Abstract
A new alvarezsaurid theropod, Xixianykus zhangi gen. et sp. nov., is described based on a partial postcranial skeleton collected from the Upper Cretaceous Majiacun Formation of Xixia County, Henan Province. The new taxon can be diagnosed by the following autapomorphies: sacral rib-transverse process complexes and zygapophyses fused to form separate anterior and posterior laminae; distinct fossa dorsal to antitrochanter on lateral surface of ilium; short ridge along posterior surface of pubic shaft near proximal end; distinct depression on lateral surface of ischium near proximal end; sharp groove along posterior surface of ischium; distal end of femur with transversely narrow ectocondylar tuber that extends considerable distance proximally as sharp ridge; transversely narrow tibial cnemial crest with sharp, ridgelike distal half; lateral margin of tibiotarsus forms step near distal end; fibula with substantial extension of proximal articular surface onto posterior face of posteriorly curving shaft; distal tarsals and metatarsals co-ossified to form tarsometatarsus; and sharp flange along anteromedial margin of metatarsal IV near proximal end. Cladistic analysis places this taxon as a basal parvicursorine within the Alvarezsauridae, a position consistent with the presence of several incipiently developed parvicursorine features in this taxon and also with its relatively early geological age. A brief analysis of vertebral functional morphology, together with data from the hindlimb, suggests that parvicursorines represent extreme cursors among non-avian dinosaurs.References
Aarde, R.J.v. (1994) Aardvark. In: McDonald, D.W. (Ed.), The Encyclopedia of Mammals, Second Edition. Oxford University Press, London, 466–467.
Alifanov, V.R. & Barsbold, R. (2009) Ceratonykus oculatus gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontological Journal, 43, 94–106.
Bonaparte, J.F. (1991) Los vertebrados fósiles de la Formación Río Colorado de Neuquén y cercanias, Cretácico Superior, Argentina. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Paleontología, 4(3), 17–123.
Carrano, M.T. (1999) What, if anything, is a cursor? Categories versus continua for determining locomotor habit in mammals and dinosaurs. Journal of Zoology, 247(1), 29–42.
Chiappe, L.M., Norell, M.A. & Clark, J.M. (1996) Phylogenetic position of Mononykus (Aves: Alvarezsauridae) from the Late Cretaceous of the Gobi Desert. Memoirs of the Queensland Museum, 39, 557–582.
Chiappe, L.M., Norell, M.A. & Clark, J.M. (1998) The skull of a relative of the stem-group bird Mononykus. Nature, 392, 275–282.
Chiappe, L.M., Norell, M.A. & Clark, J.M. (2002) The Cretaceous, short-armed Alvarezsauridae: Mononykus and its kin. In: Chiappe, L.M. & Witmer, L.M. (Eds.), Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press, Berkeley, 87–120.
Christiansen, P. & Bonde, N. (2002) Limb proportions and avian terrestrial locomotion. Journal of Ornithology, 143, 356–371.
Coombs, W.P.J. (1978) Theoretical aspects of cursorial adaptations in dinosaurs. Quarterly Review of Biology, 53, 393–418.
Currie, P.J. & Zhao, X.-J. (1993) A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences, 30(10&11), 2037–2081.
Dickman, C.R. (1994) Anteaters. In: McDonald, D.W. (Ed.), The Encyclopedia of Mammals, Second Edition. Oxford University Press, London, 772–775.
Gatesy, S.M. & Middleton, K.M. (1997) Bipedalism, flight, and the evolution of theropod locomotor diversity. Journal of Vertebrate Paleontology, 17, 308–329.
Goloboff, P.A. (1993) NONA (ver 2.0). S. M. de Tucuman, Argentina.
Hildebrand, M. & Goslow, G. (2001) Analysis of Vertebrate Structure (fifth edition). Wiley, New York, pp.
Holtz, T.R. (1994) The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology, 14, 480–519.
Hutchinson, J.R. & Allen, V. (2009) The evolutionary continuum of limb function from early theropods to birds. Naturwissenschaften, 96, 423–448.
Hutchinson, J.R. & Chiappe, L.M. (1998) The first known alvarezsaurid (Theropoda: Aves) from North America. Journal of Vertebrate Paleontology, 18(3), 447–450.
Irmis, R.B. (2007) Axial skeletal ontogeny in the Parasuchia (Archosauria: Pseudosuchia) and its implications for ontogenetic determination in archosaurs. Journal of Vertebrate Paleontology, 27(2), 350–361.
Karhu, A.A. & Rautian, A.S. (1996) A new family of Maniraptora (Dinosauria: Saurischia) from the Late Cretaceous of Mongolia. Paleontological Journal, 30(5), 583–592.
Kessler, E., Grigorescu, D. & Csiki, Z. (2005) Elopteryx revisited – a new bird-like specimen from the Maastrichtian of the Hateg Basin (Romania). Acta Palaeontologica Romaniae 5, 249–258.
Koehler, C.E. & Richardson, P.R.K. (1990) Proteles cristatus. Mammalian Species, 363, 1–6.
Longrich, N.R. & Currie, P.J. (2009) Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research, 30, 239–252.
Marsh, O.C. (1881) Principal characters of American Jurassic dinosaurs. Part IV. American Journal of Science (series 3), 21, 417–423
Nixon, K.C. (1999) Winclada (BETA) ver. 0.9.9. Ithaca, New York.
Norell, M.A., Clark, J.M. & Makovicky, P.J. (2001) Phylogenetic relationships among coelurosaurian dinosaurs. In: Gauthier, J. & Gall, L.F. (Eds.), New Perspectives on the Origin and Evolution of Birds. Yale University Press, New Haven, 49–67.
Novas, F.E. (1996) Alvarezsauridae, Cretaceous maniraptorans from Patagonia and Mongolia. Memoirs of the Queensland Museum, 39, 675–702.
Novas, F.E. (1997) Anatomy of Patagonykus puertai (Theropoda, Maniraptora, Alvarezsauridae) from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology, 17(1), 137–166.
Novas, F.E. & Pol, D. (2002) Alvarezsaurid relationships reconsidered. In: Chiappe, L.M. & Witmer, L.M. (Eds.), Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press, Berkeley, 87–120.
Ostrom, J.H. (1969) Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History, Yale University, 30, 1–165.
Owen, R. (1842) Report on British fossil reptiles. Report of the British Association for the Advancement of Science, 11 (1841), 60–294.
Perle, A., Chiappe, L.M., Barsbold, R., Clark, J.M. & Norell, M.A. (1994) Skeletal morphology of Mononykus olecranus (Theropoda: Avialae) from the Late Cretaceous of Mongolia. American Museum Novitates, 3105, 1–29.
Perle, A., Norell, M.A., Chiappe, L.M. & Clark, J.M. (1993) Flightless bird from the Cretaceous of Mongolia. Nature, 362, 623–626.
Richardson, P.K.R. & Bearder, S.K. (1994) The hyena family. In: McDonald, D.W. (Ed.), The Encyclopedia of Mammals, Second Edition. Oxford University Press, London, 154–159.
Senter, P. (2005) Function in the stunted forelimbs of Mononykus olecranus (Theropoda), a dinosaurian anteater. Paleobiology, 31, 373–381.
Sereno, P.C. (2001) Alvarezsaurids: birds or ornithomimosaurs? In: Gauthier, J. & Gall, L.F. (Eds.), New Perspectives on the Origin and Evolution of Birds. Yale University Press, New Haven, 69–98.
Sereno, P.C. (1999) The evolution of dinosaurs. Science, 284, 2137–2147.
Snively, E. & Russell, A.P. (2003) Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function. Journal of Morphology, 255, 215–227.
Soshani, J., Goldman, C.A. & Thewissen, J.G.M. (1990) Orycteropus afer. Mammalian Species, 300, 1–8.
Suzuki, S., Chiappe, L.M., Dyke, G.J., Watabe, M., Barsbold, R. & Tsogtbaatar, K. (2002) A new specimen of Shuvuuia deserti Chiappe et al., 1998 from the Mongolian Late Cretaceous with a discussion of the relationships of alvarezsaurids to other theropod dinosaurs. Contributions in Science, 494, 1–18.
Turner, A.H., Nesbitt, S.J. & Norell, M.A. (2009) A large alvarezsaurid from the Late Cretaceous of Mongolia. American Museum Novitates, 3648, 1–14.
Turner, A.H., Pol, D., Clarke, J.A., Erickson, G.M. & Norell, M.A. (2007) A basal dromaeosaurid and size evolution preceding avian flight. Science, 317, 1378–1381.
Wang, D.Y., Feng, J.C., Zhu, S.G., Wu, M., Fu, G.H., He, P., Ji, G.C., Pang, F.J., Li, G.W., Li, B.X., Li, J.K., Wang, B.X., Zhang, G.J., Qing, Z. & Guo, G.L. (2008) Dinosaur Eggs and Skeletons from Henan Province in China. Geological Publishing House, Beijing, 320 pp.
Xu, X. (2002) Deinonychosaurian fossils from the Jehol Group of western Liaoning and the coelurosaurian evolution. PhD thesis, Chinese Academy of Sciences, Beijing.
Xu, X., Wang, K.-B., Zhao, X.-J. & Li, D.-J. (in press) First ceratopsid dinosaur from China and its biogeographical implications. Chinese Science Bulletin.
Xu, X., Zhao, Q., Norell, M.A., Sullivan, C., Hone, D.W.E., Erickson, G.M., Wang, X.-L., Han, F.-L. & Guo, Y. (2009) A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin. Chinese Science Bulletin, 54, 430–435.