Abstract
Carcharodontosaurs were the largest predatory dinosaurs, and their early evolutionary history seems to be more intricate than was previously thought. Until recently, carcharodontosaurs were restricted to a group of large theropods inhabiting the Late Cretaceous Gondwanan land masses1,2, but in the last few years Laurasian evidence3,4,5 has been causing a reevaluation of their initial diversification6. Here we describe an almost complete and exquisitely preserved skeleton of a medium-sized (roughly six metres long) theropod from the Lower Cretaceous series (Barremian stage) Konservat-Lagerstätte of Las Hoyas7 in Cuenca, Spain. Cladistic analysis supports the idea that the new taxon Concavenator corcovatus is a primitive member of Carcharodontosauria6, exhibiting two unusual features: elongation of the neurapophyses of two presacral vertebrae forming a pointed, hump-like structure and a series of small bumps on the ulna. We think that these bumps are homologous to quill knobs present on some modern birds; the knobs are related to the insertion area of follicular ligaments that anchor the roots of the flight feathers (remiges) to the arm. We propose that Concavenator has integumentary follicular structures inserted on the ulna, as in modern birds. Because scales do not have follicles, we consider the structures anchored to the Concavenator arms to be non-scale skin appendages homologous to the feathers of modern birds. If this is true, then the phylogenetic bracket for the presence of non-scale skin structures homologous to feathers in theropod dinosaurs would be extended to the Neotetanurae, enlarging the scope for explaining the origin of feathers in theropods.
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Acknowledgements
We thank S. Prieto for information about the location of the specimen; C. Díaz-Romeral, R. de la Fuente, M. García Oliva and N. Santamaría for participating in the field; P. García, M. Llandres, P. Moyano and J. Saiz for specimen preparation; L. Chiappe (Natural History Museum of LA County) for comments on the manuscript; E. Feeman and Plan de Promoción de la Investigación en la Universidad Nacional de Educación a Distancia for manuscript revision; R. Martín for the flesh reconstruction of Concavenator in Fig. 2; and S. Torralba for photography. We acknowledge the Museo de las Ciencias de Castilla-La Mancha for support and funding. Part of this research was financed by project CGL2005-05614 of the Spanish Ministerio de Ciencia e Innovación.
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All authors shared all the phases and topics of the work, both in research and in writing the manuscript. F.O. and F.E. were mainly involved in the description and phylogenetical analysis, whereas J.L.S. was mainly involved in the study of the integumentary evidence. All authors discussed the results and implications and commented on the manuscript at all stages.
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Supplementary Information
This file contains Supplementary Information comprising Scores for Concavenator corcovatus, Phylogenetic analysis, Selected measurements for appendicular elements of the holotype specimen, additional references, Supplementary Figures 1-5 with legends and Supplementary Pictures 1-5 with legends. Supplementary Information section 1 was corrected on 05 May 2011. (PDF 16262 kb)
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Ortega, F., Escaso, F. & Sanz, J. A bizarre, humped Carcharodontosauria (Theropoda) from the Lower Cretaceous of Spain. Nature 467, 203–206 (2010). https://doi.org/10.1038/nature09181
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DOI: https://doi.org/10.1038/nature09181
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