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      A new species of Allodaposuchus (Eusuchia, Crocodylia) from the Maastrichtian (Late Cretaceous) of Spain: phylogenetic and paleobiological implications

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          Abstract

          Background. The Late Cretaceous is a keystone period to understand the origin and early radiation of Crocodylia, the group containing all extant lineages of crocodilians. Among the taxa described from the latest Cretaceous of Europe, the genus Allodaposuchus is one of the most common but also one of the most controversial. However, because of its fragmentary record, several issues regarding its phylogenetic emplacement and its ecology remain unsolved or unknown. The discovery of a single specimen attributed to Allodaposuchus, represented by both cranial and postcranial remains, from the Casa Fabà site (Tremp Basin, NE Spain) in the lower red unit of the Tremp Fm. (early Maastrichtian, Late Cretaceous) offers a unique opportunity to deepen in the phylogenetic relationships of the group and its ecological features.

          Methods. The specimen is described in detail, and CT scan of the skull is performed in order to study the endocranial morphology as well as paratympanic sinuses configuration. In addition, myological and phylogenetic analyses are also carried out on the specimen for to shed light in ecological and phylogenetic issues, respectively.

          Results. The specimen described herein represents a new species, Allodaposuchus hulki sp. nov ., closely related to the Romanian A. precedens. The CT scan of the skull revealed an unexpected paratympanic sinuses configuration. Allosaposuchus hulki exhibits an “anterodorsal tympanic sinus” not observed in any other extant or extinct crocodilian. The caudal tympanic recesses are extremely enlarged, and the expanded quadratic sinus seems to be connected to the middle-ear channel. Phylogenetic analyses confirm the emplacement of the informal taxonomic group ‘Allodaposuchia’ at the base of Crocodylia, being considered the sister group of Borealosuchus and Planocraniidae.

          Discussion. Although this is a preliminary hypothesis, the unique paratympanic configuration displayed by A. hulki suggests that it could possess a high-specialized auditory system. Further, the large cranial cavities could help to reduce the weight of the cranium. Concerning the postcranial skeleton, Allodaposuchus hulki shows massive and robust vertebrae and forelimb bones, suggesting it could have a bulky body. The myological study performed on the anterior limb elements supports this interpretation. In addition, several bone and muscular features seem to point at a semi-erected position of the forelimbs during terrestrial locomotion. Taking all the above results into consideration, it seems plausible to suggest that A. hulki could conduct large incursions out of the water and have a semi-terrestrial lifestyle.

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          The origin of modern crocodyliforms: new evidence from the Cretaceous of Australia.

          While the crocodyliform lineage extends back over 200 million years (Myr) to the Late Triassic, modern forms-members of Eusuchia-do not appear until the Cretaceous. Eusuchia includes the crown group Crocodylia, which comprises Crocodyloidea, Alligatoroidea and Gavialoidea. Fossils of non-crocodylian eusuchians are currently rare and, in most instances, fragmentary. Consequently, the transition from Neosuchia to Crocodylia has been one of the most poorly understood areas of crocodyliform evolution. Here we describe a new crocodyliform from the mid-Cretaceous (98-95 Myr ago; Albian-Cenomanian) Winton Formation of Queensland, Australia, as the most primitive member of Eusuchia. The anatomical changes associated with the emergence of this taxon indicate a pivotal shift in the feeding and locomotor behaviour of crocodyliforms-a shift that may be linked to the subsequent rapid diversification of Eusuchia 20 Myr later during the Late Cretaceous and Early Tertiary. While Laurasia (in particular North America) is the most likely ancestral area for Crocodylia, the biogeographic events associated with the origin of Eusuchia are more complex. Although the fossil evidence is limited, it now seems likely that at least part of the early history of Eusuchia transpired in Gondwana.
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            Crocodylian forelimb musculature and its relevance to Archosauria.

            The musculoskeletal anatomy of the crocodylian forelimb is documented to facilitate functional morphological studies of extant and extinct archosaurs. Comparative descriptions of muscles of the forelimb of several crocodylian species are presented, including attachment sites, innervation, and anatomical functions. The muscular anatomy of the crocodylian forelimb is highly conservative among the different species; however, interspecific differences do occur. Interspecific anatomical variation is interpreted functionally, and discussed in the context of the terrestrial locomotion of crocodylians as it applies to the forelimb. In addition, muscular apomorphies are identified among a phylogenetically diverse sample of extant crocodylians, providing insight into the evolution of forelimb anatomy in a clade of archosaurs possessing highly variable terrestrial locomotor behaviors. Copyright 2003 Wiley-Liss, Inc.
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              Cretaceous Crocodyliforms from the Sahara

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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                PeerJ
                PeerJ
                PeerJ
                PeerJ Inc. (San Francisco, USA )
                2167-8359
                13 August 2015
                2015
                : 3
                : e1171
                Affiliations
                [1 ]Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona , Sabadell, Catalonia, Spain
                [2 ]Departament d’Estratigrafia, Paleontologia i Geociències marines, Facultat de Geologia, Universitat de Barcelona, Carrer de Martí i Franquès s/n , Barcelona, Catalonia, Spain
                Article
                1171
                10.7717/peerj.1171
                4558081
                26339549
                5b2fa439-47c7-48ae-b1bd-ecc5118b8165
                © 2015 Blanco 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
                : 20 March 2015
                : 20 July 2015
                Funding
                Funded by: Spanish Ministry of Economy and Competitiveness (MINECO)
                Award ID: CGL2011-30069-C02-01/BTE
                Award ID: CGL2011-27869
                Funded by: Generalitat de Catalunya (Catalan Government)
                Award ID: FI-DGR 2013FI_B 01059
                Award ID: FI-DGR 2012FI_B 01221
                Funded by: Spanish Ministry of Economy and Competitiveness
                Award ID: BES-2012-057837
                This paper is a contribution to the projects CGL2011-30069-C02-01/BTE and CGL2011-27869 subsidized by the Spanish Ministry of Economy and Competitiveness (MINECO). The research of AB and ÀHL is supported by predoctoral grants (FI-DGR 2013FI_B 01059 and FI-DGR 2012FI_B 01221, respectively) awarded by the Generalitat de Catalunya (Catalan Government), whereas AV is granted by the Spanish Ministry of Economy and Competitiveness (BES-2012-057837). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Animal Behavior
                Paleontology
                Taxonomy
                Zoology

                crocodylia,endocranial morphology,locomotion,allodaposuchus,paleoecology,late cretaceous

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