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      A New Azhdarchid Pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: Implications for Azhdarchid Diversity and Distribution

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          Abstract

          We describe a new taxon of medium-sized (wing span ca. 3 m) azhdarchid pterosaur from the Upper Cretaceous Transylvanian Basin (Sebeş Formation) of Romania. This specimen is the most complete European azhdarchid yet reported, comprising a partially articulated series of vertebrae and associated forelimb bones. The new taxon is most similar to the Central Asian Azhdarcho lancicollis Nessov but possesses a suite of autapomorphies in its vertebrae that include the relative proportions of cervicals three and four and the presence of elongated prezygapophyseal pedicles. The new taxon is interesting in that it lived contemporaneously with gigantic forms, comparable in size to the famous Romanian Hatzegopteryx thambema. The presence of two distinct azhdarchid size classes in a continental depositional environment further strengthens suggestions that these pterosaurs were strongly linked to terrestrial floodplain and wooded environments. To support this discussion, we outline the geological context and taphonomy of our new specimen and place it in context with other known records for this widespread and important Late Cretaceous pterosaurian lineage.

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          A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology

          Azhdarchid pterosaurs were among the most widespread and successful of pterosaur clades, but their paleoecology remains controversial. Morphological features common to all azhdarchids include a long, shallow rostrum; elongate, cylindrical cervical vertebrae that formed a long and unusually inflexible neck; and proportionally short wings with an abbreviated fourth phalanx. While azhdarchids have been imagined as vulture-like scavengers, sediment probers, swimmers, waders, aerial predators, or stork-like generalists, most recent authors have regarded them as skim-feeders, trawling their lower jaws through water during flight and seizing aquatic prey from the water's surface. Although apparently widely accepted, the skim-feeding model lacks critical support from anatomy and functional morphology. Azhdarchids lack the many cranial specialisations exhibited by extant skim-feeding birds, most notably the laterally compressed lower jaw and shock absorbing apparatus required for this feeding style. Well-preserved azhdarchid skulls are rare, but their rostra and lower jaws appear to have been sub-triangular in cross-section, and thus dissimilar to those of skim-feeders and sediment probers. Taphonomic data indicates that azhdarchids predominately inhabited inland settings, and azhdarchid morphology indicates that they were poorly suited for all proposed lifestyles bar wading and terrestrial foraging. However, azhdarchid footprints show that their feet were relatively small, padded and slender, and thus not well suited for wading. We argue that azhdarchids were stork- or ground hornbill-like generalists, foraging in diverse environments for small animals and carrion. Proficient terrestrial abilities and a relatively inflexible neck are in agreement with this interpretation.
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            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.
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              A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution.

              Toothless pterosaurs played a key role in broadening the taxonomic, morphological and ecological diversity of Cretaceous pterosaurs. Here we report a complete, articulated skeleton of a 1.4-m-wingspan pterosaur from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China which is identified as a new genus and species, Shenzhoupterus chaoyangensis gen. et sp. nov. The new taxon is edentulous, with a relatively large skull and a remarkably large, tall nasoantorbital fenestra that extends well above the main part of the braincase. Phylogenetic analysis shows that Shenzhoupterus gen. nov. belongs in a distinct clade of azhdarchoid pterosaurs, formally recognised here as a new family, Chaoyangopteridae, that also includes Chaoyangopterus, Jidapterus and Eoazhdarcho from the Jiufotang Formation and Eopteranodon from the Yixian Formation. These new data clarify recent confusion surrounding the systematics of these Lower Cretaceous taxa and provide new insights into the evolutionary history of pterosaurs.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2013
                30 January 2013
                : 8
                : 1
                : e54268
                Affiliations
                [1 ]Department of Natural Sciences, Transylvanian Museum Society, Cluj-Napoca, Romania
                [2 ]Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Department of Geology and Paleontology, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, São Cristóvão, Brazil
                [3 ]Ocean and Earth Sciences, University of Southampton, Southampton, United Kingdom
                [4 ]Institute of Life Sciences, University of Southampton, Southampton, United Kingdom
                Team ‘Evo-Devo of Vertebrate Dentition’, France
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MV AK DN GD. Performed the experiments: MV AK DN GD. Analyzed the data: MV AK DN GD. Contributed reagents/materials/analysis tools: MV AK DN GD. Wrote the paper: MV AK DN GD.

                Article
                PONE-D-12-17833
                10.1371/journal.pone.0054268
                3559652
                23382886
                35b6054a-f3c7-4ad9-8f7d-3995d05fe388
                Copyright @ 2013

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 18 June 2012
                : 10 December 2012
                Page count
                Pages: 15
                Funding
                The authors have no support or funding to report.
                Categories
                Research Article
                Biology
                Evolutionary Biology
                Paleontology
                Vertebrate Paleontology
                Paleontology
                Vertebrate Paleontology
                Earth Sciences
                Paleontology
                Paleoecology
                Vertebrate Paleontology

                Uncategorized
                Uncategorized

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