertebral articulation morphology that is proposed to be widespread among the non-pterodactyloids. The skull of this new specimen is most similar to that of other rhamphorhynchids, Angustinaripterus longicephalus and Harpactognathus gentryii, found in terrestrial deposits. A phylogenetic analysis of 18 non-pterodactyloid pterosaurs and the Pterodactyloidea places Sericipterus wucaiwanensis with these species within the Rhamphorhynchinae and a monophyletic Rhamphorhynchidae. Unlike previous phylogenetic analyses, the Dimorphodontidae is paraphyletic, the Campylognathoididae is polyphyletic, and the Anurognathidae is the sister group of the Pterodactyloidea. Sericipterus wucaiwanensis, Angustinaripterus longicephalus, Harpactognathus gentryii represent a clade of large pterosaurs that likely lived in the terrestrial settings in which they preserved.

The first pterosaur trackways from Japan

In 1990, five well-preserved pterosaur trackways were discovered in the Kitadani Formation (Lower Cretaceous) of the Kitadani Dinosaur Quarry, Fukui Prefecture, Japan. They occur on the surface of an isolated dark grey siltstone slab (70 × 50 cm) along with amphibian and bird tracks as well as feeding marks. All pterosaur trackways (a total of 64 imprints) show a clear quadrupedal gait pattern comprising manus and pes prints which reflect their detailed anatomy of the feet. Manus and pes imprint is very small, average 22.6 mm and 21.9 mm long, respectively. The manus has an anteriorly oriented digit I imprint with a medially oriented hook-like sharp claw mark, which makes a high divarication (average 155.4°) between the posteriorly oriented digit III imprint. The high digital abduction clearly shows that digit I of the manus could be more hyper-extended anteriorly than previously thought: spreading the digits as much as possible would provide a more stable contact with substrate as well as better support the center of pterosaur body mass displaced anteriorly. The interdigital webbing of the pes imprint extends from the metatarso-phalangeal joint near to the bases of four claws. As these features clearly distinguish the Kitadani pterosaur trackways from five known ichnospecies of Pteraichnus, we assign them to a new ichnospecies, Pteraichnus nipponensis. Abundant small pterosaur ichnotaxa from Spain, Korea, and Japan indicate that many small pterodactyloid pterosaur species existed in the Early Cretaceous although there is no single skeletal datum yet.

A NEW OXFORDIAN PLIOSAURID (PLESIOSAURIA, PLIOSAURIDAE) IN THE CARIBBEAN SEAWAY

Abstract: A new pliosaurid, Gallardosaurus iturraldei gen nov. et sp. nov., was found in the Viñales area, western Cuba, in sediments of the Jagua Formation, middle–late Oxfordian. This new taxon is characterized by: wide participation of the premaxilla in the outer margin of the external naris; frontal not participating in the orbital margin; postorbital in contact with the jugal and squamosal; presence of anterior pterygoid vacuity; cultriform process of parasphenoid convex and exposed in palatal view; pterygoid flanges high; jaw articulation low relative to tooth row; trihedral teeth in cross‐section and with smooth ridges at least in the labial face. A phylogenetic analysis suggests that Gallardosaurus forms a clade with Peloneustes, the most common pliosaurid genus occurring in the Oxford Clay. The Caribbean Seaway was, at least since the Oxfordian, a corridor that permitted the interchange for marine biota between Western Tethys and Eastern Pacific realms. Among vertebrates, bony fish and long‐necked plesiosaurs prevailed. However, marine pleurodiran turtles, metriorhynchid crocodilians, ophthalmosaurian ichthyosaurs, and pliosaurids (G. iturraldei gen. nov. et sp. nov.) have also been found, as well as at least two species of pterosaurs, and one camarasaurian dinosaur. Among these reptiles there were off‐shore pelagic forms such as the ichthyosaurs and metriorhynchids, together with the pliosaurid G. iturraldei gen. nov. et sp. nov.; other taxa were presumably less pelagic, such as the pleurodiran turtles and the cryptoclidid plesiosauroids. Gallardosaurus iturraldei gen. nov. et sp. nov. would have played the role of an active predator taking advantage of nectonic fish recorded in the area.

Palaeogenomics of pterosaurs and the evolution of small genome size in flying vertebrates

The two living groups of flying vertebrates, birds and bats, both have constricted genome sizes compared with their close relatives. But nothing is known about the genomic characteristics of pterosaurs, which took to the air over 70 Myr before birds and were the first group of vertebrates to evolve powered flight. Here, we estimate genome size for four species of pterosaurs and seven species of basal archosauromorphs using a Bayesian comparative approach. Our results suggest that small genomes commonly associated with flight in bats and birds also evolved in pterosaurs, and that the rate of genome-size evolution is proportional to genome size within amniotes, with the fastest rates occurring in lineages with the largest genomes. We examine the role that drift may have played in the evolution of genome size within tetrapods by testing for correlated evolution between genome size and body size, but find no support for this hypothesis. By contrast, we find evidence suggesting that a combination of adaptation and phylogenetic inertia best explains the correlated evolution of flight and genome-size contraction. These results suggest that small genome/cell size evolved prior to or concurrently with flight in pterosaurs. We predict that, similar to the pattern seen in theropod dinosaurs, genome-size contraction preceded flight in pterosaurs and bats.

Three-dimensional geometry of a pterosaur wing skeleton, and its implications for aerial and terrestrial locomotion

This study reports on the three-dimensional spatial arrangement and movements of the skeleton of Anhanguera santanae (Pterodactyloidea: Ornithocheiridae), determined using exceptionally well-preserved uncrushed fossil material, and a rigid-body method for analysing the joints of extinct animals. The geometric results of this analysis suggest that the ornithocheirids were inherently unstable in pitch and yaw. As a result, pitch control would probably have been brought about by direct adjustment of the angle of attack of the wing, by raising or lowering the trailing edge from the root using the legs if, as is indicated in soft-tissue specimens of a number of unrelated pterosaur species, the legs were attached to the main wing membrane, or by using long-axis rotations at the shoulder or wrist to raise and lower the trailing edge from the wingtip. An analysis of the three-dimensional morphology of the wrist lends support to the idea that the pteroid – a long, slender wrist bone unique to pterosaurs that supported a membranous forewing – was directed forwards in flight, not towards the body. As a result, the forewing could have fulfilled the function of an air-brake and high-lift device, and may also have had an important role in pitch, yaw, and roll control. The joint analysis is consistent with a semi-erect quadrupedal model of terrestrial locomotion in the ornithocheirids. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 27–69.

A new species of Istiodactylus (Pterosauria, Pterodactyloidea) from the Lower Cretaceous of Liaoning, China

ABSTRACT Istiodactylus sinensis, sp. nov., from the Jiufotang Formation of Liaoning, People's Republic of China, is described on the basis of a single nearly complete and nearly osteologically adult specimen. This is the tenth pterosaur described from this formation and the eighteenth pterosaur species described from northeastern China in almost half as many years. The species is placed in the Istiodactylidae, which was previously a monospecific family of pterodactyloid pterosaurs known only from the Isle of Wight, England. The new form is distinct from the two other istiodactylid species. It is smaller, more plesiomorphic, and younger than Istiodactylus latidens, but larger and more derived than the contemporaneous Nurhachius ignaciobritoi. Istiodactylus sinensis is very similar to I. latidens, so that almost all of the previous autapomorphies of I. latidens are now synapomorphies of Istiodactylus. They differ most in that I. sinensis is much smaller than I. latidens. The length of the wingspan, skull, and most of the preserved limb elements of I. sinensis are about 63 percent of the wingspan and elements of I. latidens. This new specimen demonstrates that Istiodactylus is diagnosed by, among other characters, a dorsoventrally depressed but not laterally expanded rostrum, and the presence of a suborbital vacuity. A dorsal deflection of the alveolar margins of the jaws and a humerus between 55 percent and one and a half times the length of metacarpal IV are synapomorphies uniting the Istiodactylidae and the Anhangueridae.

A new pterodactyloid pterosaur from the Wessex Formation (Lower Cretaceous) of the Isle of Wight, England

A new pterosaur specimen comprising a partial skull and associated postcranial elements from the Lower Cretaceous Wessex Formation of Yaverland, Isle of Wight, southern England, is assigned to a new genus and species of ornithocheirid pterosaur, Caulkicephalus trimicrodon gen. et sp. nov., based on several unique features including a heterodont dentition in which the fifth, sixth and seventh teeth are reduced in size compared with those at positions 1–4 and 8–9; the presence of a frontoparietal crest and maxillopremaxillary crest that do not unite over the antorbital fenestra or cranium; a palatal ridge that extends no further forward than the eighth to ninth tooth pairs. The new taxon is the second species of pterosaur from the Wealden Group of the Wessex Basin.

Neuroanatomy of flying reptiles and implications for flight, posture and behaviour.

Comparison of birds and pterosaurs, the two archosaurian flyers, sheds light on adaptation to an aerial lifestyle. The neurological basis of control holds particular interest in that flight demands on sensory integration, equilibrium, and muscular coordination are acute. Here we compare the brain and vestibular apparatus in two pterosaurs based on high-resolution computed tomographic (CT) scans from which we constructed digital endocasts. Although general neural organization resembles birds, pterosaurs had smaller brains relative to body mass than do birds. This difference probably has more to do with phylogeny than flight, in that birds evolved from nonavian theropods that had already established trends for greater encephalization. Orientation of the osseous labyrinth relative to the long axis of the skull was different in these two pterosaur species, suggesting very different head postures and reflecting differing behaviours. Their enlarged semicircular canals reflect a highly refined organ of equilibrium, which is concordant with pterosaurs being visually based, aerial predators. Their enormous cerebellar floccular lobes may suggest neural integration of extensive sensory information from the wing, further enhancing eye- and neck-based reflex mechanisms for stabilizing gaze.

Flight characteristics of Triassic and Jurassic Pterosauria: an appraisal based on wing shape

The mass, wingspan, and wing area of pterosaurs were reconstructed. Mass was estimated by determining volume and multiplying by avian density. This method was considered appropriate only for smaller pterosaur species because there is evidence for lower density in larger species. These reconstructions were used to compare the wing shapes of Triassic and Jurassic pterosaurs with those of birds. Pterosaurs had wings of below-average loading and above-average aspect compared to the avian mean. This wing design was compatible with relatively slow and highly efficient flight, with high maneuverability. Wing area depends on the reconstruction model adopted; wings attached to the hindlimb principally reduced aspect, and secondarily reduced loading, which would improve take-off performance at the expense of efficiency. The wing shape and cranial feeding adaptations of pterosaurs were most compatible with a marine or aerial predatory adaptive zone. The reconstructed pterosaurs show a limited range of wing shape compared to birds. This may partly reflect preservational bias favoring species living in marine or lagoonal environments, but this is not a complete explanation because there is a lack of pterosaurs with wings of high loading like the marine ducks and auks. Structural, physiological, or adaptive factors may have limited pterosaur wing shape.