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Zangerlia Ukhaachelys, New Species, a Nanhsiungchelyid Turtle from the Late Cretaceous of Ukhaa Tolgod, Mongolia

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Zangerlia Ukhaachelys, New Species, a Nanhsiungchelyid Turtle from the Late Cretaceous of Ukhaa Tolgod, Mongolia PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3481, 19 pp., 6 ®gures July 25, 2005 Zangerlia ukhaachelys, New Species, a Nanhsiungchelyid Turtle from the Late Cretaceous of Ukhaa Tolgod, Mongolia WALTER G. JOYCE1 AND MARK A. NORELL2 ABSTRACT The Late Cretaceous continental deposits of Ukhaa Tolgod, Mongolia, have yielded remains of a new NANHSIUNGCHELYID turtle, Zangerlia ukhaachelys, n.sp. This taxon is based on a single individual that consists of a partial cranium, representatives of all peripherals, an almost complete plastron, and limb fragments. Zangerlia ukhaachelys is diagnosed as a new taxon by the presence of an anteromedial process of the hyoplastron that reduces the typical contact of the entoplastron with the epiplastron. Phylogenetic analysis ®rmly places Zangerlia ukhaa- chelys as sister to Zangerlia testudinimorpha and Zangerlia neimongolensis within NANHSI- UNGCHELYIDAE and con®rms the close phylogenetic relationships between Nanhsiungchelys wuchingensis and Anomalochelys angulata and among all North American representatives of Basilemys. In addition, there is modest support that all Asian representatives of NANHSI- UNGCHELYIDAE form a monophyletic clade, which is primarily diagnosed by a deep, triangular nuchal notch. From a biogeographic standpoint, it is evident that the Late Cretaceous faunas of Asia and North America are closely related; however, phylogenetic considerations dem- onstrate that faunal exchange was limited for the NANHSIUNGCHELYIDAE. INTRODUCTION the Late Cretaceous of North America. Asian Central Asian continental deposits have faunas are characterized by representatives of yielded a diverse fauna of Late Cretaceous the ADOCIDAE,CARETTOCHELYIDAE, Lindhol- turtles that are closely related to those from memydidae, NANHSIUNGCHELYIDAE, Macro- 1 Department of Geology and Geophysics, Yale University, 210 Whitney Avenue, New Haven, CT 06511 (walter. [email protected]). 2 Division of Paleontology, American Museum of Natural History ([email protected]). Copyright q American Museum of Natural History 2005 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3481 baenidae, Mongolochelyidae, and TRIONY- phylogenetically de®ned clade names as pre- CHIDAE and contain both aquatic and terres- sented by Joyce et al. (2004) and herein (see trial forms (Sukhanov, 2000). Together with below). These names are distinguished from faunas from the Late Cretaceous of North traditional rank-based taxonomic names by America, these groups are of special impor- the small capitals type style throughout the tance to the systematics of turtles, as they text. may have given rise to many living crypto- diran groups. As basal relationships of crown MATERIALS CRYPTODIRA are still under debate, a better Several turtle specimens were used for understanding of the morphology of these comparative purposes and were integrated turtles may be crucial to resolving patterns into a phylogenetic analysis. These include: of CRYPTODIRE evolution. Adocus (orig. Emys) beatus (Leidy, 1865), as Late Cretaceous deposits at Ukhaa Tolgod described by Marsh (1890), Hay (1908), have produced thousands of vertebrate fossils White (1972), and personal observation (Dashzeveg et al., 1995), many of which (WGJ) of YPM 782, holotype of A. puncta- have been exquisitely preserved. Interesting- tus Marsh, 1890; Adocus sp., as described by ly, despite the great abundance of fossil Meylan and Gaffney (1989); Zangerlia tes- mammals, lizards, and dinosaurs, only a few tudinimorpha Mlynarski, 1972, as described turtles have been discovered to date. During by Mlynarski (1972); Zangerlia neimongo- the 1993 joint expedition of the Mongolian lensis Brinkman and Peng, 1996, as de- Academy of Sciences and the American Mu- scribed by Brinkman and Peng (1996) and seum of Natural History, a partial skeleton of personal observation (WGJ) of casts of IVPP a fossil turtle was uncovered from this lo- 020788±7, holotype of Z. neimongolensis; cality and preliminarily identi®ed as Basile- ``Basilemys'' orientalis Sukhanov and Nar- mys? (Dashzeveg et al., 1995). The purpose mandakh, 1977, as described by Sukhanov of this paper is to formally describe this and Narmandakh (1977) and coded by Hir- specimen as a new species of NANHSI- ayama et al. (2001); Anomalochelys angulata UNGCHELYIDAE and to explore phylogenetic Hirayama et al., 2001, as described by Hir- relationships within this clade of turtles. ayama et al. (2001); Nanhsiungchelys wuch- INSTITUTIONAL ABBREVIATIONS: IGM, In- ingensis Yeh, 1966, as described by Yeh stitute of Geology, Mongolia, Ulannbaatar (1966) and coded by Hirayama et al. (2001); Mongolia; IVPP, Institute of Vertebrate Pa- Basilemys (orig. Compsemys) variolosa leontology and Paleoanthropology, Beijing, Cope, 1876, carapace and plastron as de- People's Republic of China; YPM, Yale Pea- scribed and depicted by Langston (1956), body Museum, New Haven, CT. cranial and other nonshell characters as TERMINOLOGY: Anatomical terms of the scored by Hirayama et al. (2001); Basilemys cranium follow those summarized by Gaff- nobilis Hay, 1911, carapace and plastron as ney (1972) and those of the shell as recom- described and depicted by Langston (1956), mended by Zangerl (1969). However, the an- postcranial characters as scored by Hirayama terior two pairs of plastral scutes are termed et al. (2001); Basilemys sinuosa Riggs, 1906, ``gulars'' and ``extragulars'' to clearly distin- as described and depicted by Riggs (1906); guish them from the similarly situated, but Basilemys praeclara Hay, 1911, as described ostensibly nonhomologous, ``gulars'' and by Brinkman and Nicholls (1993); and Bas- ``intergulars'' of other turtles (Hutchison and ilemys sp., as described by Brinkman (1998). Bramble, 1981). Because the generic assign- ment of most NANHSIUNGCHELYIDS has varied METHODS substantially over the last two decades (e.g., Sukhanov and Narmandakh, 1977; Meylan The phylogenetic review of NANHSI- and Gaffney, 1989; Brinkman and Peng, UNGCHELYIDAE developed by Hirayama et al. 1996; Sukhanov, 2000; Hirayama et al., (2001) is the basis for this analysis, but the 2001), we use the assignments used by Hir- character matrix was subjected to minor ayama et al. (2001). Wherever possible, taxa changes. In particular, additional states were that are more inclusive are referred to with added to characters 18 and 24 (17 and 20 of 2005 JOYCE AND NORELL: LATE CRETACEOUS TURTLE 3 Hirayama et al., 2001), and characters 25, 26, marily due to signi®cant amounts of missing and 32 of Hirayama et al. (2001) were split data to the cranial region of most currently into separate characters (characters 26±29, known NANHSIUNGCHELYIDS. The data matrix 33, and 35) to encompass greater morpho- was assembled using McClade 3.08 (Maddi- logical variation. In contrast, character 16 is son and Maddison, 1999) and analyzed using a hybrid of characters 22 and 24 of Hirayama PAUP 4.0b10 (Swofford, 2002). Four phylo- et al. (2001), as the original character de®ni- genetic analyses were performed that differ in tions are redundant. A list of modi®ed char- their inclusion of the geographical character acter de®nitions is provided in appendix 1. and in the ordering of the four multistate char- Four new characters were added to the acters. Characters were considered reversible analysis (6, 17, 35, 40) and two were re- and of equal weight in all analyses. Under moved from the analysis. Character 8 of Hir- parsimony settings, branches were set to be ayama et al. (2001) refers to the amount of collapsed if their minimum length was zero. ossi®cation that the canal for the carotid ar- Bootstrap values were calculated using PAUP tery exhibits posterior to the junction with 4.0b10. See appendices 1 and 2 for a com- the palatine artery. Because of the poor pres- plete list of characters used and the taxonomic ervation of the NANHSIUNGCHELYID skull ma- distribution of character states. terial, we are not con®dent in scoring the presence or absence of this character from PHYLOGENETIC NOMENCLATURE the literature. Omission of this character for- NANHSIUNGCHELYIDAE, Converted tunately has no impact on the analysis, be- Clade Name cause the derived condition is purported as present only in Nanhsiungchelys wuchingen- DEFINITION:``NANHSIUNGCHELYIDAE'' re- sis (Hirayama et al., 2001). fers to the most inclusive clade containing According to character 16 of Hirayama et Nanhsiungchelys wuchingensis Yeh, 1966, al. (2001), a unique sculpturing of the shell but not Adocus (orig. Emys) beatus (Leidy, unites all members of the NANHSIUNGCHELYI- 1865) or any species of Recent turtle. DAE into a monophyletic group. Indeed, the DISCUSSION: The name Nanhsiungchelyi- shells of all ingroup turtles are characterized dae was originally coined by Yeh (1966), but by sculpturing, but the morphology of these he only referred Nanhsiungchelys wuchin- sculptures ranges from ®ne grooves and gensis to its content. It is current taxonomic crenulations (e.g., Anomalochelys angulata) practice to assign all turtles to the taxon to uneven pits and pock-marks (e.g., Basile- Nanhsiungchelyidae that are hypothesized to mys variolosa, Zangerlia testudinimorpha), be more closely related to Nanhsiungchelys making it dif®cult to objectively compare or wuchingensis than any species of Adocus homologize them. Given that the character is (e.g., Brinkman and Peng, 1996; Sukhanov, uninformative as currently presented, and 2000; Hirayama et al., 2001). We ®x this re- that the outgroup Adocus sp. is also charac-
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