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      Discovery of a Rare Pterosaur Bone Bed in a Cretaceous Desert with Insights on Ontogeny and Behavior of Flying Reptiles

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          Abstract

          A pterosaur bone bed with at least 47 individuals (wing spans: 0.65–2.35 m) of a new species is reported from southern Brazil from an interdunal lake deposit of a Cretaceous desert, shedding new light on several biological aspects of those flying reptiles. The material represents a new pterosaur, Caiuajara dobruskii gen. et sp. nov., that is the southermost occurrence of the edentulous clade Tapejaridae (Tapejarinae, Pterodactyloidea) recovered so far. Caiuajara dobruskii differs from all other members of this clade in several cranial features, including the presence of a ventral sagittal bony expansion projected inside the nasoantorbital fenestra, which is formed by the premaxillae; and features of the lower jaw, like a marked rounded depression in the occlusal concavity of the dentary. Ontogenetic variation of Caiuajara dobruskii is mainly reflected in the size and inclination of the premaxillary crest, changing from small and inclined (∼115°) in juveniles to large and steep (∼90°) in adults. No particular ontogenetic features are observed in postcranial elements. The available information suggests that this species was gregarious, living in colonies, and most likely precocial, being able to fly at a very young age, which might have been a general trend for at least derived pterosaurs.

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          Pterosaur diversity and faunal turnover in Cretaceous terrestrial ecosystems in China.

          New specimens and an analysis of the Jehol pterosaur faunae of northeastern China show an unexpected diversity of flying reptile groups in terrestrial Cretaceous ecosystems. Here we report two new pterosaurs that are referred to European groups previously unknown in deposits of northeastern China. Feilongus youngi, from the Yixian Formation, is closely related to the Gallodactylidae and is distinguished by the presence of two independent sagittal crests and a protruding upper jaw. Nurhachius ignaciobritoi, from the Jiufotang Formation, has teeth formed by labiolingually compressed triangular crowns, only previously reported in Istiodactylus latidens from England. With these new discoveries, the Jehol pterosaurs show a wide range of groups including both primitive and derived forms that are not matched by any other deposit in the world. The discoveries also document the turnover of pterosaur faunae, with the primitive Anurognathidae and early archaeopterodactyloids being replaced by derived pterodactyloids. Furthermore, these deposits offer an opportunity to examine the interaction and competition between birds and pterosaurs--it indicates that the avian fauna during the Lower Cretaceous (and possibly most of the Mesozoic) dominated terrestrial, inland regions, whereas pterosaurs were more abundant in coastal areas.
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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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              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. Considerado um dos répteis voadores melhor conhecidos, Pteranodon foi objeto de várias revisões no século passado. Encontrado exclusivamente na Formação Niobrara e no Grupo Pierre Shale (ambas do Cretáceo Superior) 11 espécies foram atribuídas a este gênero (além das três presentemente classificadas em Nyctosaurus). Apesar da concordância entre os revisores que este número está inflado, existe discordância de quantas espécies podem ser reconhecidas. A última revisão limitou a existência de apenas duas espécies (Pteranodon longiceps and Pteranodon sternbergi), ambas exibindo dimorfismo sexual. Tomando como base diversas características observadas no crânio, alguns exemplares anteriormente referidos ao gênero Pteranodon foram re-avaliados e levaram à conclusão da existência das seguintes espécies, duas novas e descritas aqui: Pteranodon longiceps, Geosternbergia sternbergi, Geosternbergia maiseyi sp. nov. e Dawndraco kanzai gen. et sp. nov. Estas se diferenciam por feições tais como a extensão e direção da crista formada pelo frontal, da inclinação e extensão do processo posterior das pré-maxilas, da forma e tamanho da fenestra temporal inferior e da proporção do rostro. Também é discutido nesse trabalho o procedimento de como se reconhece uma espécie de pterossauro, que deve levar em conta principalmente dados morfológicos, acompanhados de informações estratigráficas e geográficas. Existe a consciência geral de que a morfologia pode variar sem que necessariamente estas mudanças estejam ligadas a questões taxonômicas. No entanto, a ausência de dados estratigráficos detalhados, aliada ao número limitado de exemplares que possam ser consideradas com um bom nível de segurança como pertencentes a uma determinada espécie, dificulta sobremaneira o nosso entendimento de como a morfologia pode variar em função de ontogenia, variações individuais e dimorfismo sexual. Apesar do presente estudo não eliminar a possibilidade de reconhecer estas variações, cuidado deve ser empregado antes que modelos sejam generalizados para os pterossauros.
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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
                2014
                13 August 2014
                : 9
                : 8
                : e100005
                Affiliations
                [1 ]Centro Paleontológico da UnC (CENPÁLEO), Universidade do Contestado, Mafra, Santa Catarina, Brazil
                [2 ]Programa de Pós-Graduação IEL-Labjor, Universidade Estadual de Campinas (UNICAMP), Campinas, São Paulo, Brazil
                [3 ]Laboratory of Systematics and Taphonomy of Fossil Vertebrates, Departamento de Geologia e Paleontologia, Museu Nacional/Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
                [4 ]Universidade Estadual de Ponta Grossa, Ponta Grossa, Paraná, Brazil
                [5 ]Departamento de Geologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil
                [6 ]Departamento de Geociências, Universidade Estadual de Ponta Grossa, Ponta Grossa, Paraná, Brazil
                Raymond M. Alf Museum of Paleontology, United States of America
                Author notes

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

                Conceived and designed the experiments: AWAK PCM LCW CEF. Analyzed the data: AWAK PCM LCW CEF CSV GBG LCG AL JHZR CCM. Wrote the paper: AWAK PCM LCW CEF CSV GBG LCG AL JHZR CCM.

                Article
                PONE-D-14-10026
                10.1371/journal.pone.0100005
                4131874
                25118592
                d4a0b256-5daf-4484-aaae-4891370aad03
                Copyright @ 2014

                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
                : 6 March 2014
                : 18 May 2014
                Page count
                Pages: 10
                Funding
                AWAK acknowledges funding from the Fundação Carlos Chagas Filho de Amparo à pesquisa do Rio de Janeiro (FAPERJ # E-26/102.737/2012) and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq # 307276/2009-9). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Paleontology
                Paleobiology
                Paleozoology
                Vertebrate Paleontology
                Earth Sciences
                Custom metadata
                The authors confirm that all data underlying the findings are fully available without restriction. All data are included within the paper.

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