Dinosaurs and other tetrapods in an Early Cretaceous bauxite-filled fissure, northwestern Romania

The bauxite mine at Cornet near Oradea in northwestern Romania produced thousands of bones in an excavation in 1978, mainly from ornithopod dinosaurs and rarer pterosaurs. Bird specimens reported previously from this fauna are equivocal. The fossils are disarticulated bones in good condition which occur highly concentrated in lenses within bauxite clays, which are dated as Berriasian (earliest Cretaceous). The bauxite represents detrital material washed into deep fissures and caves formed within a karst of uplifted Tithonian (latest Jurassic) marine limestones. The bones are generally uniform in size and shape, and they are abraded, evidence for considerable transport and for winnowing of the deposit. The area was one of several islands on the northern shore of Tethys, and it was inundated by the sea later in the Early Cretaceous. There is evidence for insular adaptations in the dinosaur faunas. The ornithopod dinosaurs may include several taxa, but they are smaller on average than an assemblage of typical Wealden ornithopods, perhaps because of dwarfing on the island. In addition, sauropods are absent and theropods are barely represented in the fauna. The fauna is geographically significant since it shows relationships with western Europe and with Asia.     

Partial mandible of a giant pterosaur from the uppermost Cretaceous (Maastrichtian) of the Hațeg Basin,Romania

We describe and interpret a posterior mandibular symphysis of a very large azhdarchid pterosaur. The specimen LPB (FGGUB ) R.2347 exhibits a series of morphological characters present in both azhdarchid and tapejarid pterosaurs, suggesting a more basal position within the clade Azhdarchidae. This fossil was collected from Maastrichtian continental deposits near V?lioara in the Ha?eg Basin, Romania, but cannot be confidently referred to the contemporaneous giant Hatzegopteryx thambema, also from V?lioara, due to the absence of overlapping skeletal elements. Remarkably, this mandibular symphysis shares a number of features the smaller azhdarchoid Bakonydraco galaczi from the Santonian of Hungary. Additional comparisons with previously described large‐sized azhdarchid mandibles indicate a certain degree of morphological and probably ecological disparity within the group. This specimen represents the largest pterosaur mandible ever found and provides insights into the anatomy of the enigmatic giant pterosaurs.     

A NEW PTEROSAUR FROM THE LIAONING PROVINCE OF CHINA, THE PHYLOGENY OF THE PTERODACTYLOIDEA, AND CONVERGENCE IN THEIR CERVICAL VERTEBRAE

Abstract:  The largest known flying organisms are the azhdarchid pterosaurs, a pterodactyloid clade previously diagnosed by the characters of their extremely elongate middle-series cervical vertebrae. The named species of the Azhdarchidae are from the Late Cretaceous. However, isolated mid-cervical vertebrae with similar dimensions and characters have been referred to this group that date back to the Late Jurassic, implying an almost 60 million year gap in the fossil record of this group and an unrecorded radiation in the Jurassic of all the major clades of the Pterodactyloidea. A new pterosaur from the Early Cretaceous of Liaoning Province of China, Elanodactylus prolatus gen. et sp. nov., is described with mid-cervical vertebrae that bear these azhdarchid characters but has other postcranial material that are distinct from the members of this group. Phylogenetic analysis of the new species and the Pterodactyloidea places it with the Late Jurassic vertebrae in the Late Jurassic–Early Cretaceous Ctenochasmatidae and reveals that the characters of the elongate azhdarchid vertebrae appeared independently in both groups. These results are realized though the large taxon sampling in the analysis demonstrating that the homoplastic character states present in these two taxa were acquired in a different order in their respective lineages. Some of these homoplastic characters were previously thought to appear once in the history of pterosaurs and may be correlated to the extension of the neck regions in both groups. Because the homoplastic character states in the Azhdarchidae and Ctenochasmatidae are limited to the mid-cervical vertebrae, these states are termed convergent based on a definition of the term in a phylogenetic context. A number of novel results from the analysis presented produce a reorganization in the different species and taxa of the Pterodactyloidea.     

The shape of pterosaur evolution: evidence from the fossil record

Abstract Although pterosaurs are a well‐known lineage of Mesozoic flying reptiles, their fossil record and evolutionary dynamics have never been adequately quantified. On the basis of a comprehensive data set of fossil occurrences correlated with taxon‐specific limb measurements, we show that the geological ages of pterosaur specimens closely approximate hypothesized patterns of phylogenetic divergence. Although the fossil record has expanded greatly in recent years, collectorship still approximates a sigmoid curve over time as many more specimens (and thus taxa) still remain undiscovered, yet our data suggest that the pterosaur fossil record is unbiased by sites of exceptional preservation (lagerstätte). This is because as new species are discovered the number of known formations and sites yielding pterosaur fossils has also increased – this would not be expected if the bulk of the record came from just a few exceptional faunas. Pterosaur morphological diversification is, however, strongly age biased: rarefaction analysis shows that peaks of diversity occur in the Late Jurassic and Early Cretaceous correlated with periods of increased limb disparity. In this respect, pterosaurs appear unique amongst flying vertebrates in that their disparity seems to have peaked relatively late in clade history. Comparative analyses also show that there is little evidence that the evolutionary diversification of pterosaurs was in any way constrained by the appearance and radiation of birds.     

Terrestrial locomotion in pterosaurs

On the basis of a well‐preserved pelvis of Anhanguera sp. from the Lower Cretaceous (Aptian) of the Chapada do Araripe, Brazil, the problem of terrestrial locomotion in pterosaurs is discussed. A three‐dimensional reconstruction of the pelvis led to a lateral, dorsal and posterior orientation of the acetabula. By use of the preserved proximal ends of the femora of the same individual, the articulation in the hip socket could be tested. The normal articulation of the femur resulted in a horizontal position of the femur shaft, probably during flight. For constructional reasons the femur could not be brought down to a vertical position. Therefore, a parasagittal swing of the femora necessary for a bird‐like stance and gait must have been impossible. It is suggested that in pterosaurs the wing membrane was attached to the upper leg, which helped in stretching, steering and cambering.

Moreover, on the basis of comparisons of the fossil preservation of pterosaurs Compsognathus and Archaeopteryx in the Solnhofen limestone, it is concluded that the femora of pterosaurs were splayed out laterally, and that they had a semi‐erect gait. They were not bipedal animals, but had to use their fore limbs as well on the ground. Nevertheless, as vertebrates extremely adapted to flight, they could not have been able quadrupeds, either.     

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.     

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.     

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 new species of Jeholornis with complete caudal integument

The Early Cretaceous long bony-tailed bird Jeholornis prima displays characters both more basal than Archaeopteryx and more derived, exemplifying the mosaic distribution of advanced avian features that characterises early avian evolution and obfuscates attempts to understand early bird relationships. The current diversity of Jeholornithiformes is controversial, since multiple possibly synonymous genera were named simultaneously. Here, we provide the first definitive evidence of a second species belonging to this clade, and erect the new taxon J. palmapenis sp. nov. This new specimen reveals the tail integument of Jeholornithiformes, the morphology of which appears to have no aerodynamic benefit suggesting this clade evolved plumage patterns that were primarily for display.     

REPTILIAN PHYSIOLOGY AND THE FLIGHT CAPACITY OF ARCHAEOPTERYX

Current scenarios frequently interpret the Late Jurassic bird Archaeopteryx as having had an avian-type physiology and as having been capable of flapping flight, but only from “the trees downward.” It putatively lacked capacity for takeoff and powered flight from the ground upward. Data from extant reptiles indicate that if Archaeopteryx were physiologically reptilian, it would have been capable of ground upward takeoff from a standstill, as well as “trees downward” powered flight. This conclusion is based largely on a previously unrecognized attribute of locomotory (skeletal) muscle in a variety of extant reptiles: During “burst-level” activity, major locomotory muscles of a number of active terrestrial taxa generate at least twice the power (watts kg^?1 muscle tissue) as those of birds and mammals. Reptilian physiological status also helps resolve the apparently uneven development of various flight support structures in Archaeopteryx (e.g., well-developed flight features but relatively unspecialized pectoral girdle, supracoracoideus muscles, etc.). Endothermy and capacity for longer-distance powered flight probably evolved only in Early Cretaceous birds, which were the first birds to have a keeled sternum, strap-like coracoid, and hypocleidium-bearing furcula.     

Mesozoic birds of China—a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper. ⁴⁰Ar/³⁹Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation. Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian) of the middle and late Early Cretaceous, respectively. In order to further evaluate the change of the avian diversity in the Jehol Biota, six new orders and families are erected based on known genera and species, which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearance of birds in the Late Jurassic. Chinese Early Cretaceous birds had experienced a significant differentiation in morphology, flight, diet and habitat. Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux. However, the attachment of metatarsal I to the medial side of metatarsal II does not preclude trunk climbing, a pre-adaptation for well developed perching life of early birds. Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight, and the adaptation to lakeshore environment had played an equally important role in the origin of ornithurine birds and their near-modern flight skill. Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet, showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g., piscivorous) dietary adaptation had appeared only in later advanced forms. The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history, and the size of the embryo and other analysis indicate it probably had a short incubation period. Leg feathers probably have a wide range of distribution in early birds, further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds. Finally, the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem. The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous. Selected from Vertebrata PalAsiatica 2006, 44 (1): 74–98     

Review of taxonomy,geographic distribution,and paleoenvironments of Azhdarchidae (Pterosauria)

The taxonomy, geographic distribution, and paleoenvironmental context of azhdarchid pterosaurs are reviewed. All purported pteranodontid, tapejarid, and azhdarchid specimens from the Cenomanian Kem Kem beds of Morocco are referred to a single azhdarchid taxon, Alanqa saharica. The four proposed autapomorphies of Eurazhdarcho langendorfensis from the lower Maastrichtian Sebeş Formation of Romania are based on misinterpretations of material and this taxon is likely a subjective junior synonym of Hatzegopteryx thambema. Among 54 currently reported azhdarchid occurrences (51 skeletal remains and 3 tracks) 13% are from lacustrine deposits, 17% from fluvial plain deposits, 17% from coastal plain deposits, 18% from estuarine and lagoonal deposits, and 35% from costal marine deposits. Azhdarchids likely inhabited a variety of environments, but were abundant near large lakes and rivers and most common in nearshore marine paleoenvironments.     

Dinoflagellate cysts and microforaminifera of the Lower Cretaceous Yatria River section,Subarctic Ural,NW Siberia (Russia). Biostratigraphy,palaeoenvironmental and palaeogeographic discussion

Rich assemblages of phytoplankton and microforaminifera (foraminifera linings) are found in the marine Lower Cretaceous section from the Yatria River (Subarctic Ural). The Lower Cretaceous (Upper Volgian to Lower Hauterivian) dinocyst and microforaminifera biostratigraphy is calibrated against the ammonite zones. The dinocyst stratigraphy differs from Arctic Canada zonation, except for the Berriasian Paragonyaulacysta? borealis zone, and shows significant similarity with the zonal subdivision of Boreal regions of Europe.

The distribution of phytoplankton and microforaminifera is related to relative sea level variations. Consecutive changes of phytoplankton associations (2 in the Berriasian, 3 in the Valanginian-Early Hauterivian, 4 in the Early Hauterivian), lateral zonations of microforaminifera (2 in the Berriasian, 2 in the Valanginian) and their relation to environmental changes in the Lower Cretaceous seas of the Subarctic Ural Basin are established.     

An unusual edentulous pterosaur from the Early Cretaceous Romualdo Formation of Brazil

Numerous taxa make up the Early Cretaceous fauna of Brazil, including Ornithocheiroidea, Tapejaridae, Thalassodromidae, Chaoyangopteridae and a purported member of Azhdarchidae. Dsungaripteridae has only been tentatively assumed to be present in the form of ‘Santanadactylus’ spixi. New study of NMSG SAO 251093 (a specimen referred to Thalassodromeus sethi) suggests it is a previously unknown species of dsungaripterid, Banguela oberlii, tax. nov., differing from Thalassodromeus and other pterosaurs from the Early Cretaceous of Brazil by a unique combination of characters, including an upturned jaw tip, a short dorsal mandibular symphyseal shelf (dmss), and an autapomorphic thin crest placed halfway along the fused mandibular symphysis without a keel along the ventral margin of the jaw. B. oberlii, tax. nov., is referred to Dsungaripteridae based on a dmss no longer than the ventral shelf, U-shaped caudal margin of the ventral shelf and lateral margins of the mandibular symphysis concave in dorsal view. B. oberlii, tax. nov., is the youngest known dsungaripterid, and expands known morphological diversity in the clade as well as the Early Cretaceous pterosaur fauna of South America.     

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.     

Gargantuavis philoinos: Giant bird or giant pterosaur?

Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcranial elements (synsacrum and pelvis, femur) from Late Cretaceous (Campanian-Maastrichtian) localities in Southern France. It has recently been suggested that these remains in fact belong to giant pterosaurs. A detailed comparison between bones referred to Gargantuavis and the corresponding skeletal elements of pterosaurs reveals considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.     

A New Small-Bodied Azhdarchoid Pterosaur from the Lower Cretaceous of England and Its Implications for Pterosaur Anatomy,Diversity and Phylogeny

Background

Pterosaurs have been known from the Cretaceous sediments of the Isle of Wight (southern England, United Kingdom) since 1870. We describe the three-dimensional pelvic girdle and associated vertebrae of a small near-adult pterodactyloid from the Atherfield Clay Formation (lower Aptian, Lower Cretaceous). Despite acknowledged variation in the pterosaur pelvis, previous studies have not adequately sampled or incorporated pelvic characters into phylogenetic analyses.

Methodology/Principal Findings

The new specimen represents the new taxon Vectidraco daisymorrisae gen. et sp. nov., diagnosed by the presence of a concavity posterodorsal to the acetabulum and the form of its postacetabular process on the ilium. Several characters suggest that Vectidraco belongs to Azhdarchoidea. We constructed a pelvis-only phylogenetic analysis to test whether the pterosaur pelvis carries a useful phylogenetic signal. Resolution in recovered trees was poor, but they approximately matched trees recovered from analyses of total evidence. We also added Vectidraco and our pelvic characters to an existing total-evidence matrix for pterosaurs. Both analyses recovered Vectidraco within Azhdarchoidea.

Conclusions/Significance

The Lower Cretaceous strata of western Europe have yielded members of several pterosaur lineages, but Aptian pterosaurs from western Europe are rare. With a pelvis length of 40 mm, the new animal would have had a total length of c. 350 mm, and a wingspan of c. 750 mm. Barremian and Aptian pterodactyloids from western Europe show that small-bodied azhdarchoids lived alongside ornithocheirids and istiodactylids. This assemblage is similar in terms of which lineages are represented to the coeval beds of Liaoning, China; however, the number of species and specimens present at Liaoning is much higher. While the general phylogenetic composition of western European and Chinese communities appear to have been approximately similar, the differences may be due to different palaeoenvironmental and depositional settings. The western Europe pterodactyloid record may therefore be artificially low in diversity due to preservational factors.     

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.     

An Unusual Pterosaur Specimen (Pterodactyloidea, ?Azhdarchoidea) from the Early Cretaceous Romualdo Formation of Brazil,and the Evolution of the Pterodactyloid Palate

A new and unusual specimen of a probable azhdarchoid pterosaur is described for the Early Cretaceous (Albian) Romualdo Formation of Brazil. The specimen consists of a palate that, although fragmentary, has a unique morphology differing from all other known pterosaurs with preservation of palatal elements. The new specimen probably indicates the presence of a yet undescribed pterodactyloid taxon for Romualdo Formation and brings new information on pterosaur diversity of this sedimentary unity. Mainly due to the rarity of pterodactyloid specimens with palate preservation, this structure has been overlooked in this clade. Here, we reassess the palatal anatomy of Pterodactyloidea, revealing an intriguing variety of morphotypes and evolutionary trends, some of them described here for the first time. The morphological disparity displayed by different pterodactyloid taxa may be further evidence of the presence of diverse feeding strategies within the clade.     

The first specimen of Archaeopteryx from the Upper Jurassic Mörnsheim Formation of Germany

From an initial isolated position as the oldest evolutionary prototype of a bird, Archaeopteryx has, as a result of recent fossil discoveries, become embedded in a rich phylogenetic context of both more and less crownward stem-group birds. This has prompted debate over whether Archaeopteryx is simply a convergently bird-like non-avialan theropod. Here we show, using the first synchrotron microtomographic examination of the genus, that the eighth or Daiting specimen of Archaeopteryx possesses a character suite that robustly constrains it as a basal avialan (primitive bird). The specimen, which comes from the Mörnsheim Formation and is thus younger than the other specimens from the underlying Solnhofen Formation, is distinctive enough to merit designation as a new species, Archaeopteryx albersdoerferi sp. nov., but is recovered in close phylogenetic proximity to Archaeopteryx lithographica. Skeletal innovations of the Daiting specimen, such as fusion and pneumatization of the cranial bones, well vascularized pectoral girdle and wing elements, and a reinforced configuration of carpals and metacarpals, suggest that it may have had more characters seen in flying birds than the older Archaeopteryx lithographica. These innovations appear to be convergent on those of more crownward avialans, suggesting that Bavarian archaeopterygids independently acquired increasingly bird-like traits over time. Such mosaic evolution and iterative exploration of adaptive space may be typical for major functional transitions like the origin of flight.