Ornithischia is one of the two major groups of dinosaurs, with heterodontosauridae as one of its major clades. Heterodontosauridae is characterized by small, gracile bodies and a problematic phylogenetic position1,2. Recent phylogenetic work indicates that it represents the most basal group of all well-known ornithischians3. Previous heterodontosaurid records are mainly from the Early Jurassic period (205–190 million years ago) of Africa1,3. Here we report a new heterodontosaurid, Tianyulong confuciusi gen. et sp. nov., from the Early Cretaceous period (144–99 million years ago) of western Liaoning Province, China. Tianyulong extends the geographical distribution of heterodontosaurids to Asia and confirms the clade’s previously questionable temporal range extension into the Early Cretaceous period. More surprisingly, Tianyulong bears long, singular and unbranched filamentous integumentary (outer skin) structures. This represents the first confirmed report, to our knowledge, of filamentous integumentary structures in an ornithischian dinosaur.
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Norman, D. B., Sues, H.-D., Witmer, L. M. & Coria, R. A. in The Dinosauria 2nd edn (eds Weishampel, D. B., Dodson, P. and Osmólska, H.) 393–412 (Univ. California Press, 2004)
Xu, X., Forster, C. A., Clark, J. M. & Mo, J. A basal ceratopsian with transitional features from the Late Jurassic of northwestern China. Proc. R. Soc. Lond. B 273, 2135–2140 (2006)
Butler, R. J., Upchurch, P. & Norman, D. B. The phylogeny of the ornithischian dinosaurs. J. Syst. Palaeontol. 6, 1–40 (2008)
Zhou, Z., Barrett, P. M. & Hilton, J. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421, 807–814 (2003)
Ji, Q. et al. Mesozoic Jehol Biota of Western Liaoning, China (Geological Publishing House, 2004)
Weishampel, D. B. & Witmer, L. M. in The Dinosauria 1st edn (eds Weishampel, D. B., Dodson, P. & Osmólska, H.) 486–497 (Univ. California Press, 1990)
Lingham-Soliar, T., Feduccia, A. & Wang, X. A new Chinese specimen indicates that ‘protofeathers’ in the Early Cretaceous theropod dinosaur Sinosauropteryx are degraded collagen fibres. Proc. R. Soc. Lond. B 274, 1823–1829 (2007)
Feduccia, A., Lingham-Soliar, T. & Hinchliffe, J. R. Do feathered dinosaurs exist? Testing the hypothesis on neontological and paleontological evidence. J. Morphol. 266, 125–166 (2005)
Mayr, G., Peters, D. S., Plodowski, G. & Vogel, O. Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus. Naturwissenschaften 89, 361–365 (2002)
Zhang, F., Zhou, Z. & Dyke, G. J. Feathers and ‘feather-like’ integumentary structures in Liaoning birds and dinosaurs. Geol. J. 41, 395–404 (2006)
Xu, X. Feathered dinosaurs from China and the evolution of major avian characters. Integ. Zool. 1, 4–11 (2006)
Zhang, F. et al. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature 455, 1105–1108 (2008)
Zhang, F., Zhou, Z. & Benton, M. J. A primitive confuciusornithid bird from China and its implications for early avian flight. Sci. China Ser. D 51, 625–639 (2008)
Zhang, F. & Zhou, Z. A primitive enantiornithine bird and the origin of feathers. Science 290, 1955–1960 (2000)
Zheng, X., Zhang, Z. & Hou, L. A new enantiornitine bird with four long rectrices from the Early Cretaceous of northern Hebei, China. Acta Geol. Sin.-Engl. 81, 703–708 (2007)
Currie, P. J. & Chen, P. J. Anatomy of Sinosauropteryx prima from Liaoning, northeastern China. Can. J. Earth Sci. 38, 1705–1727 (2001)
Xu, X., Zhou, Z.-H. & Prum, R. O. Branched integumental structures in Sinornithosaurus and the origin of feathers. Nature 410, 200–204 (2001)
Butler, R. J., Porro, L. B. & Norman, D. B. A juvenile skull of the primitive ornithischian dinosaur Heterodontosaurus tucki from the ‘Stormberg’ of southern Africa. J. Vertebr. Paleontol. 28, 702–711 (2008)
Naish, D. & Martill, D. M. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: Ornithischia. J. Geol. Soc. Lond. 165, 613–623 (2008)
Sánchez-Hernández, B., Benton, M. J. & Naish, D. Dinosaurs and other fossil vertebrates from the Late Jurassic and Early Cretaceous of the Galve area, NE Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 249, 180–215 (2007)
Norman, D. B. & Barrett, P. M. Ornithischian dinosaurs from the Lower Cretaceous (Berriasian) of England. Spec. Pap. Palaeontol. 68, 161–189 (2002)
Sharov, A. G. An unusual reptile from the Lower Triassic of Fergana. Paleontol. J. 1970, 112–116 (1970)
Voigt, S. et al. Feather-like development of Triassic diapsid skin appendages. Naturwissenschaften 96, 81–86 (2009)
Wang, X., Zhou, Z., Zhang, F. & Xu, X. A nearly completely articulated rhamphorhynchoid pterosaur with exceptionally well-preserved wing membranes and ‘hairs’ from Inner Mongolia, northeast China. Chin. Sci. Bull. 47, 226–230 (2002)
Ji, Q. & Yuan, C.-X. Discovery of two kinds of protofeathered pterosaurs in the Mesozoic Daohugou Biota in the Ningcheng region and its stratigraphic and biologic significances. Geol. Rev. 48, 221–224 (2002)
Reisz, R. R. & Sues, H. D. The ‘feathers’ of Longisquama. Nature 408, 428 (2000)
Jones, T. D. et al. Nonavian feathers in a late Triassic archosaur. Science 288, 2202–2205 (2000)
Prum, R. O. & Brush, A. H. The evolutionary origin and diversification of feathers. Q. Rev. Biol. 77, 261–295 (2002)
Yu, M., Wu, P., Widelitz, R. B. & Chuong, C.-M. The morphogenesis of feathers. Nature 420, 308–312 (2002)
Goloboff, P. A., Farris, J. S. & Nixon, K. C. TNT, a free program for phylogenetic analysis. Cladistics 24, 774–786 (2008)
We thank P.-J. Chen for arranging the study, Y.-Q. Zhang for preparing the specimen, J. D. Harris for improving the manuscript, and L.-D. Xing for drawing Fig. 1d. H.-L.Y. was supported by grants from the National Natural Science Foundation of China (40672007), the 973 Project and the Basic Outlay of Scientific Research Work from the Ministry of Science and Technology, and the Hundred Talents Project of the Ministry of Land and Resources of China.
Author Contributions X.-T.Z. designed the project. H.-L.Y., X.X. and Z.-M.D. performed the research. H.-L.Y. wrote the manuscript.
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