Air Space Proportion in Pterosaur Limb Bones Using Computed Tomography and Its Implications for Previous Estimates of Pneumaticity

Air Space Proportion (ASP) is a measure of how much air is present within a bone, which allows for a quantifiable comparison of pneumaticity between specimens and species. Measured from zero to one, higher ASP means more air and less bone. Conventionally, it is estimated from measurements of the internal and external bone diameter, or by analyzing cross-sections. To date, the only pterosaur ASP study has been carried out by visual inspection of sectioned bones within matrix. Here, computed tomography (CT) scans are used to calculate ASP in a small sample of pterosaur wing bones (mainly phalanges) and to assess how the values change throughout the bone. These results show higher ASPs than previous pterosaur pneumaticity studies, and more significantly, higher ASP values in the heads of wing bones than the shaft. This suggests that pneumaticity has been underestimated previously in pterosaurs, birds, and other archosaurs when shaft cross-sections are used to estimate ASP. Furthermore, ASP in pterosaurs is higher than those found in birds and most sauropod dinosaurs, giving them among the highest ASP values of animals studied so far, supporting the view that pterosaurs were some of the most pneumatized animals to have lived. The high degree of pneumaticity found in pterosaurs is proposed to be a response to the wing bone bending stiffness requirements of flight rather than a means to reduce mass, as is often suggested. Mass reduction may be a secondary result of pneumaticity that subsequently aids flight.     

On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness

The size and flight mechanics of giant pterosaurs have received considerable research interest for the last century but are confused by conflicting interpretations of pterosaur biology and flight capabilities. Avian biomechanical parameters have often been applied to pterosaurs in such research but, due to considerable differences in avian and pterosaur anatomy, have lead to systematic errors interpreting pterosaur flight mechanics. Such assumptions have lead to assertions that giant pterosaurs were extremely lightweight to facilitate flight or, if more realistic masses are assumed, were flightless. Reappraisal of the proportions, scaling and morphology of giant pterosaur fossils suggests that bird and pterosaur wing structure, gross anatomy and launch kinematics are too different to be considered mechanically interchangeable. Conclusions assuming such interchangeability--including those indicating that giant pterosaurs were flightless--are found to be based on inaccurate and poorly supported assumptions of structural scaling and launch kinematics. Pterosaur bone strength and flap-gliding performance demonstrate that giant pterosaur anatomy was capable of generating sufficient lift and thrust for powered flight as well as resisting flight loading stresses. The retention of flight characteristics across giant pterosaur skeletons and their considerable robustness compared to similarly-massed terrestrial animals suggest that giant pterosaurs were not flightless. Moreover, the term 'giant pterosaur' includes at least two radically different forms with very distinct palaeoecological signatures and, accordingly, all but the most basic sweeping conclusions about giant pterosaur flight should be treated with caution. Reappraisal of giant pterosaur material also reveals that the size of the largest pterosaurs, previously suggested to have wingspans up to 13 m and masses up to 544 kg, have been overestimated. Scaling of fragmentary giant pterosaur remains have been misled by distorted fossils or used inappropriate scaling techniques, indicating that 10-11 m wingspans and masses of 200-250 kg are the most reliable upper estimates of known pterosaur size.     

A morphospace‐based test for competitive exclusion among flying vertebrates: did birds,bats and pterosaurs get in each other's space?

Three vertebrate groups – birds, bats and pterosaurs – have evolved flapping flight over the past 200 million years. This innovation allowed each clade access to new ecological opportunities, but did the diversification of one of these groups inhibit the evolutionary radiation of any of the others? A related question is whether having the wing attached to the hindlimbs in bats and pterosaurs constrained their morphological diversity relative to birds. Fore‐ and hindlimb measurements from 894 specimens were used to construct a morphospace to assess morphological overlap and range, a possible indicator of competition, among the three clades. Neither birds nor bats entered pterosaur morphospace across the Cretaceous–Paleogene (Tertiary) extinction. Bats plot in a separate area from birds, and have a significantly smaller morphological range than either birds or pterosaurs. On the basis of these results, competitive exclusion among the three groups is not supported.     

A New Model for the Evolution of the Pterosaur Wing--with a twist

A new hypothesis for the evolution of the pterosaur wing is presented. It is based on the observation that many aspects of pterosaur non-wing anatomy were present in their non-volant, prolacertiform sister taxa. In pterosaurs alone, metacarpal IV was twisted 90° medially, so the wing digit flexed in the plane of the metacarpus, rather than towards the palm as in other tetrapods. While grappling a tree using manual digits I-III, a pre-pterosaur with this character would have been free to flex and extend digit IV in the plane tangential to the circumference of the tree. The addition of a small dermal extension that opened like a Japanese fan would essentially have completed the development of the proto-wing. Its subsequent increase in size was brought on by selective competition. Successful powered flight would have been possible only after a critical wing size had been achieved.     

Limb disparity and wing shape in pterosaurs

The limb proportions of the extinct flying pterosaurs were clearly distinct from their living counterparts, birds and bats. Within pterosaurs, however, we show that further differences in limb proportions exist between the two main groups: the clade of short-tailed Pterodactyloidea and the paraphyletic clades of long-tailed rhamphorhynchoids. The hindlimb to forelimb ratios of rhamphorhynchoid pterosaurs are similar to that seen in bats, whereas those of pterodactyloids are much higher. Such a clear difference in limb ratios indicates that the extent of the wing membrane in rhamphorhynchoids and pterodactyloids may also have differed; this is borne out by simple ternary analyses. Further, analyses also indicate that the limbs of Sordes pilosus, a well-preserved small taxon used as key evidence for inferring the extent and shape of the wing membrane in all pterosaurs, are not typical even of its closest relatives, other rhamphorhynchoids. Thus, a bat-like extensive hindlimb flight membrane, integrated with the feet and tail may be applicable only to a small subset of pterosaur diversity. The range of flight morphologies seen in these extinct reptiles may prove much broader than previously thought.     

Constraints on the wing morphology of pterosaurs

Animals that fly must be able to do so over a huge range of aerodynamic conditions, determined by weather, wind speed and the nature of their environment. No single parameter can be used to determine-let alone measure-optimum flight performance as it relates to wing shape. Reconstructing the wings of the extinct pterosaurs has therefore proved especially problematic: these Mesozoic flying reptiles had a soft-tissue membranous flight surface that is rarely preserved in the fossil record. Here, we review basic mechanical and aerodynamic constraints that influenced the wing shape of pterosaurs, and, building on this, present a series of theoretical modelling results. These results allow us to predict the most likely wing shapes that could have been employed by these ancient reptiles, and further show that a combination of anterior sweep and a reflexed proximal wing section provides an aerodynamically balanced and efficient theoretical pterosaur wing shape, with clear benefits for their flight stability.     

How the pterosaur got its wings

Throughout the evolutionary history of life, only three vertebrate lineages took to the air by acquiring a body plan suitable for powered flight: birds, bats, and pterosaurs. Because pterosaurs were the earliest vertebrate lineage capable of powered flight and included the largest volant animal in the history of the earth, understanding how they evolved their flight apparatus, the wing, is an important issue in evolutionary biology. Herein, I speculate on the potential basis of pterosaur wing evolution using recent advances in the developmental biology of flying and non‐flying vertebrates. The most significant morphological features of pterosaur wings are: (i) a disproportionately elongated fourth finger, and (ii) a wing membrane called the brachiopatagium, which stretches from the posterior surface of the arm and elongated fourth finger to the anterior surface of the leg. At limb‐forming stages of pterosaur embryos, the zone of polarizing activity (ZPA) cells, from which the fourth finger eventually differentiates, could up‐regulate, restrict, and prolong expression of 5′‐located Homeobox D (Hoxd) genes (e.g. Hoxd11, Hoxd12, and Hoxd13) around the ZPA through pterosaur‐specific exploitation of sonic hedgehog (SHH) signalling. 5′Hoxd genes could then influence downstream bone morphogenetic protein (BMP) signalling to facilitate chondrocyte proliferation in long bones. Potential expression of Fgf10 and Tbx3 in the primordium of the brachiopatagium formed posterior to the forelimb bud might also facilitate elongation of the phalanges of the fourth finger. To establish the flight‐adapted musculoskeletal morphology shared by all volant vertebrates, pterosaurs probably underwent regulatory changes in the expression of genes controlling forelimb and pectoral girdle musculoskeletal development (e.g. Tbx5), as well as certain changes in the mode of cell–cell interactions between muscular and connective tissues in the early phase of their evolution. Developmental data now accumulating for extant vertebrate taxa could be helpful in understanding the cellular and molecular mechanisms of body‐plan evolution in extinct vertebrates as well as extant vertebrates with unique morphology whose embryonic materials are hard to obtain.     

Respiratory Evolution Facilitated the Origin of Pterosaur Flight and Aerial Gigantism

Pterosaurs, enigmatic extinct Mesozoic reptiles, were the first vertebrates to achieve true flapping flight. Various lines of evidence provide strong support for highly efficient wing design, control, and flight capabilities. However, little is known of the pulmonary system that powered flight in pterosaurs. We investigated the structure and function of the pterosaurian breathing apparatus through a broad scale comparative study of respiratory structure and function in living and extinct archosaurs, using computer-assisted tomographic (CT) scanning of pterosaur and bird skeletal remains, cineradiographic (X-ray film) studies of the skeletal breathing pump in extant birds and alligators, and study of skeletal structure in historic fossil specimens. In this report we present various lines of skeletal evidence that indicate that pterosaurs had a highly effective flow-through respiratory system, capable of sustaining powered flight, predating the appearance of an analogous breathing system in birds by approximately seventy million years. Convergent evolution of gigantism in several Cretaceous pterosaur lineages was made possible through body density reduction by expansion of the pulmonary air sac system throughout the trunk and the distal limb girdle skeleton, highlighting the importance of respiratory adaptations in pterosaur evolution, and the dramatic effect of the release of physical constraints on morphological diversification and evolutionary radiation.     

A survey of pathologies of large pterodactyloid pterosaurs

Examples of four types of pathologies have been found on specimens of large pterodactyloid pterosaurs. Osteoarthritis of the intersyncarpal, carpometacarpal, or metacarpophalangeal joints, indicated by grooving of the articular surfaces in the direction of joint motion, is present on at least four pterosaur specimens from the Cambridge Greensand. This pathology has not been found in Pteranodon despite comparably large samples of specimens. Pteranodon most commonly exhibits deformed bones and lumps apparently resulting from injury, necrosis, and subsequent repair. A small number of pathological bone growths apparently not associated with injuries were found, as were healed fractures of a fourth wing phalanx and a pedal phalanx. The absence of healed fractures of other elements suggests that they were either uncommon or catastrophic and fatal. This study suggests that pathologies were not uncommon in large pterosaurs.     

ON THE RECONSTRUCTION OF PTEROSAURS AND THEIR MANNER OF FLIGHT, WITH NOTES ON VORTEX WAKES

Classical pterosaur reconstructions are variants on a ‘bat-analogy’, whereby the wing is conceived as a simple membrane with no inherent bending strength, stretched between the arm and leg skeletons. The legs are considered to be splayed out to the sides, as in bats, so that the animal would have to adopt a quadrupedal stance on the ground, supported on its feet and the metacarpo-phalangeal joints. In recent years an alternative ‘bird-analogy’ has come to be generally accepted. This hypothesis, most elements of which are due to Padian (1983 a, b) calls for the animal to stand upright on its legs like a bird. The wings are independent of the legs, as in birds, are stiffened by skeletal fibres in the membrane, and have a very narrow, sharply pointed shape. There are difficulties in reconciling the bird-analogy with the evidence. The long-tailed rhamphorhynchs might conceivably have balanced their weight about their hip joints but this would not have been possible for the short-tailed pterodactyls. The bird pelvis shows modifications which permit bipedal standing in spite of the reduction of the tail, but no equivalent adaptations are seen in pterodactyls. Besides, all known pterosaur pelvises, except that of the giant pterodactyl Pteranodon were open ventrally, which would have precluded the legs from being brought to a parasagittal position, as required for bipedal walking. The notion that the wing was not attached to the legs is based on negative evidence, in that no clear impressions of the inner end of the wing membrane are preserved in the fossils. However one pterodactyl fossil shows a membrane edge approaching the ankle joint. In fossils that are preserved with the wings forward, the legs have been pulled forwards by the ankles. A tendon connecting the ankle to the wing tip is consistent with the evidence. The ‘fibres’ in the wing membranes are actually impressions of surface ridges, with no internal structure, and are better interpreted as surface wrinkles in the skin, caused by contraction of elastic fibres within the membrane. The bird analogy also results in a very unsatisfactory wing from an aerodynamic point of view. The structure of an animal wing is best understood in terms of the type of vortex wake it is adapted to generate. Hummingbirds, and insects capable of economical hovering, have wings that can be inverted on the upstroke, and when hovering, generate a wake consisting of two vortex rings per wingbeat cycle. The span of such wings is fixed, which implies that they create a ‘ladder wake’ in cruising flight, consisting of a pair of undulating wing-tip vortices, joined by a transverse vortex at each transition from downstroke to upstroke and back. Normal birds cannot invert their wings, and so are less efficient in hovering, but they can shorten the wing during the upstroke in cruising flight. This creates a ‘concertina wake’, with no transverse vortices. Hummingbirds show very limited migration performance, compared with normal birds, with the implication that a wing capable of creating a concertina wake is more economical in cruising flight than one creating a ladder wake, and is an essential adaptation for long-distance migration. A revised reconstruction of the pterosaur wing starts from the observations that, contrary to the currently popular bird-analogy, pterosaurs were not bipedal, their wings did not contain stiffening fibres but did contain elastic fibres, and the trailing edge of the membrane was supported by a tendon joining the tip of the wing finger to the ankle. A hypothetical arrangement of elastic fibres, that accounts well for the observed pattern of wrinkles in contracted wings, also allows the planform shape of the wing to be adjusted in much the same way as seen in birds, although using a completely different mechanism. It opens the possibility that pterosaurs could fly with a concertina wake, and thus could have been long-distance migrators like modern birds. Although this hypothetical wing is mechanically somewhat bat-like, it is not a return to the classical bat-analogy. It would not have the high degree of control over profile shape, which gives bats their outstanding manoeuvrability. On the other hand bats do not have the degree of control over their wingspan that is suggested here for pterosaurs, and consequently are not notable for migration performance.     

Wing size but not wing shape is related to migratory behavior in a soaring bird

Both wing size and wing shape affect the flight abilities of birds. Intra and inter‐specific studies have revealed a pattern where high aspect ratio and low wing loading favour migratory behaviour. This, however, have not been studied in soaring migrants. We assessed the relationship between the wing size and shape and the characteristics of the migratory habits of the turkey vulture Cathartes aura, an obligate soaring migrant. We compared wing size and shape with migration strategy among three fully migratory, one partially migratory and one non‐migratory (resident) population distributed across the American continent. We calculated the aspect ratio and wing loading using wing tracings to characterize the wing morphology. We used satellite‐tracking data from the migratory populations to calculate distance, duration, speed and altitude during migration. Wing loading, but not aspect ratio, differed among the populations, segregating the resident population from the completely migratory ones. Unlike what has been reported in species using flapping flight during migration, the migratory flight parameters of turkey vultures were not related to the aspect ratio. By contrast, wing loading was related to most flight parameters. Birds with lower wing loading flew farther, faster, and higher during their longer journeys. Our results suggest that wing morphology in this soaring species enables lower‐cost flight, through low wing‐loading, and that differences in the relative sizes of wings may increase extra savings during migration. The possibility that wing shape is influenced by foraging as well as migratory flight is discussed. We conclude that flight efficiency may be improved through different morphological adaptations in birds with different flight mechanisms.     

Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method

Similarities between the anatomies of living organisms are often used to draw conclusions regarding the ecology and behaviour of extinct animals. Several pterosaur taxa are postulated to have been skim-feeders based largely on supposed convergences of their jaw anatomy with that of the modern skimming bird, Rynchops spp. Using physical and mathematical models of Rynchops bills and pterosaur jaws, we show that skimming is considerably more energetically costly than previously thought for Rynchops and that pterosaurs weighing more than one kilogram would not have been able to skim at all. Furthermore, anatomical comparisons between the highly specialised skull of Rynchops and those of postulated skimming pterosaurs suggest that even smaller forms were poorly adapted for skim-feeding. Our results refute the hypothesis that some pterosaurs commonly used skimming as a foraging method and illustrate the pitfalls involved in extrapolating from limited morphological convergence.     

A New Pterosaur from the Jurassic of Cuba

Cacibupteryx caribensis gen. et sp. nov. is a new pterosaur of the family Rhamphorhynchidae found in western Cuba, in rocks of the Jagua Formation (Middle–Upper Oxfordian). The holotype, a skull and part of the left wing, is one of the few Jurassic pterosaurs that is well preserved in three dimensions. The new taxon shares characters with early and late Jurassic pterosaurs, and is one of the few late Jurassic taxa from western Laurasia and Gondwana. Furthermore, Cacibupteryx joins Nesodactylus hesperius Colbert from Cuba, and Sordes pilosus Sharov, from Kazakhstan as the most complete pterosaur recorded from the Middle–Upper Oxfordian. Cacibupteryx caribensis is one of the largest Jurassic pterosaurs known, and its skull possesses several distinct characters, including relatively broad roof elements (mainly frontal and parietal bones), a jugal with a prominent recess, occipital table trapezoidal in shape with the maximum width between the quadrate bones, and a small fenestra located in the posterior part of the pterygoid bones. In the Oxfordian, the Caribbean Corridor separated Laurasia and western Gondwana. The diversity of the marine herpetofauna found in the Jagua Vieja Member (Jagua Formation), and of teleostean fish, confirms that the corridor was an effective seaway over which flew at least Nesodactylus and Cacibupteryx .     

New models for the wing extension in pterosaurs

All powered flying animals have to face the same energetic problems: operating the wings during steady flight with muscles that require constant energy input and neural control to work. Accordingly the extant flying vertebrates have apparently found very similar solutions to parts of these issues – the biomechanical automatism built in their skeletal, muscular and connective tissue system. Based on these extant analogues (birds and bats) two new models are presented here for the mechanism of the distal wing extension in pterosaurs, an extinct group of flying vertebrates. The elongate fourth finger which solely supported their extensive flight membrane was a long lever arm that experienced significant loads and for which a reduction in muscle mass through automatisation would have been strongly beneficial. In the first model we hypothesize the presence of a propatagial ligament or ligamentous system which, as a result of the elbow extension, automatically performs and maintains the extension of the wing finger during flight and prohibits the hyperextension of the elbow. The second model has a co-operating bird-like propatagial ligamentous system and bat-like tendinous extensor muscle system on the forearm of the hypothetical pterosaur. Both models provide strong benefits to an animal with powered flight: (1) reduction of muscles and weight in the distal wing; (2) prevention of hyper extension of the elbow against drag; (3) automating wing extension and thereby reducing metabolic costs required to operate the pterosaurian locomotor apparatus. These models, although hypothetical, fit with the existing fossil evidence and lay down a basis for further biomechanical and/or aerodynamical investigations.     

Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guiñazui

Life-history parameters of pterosaurs such as growth and ontogenetic development represent an enigma. This aspect of pterosaur biology has remained perplexing because few pterosaur taxa are represented by complete ontogenetic series. Of these, Pterodaustro is unique in that besides being represented by hundreds of individuals with wing spans ranging from 0.3 to 2.5m, it includes an embryo within an egg. Here we present a comprehensive osteohistological assessment of multiple skeletal elements of a range of ontogenetic sizes of Pterodaustro, and we provide unparalleled insight into its growth dynamics. We show that, upon hatching, Pterodaustro juveniles grew rapidly for approximately 2 years until they reached approximately 53% of their mature body size, whereupon they attained sexual maturity. Thereafter, growth continued for at least another 3-4 years at comparatively slower rates until larger adult body sizes were attained. Our analysis further provides definitive evidence that Pterodaustro had a determinate growth strategy.     

A Catalog of Pterosaur Pedes for Trackmaker Identification

The matching of ichnites to extinct trackmakers has been done successfully with a variety of taxa, from basal hominids to basal tetrapods. Traces attributed to pterosaurs have been studied for more than 50 years, but little interest has been shown in the pedes themselves. While ichnites can vary greatly in their correspondence to their trackmaker, most pterosaur tracks appear to preserve sufficient detail to assess their origins. This report presents a catalog of pterosaur pedal skeletons that can be matched to a wider spectrum of ichnites, including digitigrade and bipedal ichnites previously not associated with pterosaurs. A variety of pedal characters separate several putative genera into distinct clades, some only distantly related to one another. Distinct pedal characters indicate certain tiny pterosaurs were not juveniles of dissimilar adults, but were separate taxa and likely adults themselves. A squamate and fenestrasaur origin for pterosaurs is supported. These new insights overturn long-standing paradigms. The pterosaur pes contains a wealth of data that should not be ignored. Application of this data enables a more precise identification of both skeletal taxa and ichnotaxa.     

辽西早白垩世九佛堂组两种新的翼手龙类化石(英文)

简要报道了辽西热河群上部九佛堂组两件新的翼手龙类化石 ,即夜翼龙科(Nyctosauridae)的张氏朝阳翼龙 (新属、新种 )Chaoyangopteruszhangigen .etsp .nov.和古魔翼龙科 (Anhangueridae)的顾氏辽宁翼龙 (新属、新种 )Liaoningopterusguigen .et.sp .nov.。前者为保存较完整的化石骨架 ,后者为一大型翼龙的头骨和部分头后骨骼化石。朝阳翼龙是夜翼龙科在亚洲大陆的首次确切的化石记录 ,也是层位最低和保存最完整的化石骨架。朝阳翼龙具有4节翼指骨 ,手指爪粗大弯曲 ,这些发现补充和修正了前人认为的夜翼龙科只有 3节翼指骨 ,手指爪退化缺失等一些重要的形态学特征。朝阳翼龙与该科的Nyctosaurusgracilis头后骨骼相比 ,具有许多不同的特征 ,如胫骨特长 ,远长于股骨 ,翼掌骨和第 1翼指骨相对较短 ,肩胛骨短于乌喙骨等。辽宁翼龙是我国已发现的个体最大的翼龙化石 ,发育前上颌骨和齿骨弧形脊突这一古魔翼龙科的重要鉴别特征。与该科的其他成员相比 ,辽宁翼龙上、下颌的牙齿较少 ,仅分布在其前部 ,齿列约占上、下颌长度的 1 / 2。上颌第 1、3齿小 ,第 2、4齿巨大 ,其中第 4齿最大 ,为已知翼龙中最大的牙齿。牙齿具有明显的替换现象。夜翼龙科的成员仅分布于美洲大陆的晚白垩世地层中 ,而古魔翼龙科的成员则是     

Palaeohistology of the bones of pterosaurs (Reptilia: Archosauria): anatomy, ontogeny, and biomechanical implications

Thin sections from long bones of specimens representing pterosaurs ranging from the Early Jurassic to the latest Cretaceous provide a profile of bone histology across a range of sizes, skeletal elements, growth stages, and phylogenetic positions. Most pterosaur bone is fibro-lamellar, organized in an unusual way that suggests high growth rates through ontogeny. Fibro-lamellar deposits are finished by a relatively abrupt deceleration or cessation of growth represented by lamellar, poorly vascularized subperiosteal bone in what appear to be adults. Pterosaurs had the thinnest bone walls of any tetrapods; they complemented high rates of periosteal deposition with almost equally high rates of endosteal erosion. Pterosaurs show a great variety of histologic features that include articular calcified cartilage, sub-chondral bone plates, trabecular bone struts and related internal supports, and secondary deposition and remodeling of bone. They remodeled their bones internally by (1) depositing endosteal bone coatings on the inner cortex and over struts of pre-existing internal bone, (2) secondarily filling bone spaces, and (3) Haversian reworking. The construction of these struts reflects both developmental patterns of bone construction and biomechanical function. Alternating plywood-like layers of bone, heretofore undescribed in tetrapods, provided strength, as did the obliquely oriented system of reticular blood vessels in the bones. The distribution and ontogenetic features of pterosaur bone tissues, when combined with other evidence, suggest generally high growth rates, high metabolic levels, altricial birth, and extended parental care.     

First record of a pterosaur landing trackway

The terrestrial progression of pterosaurs, the flying reptiles of the Mesozoic Era, has been debated for over two centuries. The recent discovery of quadrupedal pterodactyloid pterosaur tracks from Late Jurassic sediments near Crayssac, France, shows that the hindlimbs moved parasagittally, as in mammals, birds and other dinosaurs, and the hypertrophied forelimbs could make tracks both close to the body wall and far outside it. Their manus tracks are unique in form, position and kinematics, which would be expected because the forelimbs were used for flight. Here, we report the first record of a pterosaur landing track, which differs substantially from typical walking trackways. The individual landed on both hind feet in parallel fashion, dragged its toes slightly as it left the track, landed again almost immediately and placed the hindfeet parallel again, then placed its forelimbs on the ground, took another short step with both hindlimbs and adjusted its forelimbs, and then began to walk off normally. The trackway shows that pterosaurs stalled to land, a reflection of their highly developed capacity for flight control and manoeuverability.     

Do different disparity proxies converge on a common signal? Insights from the cranial morphometrics and evolutionary history of Pterosauria (Diapsida: Archosauria)

Disparity, or morphological diversity, is often quantified by evolutionary biologists investigating the macroevolutionary history of clades over geological timescales. Disparity is typically quantified using proxies for morphology, such as measurements, discrete anatomical characters, or geometric morphometrics. If different proxies produce differing results, then the accurate quantification of disparity in deep time may be problematic. However, despite this, few studies have attempted to examine disparity of a single clade using multiple morphological proxies. Here, as a case study for this question, we examine the disparity of the volant Mesozoic fossil reptile clade Pterosauria, an intensively studied group that achieved substantial morphological, ecological and taxonomic diversity during their 145+ million-year evolutionary history. We characterize broadscale patterns of cranial morphological disparity for pterosaurs for the first time using landmark-based geometric morphometrics and make comparisons to calculations of pterosaur disparity based on alternative metrics. Landmark-based disparity calculations suggest that monofenestratan pterosaurs were more diverse cranially than basal non-monofenestratan pterosaurs (at least when the aberrant anurognathids are excluded), and that peak cranial disparity may have occurred in the Early Cretaceous, relatively late in pterosaur evolution. Significantly, our cranial disparity results are broadly congruent with those based on whole skeleton discrete character and limb proportion data sets, indicating that these divergent approaches document a consistent pattern of pterosaur morphological evolution. Therefore, pterosaurs provide an exemplar case demonstrating that different proxies for morphological form can converge on the same disparity signal, which is encouraging because often only one such proxy is available for extinct clades represented by fossils. Furthermore, mapping phylogeny into cranial morphospace demonstrates that pterosaur cranial morphology is significantly correlated with, and potentially constrained by, phylogenetic relationships.