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Norntatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NorntatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2941, 60 pp., 22 figs., 3 tables June 26, 1989 The Skeletal Morphology of the Cretaceous Cryptodiran Turtle, Adocus, and the Relationships of the Trionychoidea PETER A. MEYLAN1 AND EUGENE S. GAFFNEY2 CONTENTS Abstract .................... ............................................ 2 Introduction .................................................................. 2 Materials and Methods .................................................................. 4 Materials ...................................................... 4 Methods ................................I.....S5 Abbreviations ................. .................... 6 Descriptions ofAdocus with Comparisons to Other Members of the Trionychoidea ..... ....... 6 Skull ................... ............................................. 6 Skull Roof ................................................................. 6 Palatal Elements ................................................................. 14 Palatoquadrate and Braincase ........................................................ 17 Canalis Carotici Internus and Related Structures ........... ............................ 19 Lower Jaw ................................................................... 21 Carapace .......................... ....................................... 24 Carapace Form ................................................................. 24 Nuchal .................................................................. 24 Neurals .................................................................. 24 Costals .................................................................. 27 Peripherals .................................................................. 27 Suprapygals .................................................................. 28 Carapacial Scutes ................................................................. 29 Internal Shell Morphology ............................................................ 29 Plastron .................................................................. 30 Plastral Bones ................................................................. 30 Plastral Scales ................................................................. 32 Inframarginal Scales ................................................................. 35 The Cervical Vertebrae ................................................................ 38 Pectoral Girdle and Forelimb ........................................................... 39 Pelvic Girdle .................................................................. 41 'Research Associate, Department of Vertebrate Paleontology, American Museum of Natural History. 2 Curator, Department of Vertebrate Paleontology, American Museum of Natural History. Copyright © American Museum of Natural History 1989 ISSN 0003-0082 / Price $5.50 2 AMERICAN MUSEUM NOVITATES NO. 2941 Discussion ............................................................................. 43 The Trionychoidea .................................................................... 43 The Trionychoidae .................................................................... 48 The Adocidae ........................................................................ 49 The Nanhsiungchelyidae, Peltochelys, and the Trionychia ................................. 50 The Nanhsiungchelyidae ............................................................... 51 Peltochelys and the Trionychia ......................................................... 52 The Trionychia ....................................................................... 52 The Kinosternoidae ................................................................... 52 The Dermatemydidae, Agomphus, and the Kinosternia .................................... 54 The Dermatemydidae ................................................................. 54 Agomphus and the Kinostermia ......................................................... 55 The Kinosternia ...................................................................... 55 The Kinosternidae .................................................................... 56 Acknowledgments. 56 References ............................................................................. 56 ABSTRACT The osteology of the Cretaceous turtle, Adocus, and the sister-group ofthose four taxa, the Adoci- is described, based on a nearly complete specimen dae. The epifamily Kinosternoidae includes the from the Hell Creek Formation of Fallon County, genus Hoplochelys as the sister-group to a mono- Montana. An analysis of this morphology, com- phyletic Kinosternidae; the possible sister-group bined with comparisons with all other fossil and to those two taxa, Agomphus; the sister-group to living trionychoids, provides the basis for a phy- those three taxa, a restricted Dermatemydidae (in- logenetic analysis of the superfamily Trionychoi- cluding only Baptemys and Dermatemys); and the dea, which consists of two large, monophyletic sister-group to those four taxa, Emarginachelys. groups, the epifamilies Trionychoidae and Kino- The inclusion of fossils in this study results in a sternoidae. The epifamily Trionychoidae includes hypothesis for the relationships among the living the sister families Trionychidae and Carettoche- families that is fundamentally different from that lyidae; their sister-group, Peltochelys; the sister- based on a previous study of the living families group ofthose three taxa, the Nanhsiungchelyidae; alone. INTRODUCTION During his short career, George Baur (1859- dae (his Trionychia) was "not an original, but 1898) proposed several hypotheses for rela- a highly specialized group" and that it was tionships among turtles that were novel for related via the Carettochelyidae to stauro- the day but have proved over the years to be typine and kinosternine kinostemids and well supported. In contrast to Cope (1871), dermatemydids (Baur, 1891 c). Thus he pre- Dollo (1886), Boulenger (1889), and Lydek- dicted the existence of the Trionychoidea of ker (1889), he considered Dermochelys to be Gaffney (1975, 1984), a superfamily whose a derived chelonioid (Baur, 1889a) and not recognition is supported by the work of Al- the sister-group to all living turtles (i.e., the brecht (1967, 1976), McDowell (1961), and Atheca). He challenged the ideas of Owen Meylan (1987). (1881) and Boulenger (1887b) and suggested The subject of this paper is the relation- that Meiolania was a cryptodiran turtle (Baur, ships of the members of the Trionychoidea 1889b). And he suggested that Glyptops based primarily on the study of a remarkable should be placed in the family Pleurosterni- new fossil of Adocus. Although membership dae, close to the Baenidae (Baur, 1891a, of living families in this superfamily is suf- 1891 b). All of these ideas have recently been ficiently established, there are two funda- supported by attempts to develop complete mentally different hypotheses for interrela- phylogenetic hypotheses for turtles (Gaffney tionships among them (Gaffney, 1975, 1984; 1972, 1975, 1984; Gaffney and Meylan, Meylan, 1987) and the inclusion of extinct 1988). Baur also recognized that Trionychi- forms has not been completely explored 1 989 MEYLAN AND GAFFNEY: ADOCUS 3 (Hutchison and Bramble, 1981). A nearly complete and well-preserved specimen ofthe genus Adocus provides critical new data and the incentive for a detailed review of the re- lationships of fossil and living Trionychoi- dea. The specimen described below (CCM 60- 15) was made available through the generos- ity ofMarshall Lambert ofthe Carter County Museum, Ekalaka, Montana. It was located and collected by his son, Brice, during the summer of 1961 from the Late Cretaceous (Lancian), Hell Creek Formation in the NW 1/4 ofSec 32 T6N R60E, Fallon County, Mon- tana (figs. 1, 2). Details of the shell morphology of this 00. specimen support its assignment to the genus Adocus of Cope (1868). The type species of Cope's Adocus is Emys beatus Leidy, 1865, based on fragmentary remains from the Cre- taceous Greensands of New Jersey. Cope (1868) gave this species a new generic name, Adocus, and suggested that it resembled Stau- rotypus and Dermatemys. He later (Cope, 1870) coined the family name Adocidae for this new genus. He included the genera Zy- goramma and Homorophus in this family, both ofwhich appear to be synonyms ofAdo- cus. Wieland (1904) provided the first thorough Fig. 1. Collecting the skeleton of Adocus sp. description of the shell of Adocus based on (Carter County Museum 60-15) in the Hell Creek Marsh's (1890) Adocus punctatus. Wieland Formation, Fallon County, Montana, 1961. Above, also recognized the affinity ofAdocus to Der- Brice Lambert exposing carapace; below, dorsal matemys, Staurotypus, and Claudius. Hay view of carapace in situ showing that the skeleton (1908a) first referred Adocus to the Derma- was found dorsal side up. (Photographs courtesy temydidae, and considered it the least mod- of Marshall Lambert, Carter County Museum) ified member of the family. Gilmore (1919), who described five new species ofAdocus from the Cretaceous-Tertiary boundary of New recognized Cope's Adocidae. Mlynarski Mexico, considered Adocus, and also Hop- (1976) included Adocus in the family Der- lochelys, to be representatives of the Der- matemydidae but recognized the subfamily matemydidae. White (1972) reviewed the Adocinae for Adocus, Basilemys, Peishan- New Jersey Cretaceous Greensand
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