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      New information on the Wukongopteridae (Pterosauria) revealed by a new specimen from the Jurassic of China

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          Abstract

          The Wukongopteridae is an important pterosaur group discovered from Yanliao Biota, because it combines character states seen in non-pterodactyloid and pterodactyloid pterosaurs. So far, the Wukongopteridae contains three genera: Wukongopterus, Darwinopterus and Kunpengopterus; representing five species. Here we report on a new specimen, IVPP V 17959, that can be undoubtedly referred to the Wukongopteridae based on the presence of a confluent nasoantorbital fenestra, elongated cervical vertebrae (convergent with Pterodactyloidea) and a long tail enclosed by rod-like bony extensions of the zygapophyses. Traits distinguishing this new specimen from other wukongopterid pterosaurs include a premaxilla with a low ossified anterodorsal crest, a nasal bearing the most elongated process known in the Wukongopteridae, and a lacrimal that has a foramen in its middle portion. The new kind of premaxillary crest preserved in IVPP V 17959 suggests that the presence or absence of a premaxillary crest might be an interspecific feature within the Wukongopteridae. A phylogenetic analysis including all wukongopterid pterosaurs recovers IVPP V 17959 in a polytomy with Wukongopterus and the species of Darwinopterus, having Kunpengopterus in a more basal position. The postcranial skeleton of IVPP V 17959 has ontogenetically mature characteristics including a completely fused scapula and coracoid, fused proximal and distal carpal series, and an ossified extensor tendon process of the first wing phalanx, allowing its classification as ontogenetic stage five. Furthermore, the atlas and axis are separated in IVPP V 17959, which indicates that these two bones probably are not fused in skeletally mature wukongopterid individuals.

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          The earliest pterodactyloid and the origin of the group.

          The pterosaurs were a diverse group of Mesozoic flying reptiles that underwent a body plan reorganization, adaptive radiation, and replacement of earlier forms midway through their long history, resulting in the origin of the Pterodactyloidea, a highly specialized clade containing the largest flying organisms. The sudden appearance and large suite of morphological features of this group were suggested to be the result of it originating in terrestrial environments, where the pterosaur fossil record has traditionally been poor [1, 2], and its many features suggested to be adaptations to those environments [1, 2]. However, little evidence has been available to test this hypothesis, and it has not been supported by previous phylogenies or early pterodactyloid discoveries. We report here the earliest pterosaur with the diagnostic elongate metacarpus of the Pterodactyloidea, Kryptodrakon progenitor, gen. et sp. nov., from the terrestrial Middle-Upper Jurassic boundary of Northwest China. Phylogenetic analysis confirms this species as the basalmost pterodactyloid and reconstructs a terrestrial origin and a predominantly terrestrial history for the Pterodactyloidea. Phylogenetic comparative methods support this reconstruction by means of a significant correlation between wing shape and environment also found in modern flying vertebrates, indicating that pterosaurs lived in or were at least adapted to the environments in which they were preserved.
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            Pterosaur phylogeny and comments on the evolutionary history of the group

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              Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull.

              The fossil record is a unique source of evidence for important evolutionary phenomena such as transitions between major clades. Frustratingly, relevant fossils are still comparatively rare, most transitions have yet to be documented in detail and the mechanisms that underpin such events, typified by rapid large scale changes and for which microevolutionary processes seem insufficient, are still unclear. A new pterosaur (Mesozoic flying reptile) from the Middle Jurassic of China, Darwinopterus modularis gen. et sp. nov., provides the first insights into a prominent, but poorly understood transition between basal, predominantly long-tailed pterosaurs and the more derived, exclusively short-tailed pterodactyloids. Darwinopterus exhibits a remarkable 'modular' combination of characters: the skull and neck are typically pterodactyloid, exhibiting numerous derived character states, while the remainder of the skeleton is almost completely plesiomorphic and identical to that of basal pterosaurs. This pattern supports the idea that modules, tightly integrated complexes of characters with discrete, semi-independent and temporally persistent histories, were the principal focus of natural selection and played a leading role in evolutionary transitions.
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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                peerj
                peerj
                PeerJ
                PeerJ Inc. (San Francisco, USA )
                2167-8359
                7 July 2016
                2016
                : 4
                : e2177
                Affiliations
                [1 ]Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology , Beijing, China
                [2 ]University of Chinese Academy of Sciences , Beijing, China
                [3 ]Department of Geology and Paleontology, National Museum/UFRJ , Rio de Janeiro, Brazil
                Article
                2177
                10.7717/peerj.2177
                4941781
                27441118
                c559114a-21bf-4581-b625-2c1720e9bc54
                ©2016 Cheng et al.

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

                History
                : 14 January 2016
                : 5 June 2016
                Funding
                Funded by: National Key Basic Research Program of China
                Award ID: 2012CB821900
                Funded by: National Natural Science Foundation of China
                Award ID: 41572020
                Funded by: Major Research plan of the National Natural Science Foundation of China
                Award ID: 91514302
                Funded by: Hundred Talents Project of Chinese Academy of Sciences
                Funded by: National Science Fund for Distinguished Young Scholars
                Award ID: 40825005
                Funded by: Fundação Carlos Chagas Filho de Amparo à pesquisa do Rio de Janeiro
                Award ID: FAPERJ # E-26/102.737/2012
                Funded by: Conselho Nacional de Desenvolvimento Científico e Tecnológico
                Award ID: CNPq # 304780/2013-8
                This study was supported by the National Key Basic Research Program of China (2012CB821900), the National Natural Science Foundation of China (41572020), the Major Research plan of the National Natural Science Foundation of China (91514302), the Hundred Talents Project of Chinese Academy of Sciences, and the National Science Fund for Distinguished Young Scholars (40825005). AWAK acknowledges founding 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 # 304780/2013-8). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Paleontology

                wukongopteridae,pterosauria,yanliao biota,china,late jurassic

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