Comments on the cervical vertebrae of the Tapejaridae (Pterosauria, Pterodactyloidea) with description of new specimens

BackgroundIn the Early Cretaceous Jehol Biota, the toothless pterosaurs flourished with the chaoyangopterids and tapejarids playing a key role in understanding the early diversity and evolution of the Azhdarchoidea. Unlike the more diverse tapejarids, the rarer chaoyangopterids are characterized by a long and low rostrum, supporting a close relationship with the huge azhdarchids. Unfortunately, our knowledge is still limited in the osteology, paleoecology, and taxonomy of the Chaoyangopteridae. As one of the best preserved skeletons, the type and only specimen of Jidapterus edentus provides an opportunity to understand the morphology and paleoecology of the chaoyangopterids.ResultsOur study of the osteology of Jidapterus edentus reveals valuable information about the morphology of the Chaoyangopteridae such as a rostrum with a curved dorsal profile, high Rostral Index (RI), larger angle between the dorsal and postorbital processes of the jugal, sequentially shorter fourth to seventh cervical vertebrae, sternum with a plate wider than long, contact of the metacarpal I with the distal syncarpal, pneumatic foramen on first wing phalanx, hatchet-like postacetabular process with unconstricted neck and small dorsal process, distinctly concave anterior margin of pubis, subrectangular pubic plate with nearly parallel anterior and posterior margins, longer proximal phalanges of pedal digits III and IV, as well as reduced and less curved pedal unguals. These features further support the validity of Jidapterus edentus as a distinct species and the close relationship of the chaoyangopterids with the azhdarchids. Paleoecologically, the chaoyangopterids are probably like the azhdarchids, more terrestrial than the contemporaneous and putatively arboreal tapejarids, which may have been limited to the forest-dominated ecosystem of the Jehol Biota.DiscussionThe osteology of Jidapterus edentus further supports the close relationship of the Chaoyangopteridae with the Azhdarchidae in sharing a high RI value and reduced and mildly-curved pedal unguals, and it also implies a possible paleoecological similarity in their terrestrial capability. Combined with the putatively arboreal and herbivorous tapejarids, this distinct lifestyle of the chaoyangopterids provides new insights into the diversity of pterosaurs in the ecosystem of the Jehol Biota.

The literature on crocodylian anatomy presents the transverse process in an ambiguous meaning, which could represent all lateral expansions derived from the neural arch, including vertebrae from cervical to caudal series, or in a more restrictive meaning, being applied only to lumbar vertebrae. The lateral expansion of sacral and caudal vertebrae usually referred to as the transverse process has been discovered to be fused ribs, bringing more ambiguity to this term. Therefore, with the lack of a definition for transverse process and other associated terms, the present work aims to propose a nomenclatural standardization, as well as definitions and biological meaning, for vertebral rib related structures. Vertebra obtained from museum collections from a total of 87 specimens of 22 species of all extant Crocodylia genera were studied. All vertebrae, except cervical and first three dorsal, exhibit transverse processes. The transverse process is more developed in dorsal and lumbar vertebrae than in sacral and caudal vertebrae in which it is suppressed by the fused ribs. The serial homology hypotheses here proposed can also be aplied to other Crurotarsi and saurischian dinosaurs specimens. This standardization clarifies the understand of the serial homology among those homotypes, and reduces the ambiguity and misleadings in future work comparisons. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc.

Birds and pterosaurs have pneumatic bones, a feature likely related to their flight capabilities but whose evolution and origin is still poorly understood. Pneumatic foramina are present on the external surface of the bone and are reliable indicators of post-cranial skeletal pneumatization present in Pterosauria, Eusauropoda, and Neotheropoda. Here, we carried out a qualitative analysis of the position, size and number of pneumatic foramina of the cervical and thoracic/dorsal vertebrae of pterosaurs and birds, as they have the potential to challenge hypotheses about the emergence and evolution of the respiratory trait in these groups. We also discussed differences between pneumatic and vascular foramina for identification purposes. Besides phylogenetic representativeness, the pterosaur taxonomic sampling considered the preservation of specimens and, for birds, their life habit, as this relates to the level of pneumatization. Pneumatic foramina on the lateral faces of the centrum of the mid-cervical vertebrae of pterosaurs and birds differ in position and size, and those adjacent to the neural canal additionally differ in number. The avian posterior cervical vertebrae show a higher number of pneumatic foramina in comparison to their mid-cervicals, while the opposite is true for pterosaurs, suggesting differences in the cervical air sac of these clades. Pneumatic foramina were found at the base of the transverse processes of the notarial vertebrae of birds, while they were absent from some of the pterosaurs analyzed here, revealing the presence of a pneumatic hiatus in the vertebral column that might be explained due to the distance of this structure to the cervical air sac. These findings indicate that, although the overall skeletal pneumatization of pterosaurs and birds present deep homologies, some pneumatic features occurred convergently because variation in the number of pneumatic foramina along the vertebral column is related to the position of the air sacs in pterosaurs and birds and/or the habit of each species. There is an evident reduction of the pneumatic foramina in birds that have aquatic foraging and an increase in the ones which perform static soaring. Although we did not find any external anatomical difference between pneumatic and vascular foramina, we observed that vascular foramina occur at specific sites and thus identification on the basis of location is reliable.