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Sineoamphisbaena Hexatabularis, an Amphisbaenian (Diapsida

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Sineoamphisbaena Hexatabularis, an Amphisbaenian (Diapsida 541 Sineoamphisbaena hexatabularis, an amphisbaenian (Diapsida: Squamata) from the Upper Cretaceous redbeds at Bayan Mandahu (Inner Mongolia, People's Republic of China), and comments on the phylogenetic relationships of the Amphisbaenia Xiao-Chun Wu, Donald B. Brinkman, and Anthony P. Russell Abstract: Sineoamphisbaena hexatabularis Wu et al., 1993 is the earliest known amphisbaenian represented by well-preserved cranial and postcranial material. It reveals a mosaic of generalized lizard-like features and amphisbaenian characters. Most distinctive of the latter are features of cranial consolidation adaptive for a fossorial way of life. Phylogenetic analyses strongly confirm the monophyly of the Amphisbaenia inclusive of S. hexatabularis. The Amphisbaenia is diagnosed by a suite of apomorphic characters. The available evidence suggests a probable Amphisbaenia-Macrocephalosauridae relationship within the Scincomorpha. This is supported primarily by the unique modifications of the palate and temporal region of the skull. It is argued here that the Amphisbaenia evolved in Central Asia during the Cretaceous, in response to the transition from a perennial lacustrine environment to a dry, semiarid eolian environment. The relatively primitive morphology indicates that S. hexatabularis was not permanently subterranean. The further derived modifications of later forms are associated with tunneling in an environment of more compact soils. RCsumC : Le plus ancien amphisbenien connu, reprtsentC par du mattriel crdnien et postcrinien bien prCservC, est Sineoamphisbaena hexatabularis Wu et al., 1993. I1 montre un assemblage de traits generaux appartenant aux lCzards et des caractkres d'amphisbCniens. Ce qui distingue le plus ces derniers, ce sont les particularitCs qui consolident l'adaptation du crine au mode de vie de reptile fouisseur. Les analyses phylogCnCtiques confirment largement la monophylie des Amphisbaenia incluant S. hexatabularis. Les Amphisbaenia sont identifiCs par un ensemble de caractbres apomorphiques. Les donnees disponibles suggbrent un rapprochement probable des Ampbisbaenia-Macrocephalosauridae a I'intCrieur des Scincomorphes. Ce qui est CtayC principalement par les modifications uniques du palatin et de la rtgion temporale For personal use only. du crine. I1 est suggCrC ici que les Amphisbaenia ont CvoluC en Asie centrale durant le CrCtacC, en rkponse a la transition d'un milieu lacustre stable depuis longtemps vers un milieu sec, semiaride et Colien. La morphologie relativement primitive indique que S. hexatabularis ne vivait pas en permanence sous terre. Les modifications dCrivCes ulterieures des formes plus tardives sont associCes au creusement de tunnels dans un milieu constituC de sols plus compacts. [Traduit par la rCdaction] Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by 129.107.48.242 on 04/20/11 Received February 8, 1995. Accepted December 13, 1995. X.-C. Wu.' Vertebrate Morphology Research Group, Department of Biological Sciences, The University of Calgary, 2500 University Drive NW, Calgary, AB T2N 1N4, Canada, and Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, P.O. Box 643, Beijing 100044, People's Republic of China. D.B. Brinkman. Royal Tyrrell Museum of Palaeontology, P.O. Box 7500, Drumheller, AB TOJ OYO, Canada. A.P. Russell. Vertebrate Morphology Research Group, Department of Biological Sciences, The University of Calgary, 2500 University Drive NW, Calgary, AB T2N 1N4, Canada. ' Corresponding author (e-mail: rtmp@dns .magtech.ab. ca) . lan. J. Earth Sci. 33: 541 -577 (1996). 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AOSSaN f 1661 .3!ozoua3 ayl ~IO.IJ I@ alam exs psoj pa~asa~ddlalald .Ie la eyqgsv f1661 ey3!udp~a-yns~oa)e!sv pIiua3 jo -1.~03lsour ayl 'dlsno~naq'ajg jo dem ppossoj e 01 paldepe snoa3qal3 lamq lsoulraddn ayl pue 'u~dspm ego%uo~ 'sawumnbs ayy-mom 'paqpgads 6~3yale smyuaeqsyyduqf jo snoasqala ~addnaql u~oypapodal dpua3a~ slam For personal use only. Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by 129.107.48.242 on 04/20/11 Wu et al. Fig. 1. (A) Regional locality map of the Gobi Basin showing the location of the Bayan Mandahu. (B) Localities of fossil vertebrates near the Bayan Mandahu seasonal herb station. The specimens of Sineoamphisbaena hexatabularis were collected from bedrock near the North Canyon site. Solid lines indicate ephemeral drainage. Modified from Figs. 1 and 3 of Jerzykiewicz et al. (1993). For personal use only. { To Urad Hou Qi amphisbaenian. Its relative primitiveness provides insights nized within the redbeds (Fig. lB), representing a proximal- into the morphology, early history, and evolution of the to-distal depositional gradient from the "Paleo" Lang Shan Amphisbaenia. In the present paper, we first describe the in the south to an extensive sand sea in the north (Eberth osteological anatomy of S. hexatabularis, and then reevalu- 1993). The amphisbaenian specimens studied here were col- ate previous hypotheses of the phylogenetic relationships of lected from zone 3, in which protoceratopsian and ankylo- the group as well as discuss its origin and early adaptation. saurian (Pinacosaurus) dinosaurs are dominant, and dinosaur eggs, lizards, and mammals are common. Zone 3 is inter- Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by 129.107.48.242 on 04/20/11 Geological setting preted as the southern margin of an extensive sand sea and is characterized by a large number of eolian dune deposits, The Bayan Mandahu redbeds are well exposed about 50 km but also contains structureless deposits and rare lacustrine northwest of the Lang Shan mountain range (Fig. 1). The deposits (Eberth 1993). redbeds are essentially flat lying (with a regional dip < 3" W) and outcrop along a 12 km, north-south-trending escarp- Materials and methods ment, about 8 km north and slightly east of the Bayan Mandahu seasonal shepherd station. The entire stratigraphic The two available specimens were both collected in the thickness of the Bayan Mandahu redbeds unit is unknown. North Canyon area (Fig. 1B). The articulated skeleton was Jerzykiewicz et al. (1993) interpreted the redbeds as chrono- preserved in a natural posture and reported as an embryonic stratigraphically equivalent to the Djadokhta Formation of ankylosaur in the popular literature (Cui 1991). This skeleton south-central Mongolia, which is regarded to be Campanian consists of the skull without the mandible, associated first in age (Lillegraven and McKenna 1986; Jerzykiewicz and 19 vertebrae with 11 (right) and 10 (left) ribs, the left Russell 1991). Three coeval sedimentary zones are recog- pectoral girdle with the humerus, the distal portion of the 544 Can. J. Earth Sci. Vol. 33, 1996 Fig. 2. Holotype specimen of Sineoamphisbaena hexatabularis (IVPP V10.593 (= V10593-I, Wu et al. 1993)) in (A) dorsal and (B) ventral views. For personal use only. Can. J. Earth Sci. Downloaded from www.nrcresearchpress.com by 129.107.48.242 on 04/20/11 right humerus and part of the right coracoid, and the placed ventrally. The ramus is missing its retroarticular posterior portion of the interclavicle (Fig. 2). The cranium process, and the ventral edge of its dentary is damaged. is missing the anterior tip of the snout, and its occipital The specimens are housed in the Institute of Vertebrate surface is slightly damaged. The neural arch is incomplete Paleontology and Paleoanthropology (IVPP), Academia and the ribs are not preserved on vertebrae 3 -7. Most of the Sinica, Beijing, People's Republic of China. According to ribs are represented by their proximal portion. The second the regulations of IVPP, different individuals of a species specimen, an isolated skull, was picked up from the surface should not share the same accession number, even if they (Fig. 3). The dermal roof elements of its cranium are heavily have been collected from the same locality. Therefore, the eroded, exposing the brain cavity, exaggerating the size of specimen number of the holotype has been changed to IVPP the external nares, and creating a pair of holes on the fron- V10593 (= V10593-1, Wu et al. 1993) and that of the para- tals. In addition, the cranium is missing its palatal elements type has been changed to IVPP V10612 (= V10.593-2, Wu and occipital condyle. The ventral surface of the braincase is et al. 1993). Specimens of many living and fossil amphis- broken, and its left quadrate has shifted anteromedially. The baenians were examined for comparative purposes and the postdentary portion of the left rarnus of the mandible is dis- generation of the data matrix for the phylogenetic study; Wu et al. these were obtained from Pratt Museum of Arnherst College Fig. 3. Stereophotographs of the skull of the paratype of (ACM); American Museum of Natural History (AMNH); Sineoamphisbaena hexatabularis (IVPP V10612 (= V10593-2, Carnegie Museum of Natural History (CMNH); Museum of Wu et al.
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