ERNEST WILLI,A.MS BULLEIN OF TlHE AMERICAN.MUSEUM" OF" "NATURAL HISTORY-'. VOLM 95: ARTICLE NEW OK 1950f'~ TESTUDO CUBENSIS AND THE EVOLUTION OF WESTERN HEMISPHERE TORTOISES Page 000_02 is blank TESTUDO CUBENSIS AND THE EVOLUTION OF WESTERN HEMISPHERE TORTOISES ERNEST WILLIAMS Harvard University BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 95 : ARTICLE 1 NEW YORK 1950 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 95, article 1, pages 1-36, text figures 1, 2, plates 1-8, tables 1-3 Issued March 30, 1950 Price: $.75 a copy CONTENTS INTRODUCTION ... ....... .. .. .. 7 Historical Background ....... ........ .. 7 SYSTEMATIC DESCRIPTION .. 9 DISCUSSION .10 Geologic Occurrence and Condition of the Material . 1 Character Analysis. .11 Interrelationships of Western Hemisphere Testudines and the Phylogeny of tTestudo cubensis . 16 Miocene Testudines . 17 Oligocene Ancestors . 19 Eocene Prototestudines .20 Pliocene and Pleistocene Descendants. .21 The Phyletic Position of tTest-udo cubensis .. 23 The Nomenclature of Western Hemisphere Testudines .24~ The Neotropical Tortoises . .. .. .. .. 24~ The Nearctic Tortoises .25 Species and Genera Incertae Sedis .26 CHECI LIST OF NEW WORLD TESTUDINES AND SUPPOSED TESTUDINES 29' SUMMARY ...... .. .... .. .. 32 ACKNOWLEDGMENTS.33 BIBLIOGRAPHY . 34: 99 4-: O Its i. L e a a tv 4- 0 . D eV a CdO ,p O a cdO 0 0 Co)0 bcn .cn J &C O44._t, 96- *> ;-1 0) c :a v 8) co~ c, Co 3e 0 io Q 0) c0 co Co (,rn c O .c -C^o0- 6 INTRODUCTION IN THE MORE THAN 80 years since the descrip- living P. decussata. From Jamaica a single tion by Leidy of testudinate fossil renains partial shell records the existence of Pseu- from Cuba and Sombrero Island, few addi- demys terrapen contemporaneously with cer- tions have been made to the knowledge of tain Jamaican fossil mammals sometimes this West Indian fauna. During this period regarded as older than most other so-called enough material has accumulated at the Pleistocene West Indian mammals, while American Museum of Natural History to from Puerto Rico a great number of frag- demonstrate that the testudinate fauna of the ments give evidence of a large and thick- West Indian Pleistocene was much richer shelled emydine not at present generically than has ever been suspected. From Cuba determinable. Finally from the caves of Mona there is now available, in part from the same Island elements of a turtle have been col- deposits that provided the ground sloths lected which in limbs and cervical vertebrae named by Matthew (1931), not only consider- closely resemble Testudo but from the palate able material of Leidy's tTestudo cubensis and certain features of the shell must belong but a number of partial shells and fragments to an undescribed genus of uncertairn relation- of a tPseudemys evidently related to the still ships. HISTORICAL BACKGROUND Vertebrate fossils were first found in Cuba partly studied by him and partly entrusted in April, 1860, in a thermal spring at the to the American Museum of Natural History Bafnos de Ciego Montero by a student at the for more thorough research in conjunction University of Havana, Jose de Figueroa. He with further exploitation of the localities. brought them to Felipe Poey, who presented In 1911 Barnum Brown in association with them on the fifteenth of September, 1861, de la Torre completed collections at the before the Academy of Sciences at Havana. locality in the Sierra de Jatibonico and also Poey sent several of the fossils to Leidy who obtained a large sample from Ciego Montero. in 1868 described tTestudo cubensis from a In 1918 Brown completed the excavation of single partial pleural' plate found at Ciego the Ciego Montero spring. At both localities Montero. Carlos de la Torre, who succeeded material was gathered not only of t T. cubensis Poey in the chair of zoology at Havana, but also of tPseudemys cf. P. decussata. continued collecting at the Ciego Montero Three other localities in Cuba have yielded spring. He also devoted much time to a new testudinate remains. A fragment of a Pseu- locality in the Sierra de Jatibonico, a fissure demys plastron was found by H. E. Anthony discovered by a refugee Cuban patriot, in 1917 in a cave at Daiquiri, Oriente Prov- Ramon Gonzalez, during the Cuban fight for ince, and fragments of a small specimen of the liberation from Spain. In 1910 de la Torre same genus were collected by Barnum Brown reported some of his findings to the Inter- in the Cueva de los Machos near Cienfuegos, national Geological Congress at Stockholm. Santa Clara Province. In a tar pit near Hato The material collected by de la Torre was Nuevo, Matanzas Province, Roy E. Dicker- son and P. J. Bermudez in 1933 found, be- 1 The terminology here employed endeavors to be sides some bones of domestic animals, ground consistent in distinguishing horny scutes from bony sloth claws, and small rodent bones, a few plates. The following are the corresponding terms for the sets of elements in the carapace: plates and a femur of tT. cubensis. The lat- SCUTES PLATES ter fragments from a source apparently vertebral neural deserving of further investigation were do- costal pleural nated to the American Museum of Natural marginal peripheral nuchal nuchal History by Dickerson. supracaudal pygal Leidy, in the same brief note in which he Plastral nomenclature is already consistent and described tT. cubensis, erected the name unambiguous. tEmys sombrerensis for a partial plastron 7 BULLETIN AMERICAN MUSEIUM OF NATURAL HISTORY VOL. 95 found during excavations for fertilizer on at the moment of publication of Schmidt's tiny Sombrero Island. Leidy was uncertain paper, Anthony was in the process of explor- whether the form was an Emys or a Testudo. ing the caves of Mona. The only fossil verte- The type is now lost, and the description is brate he found, aside from a very few bones not determinable. The specimen was not of a small mammal (tlsolobodon) probably Emys as the genus is now understood. That of human importation, was a new and peculiar it was Pseudemys is not probable either; genus of tortoise. the described depth of the xiphiplastral notch Two years previously from Puerto Rico suggests Testudo as more likely. itself (the exact locality and circumstances In 1920 a third island was added to the list of the find uncertain) still another form was of those on which fossil turtles were to be added to the growing collection at the Ameri- found. In that year H. E. Anthony procured can Museum of Natural History. This form, a shell of Pseudemys terrapen from the breccia the gift of Sefnor Rabell Cabrera, was repre- of the same cave in Jamaica in which he sented by numerous fragments of plastron, found tClidomys, tSpirodontomys, tSpeoxe- pelvis, and of the buttress region of the cara- nus, and tAlterodon (Anthony, 1920a). pace, but was unfortunately so incomplete The abundant mammalian remains in West as to afford no generic characters. Indian caves reported principally by Anthony Of this material only tTestudo cubensis stirred considerable interest at this time. In Leidy is described in the present paper, 1926 K. P. Schmidt, discussing the modern special emphasis being placed on the relation- herpetofauna (in which no turtles are in- ships of that form and on its position in the cluded) of the small island of Mona off phylogeny of Western Hemisphere tortoises. Puerto Rico, suggested that the numerous It is intended to discuss the other forms in limestone caves on that island would prob- succeeding papers. ably afford much of interest. As it happened, SYSTEMATIC DESCRIPTION ORDER TESTUDINATA ADDITIONAL MATERIAL: Fragments of cara- FAMILY TESTUDINIDAE pace, plastron, and limb bones from the SUBFAMILY TESTUDININAE Casimba locality in the collection of the American Museum of Natural History; frag- DEFINITION: Testudinids with never more ments (Moreno collection) from the Casimba than two phalanges per digit on fore or hind locality in the collections of the Museum of foot. Comparative Zo6logy, Harvard College; a GENUS TESTUDO LINNi, 1758 femur and plate fragments from the tar GENOTYPE: Testudo graeca Linn6, 1758. pit in Matanzas Province in the collections NEW RESTRICTED DEFINITION: A genus of of the American Museum of Natural History. Testudininae world-wide in distribution, DIAGNOSIS: A species of 7iestudo distin- with the alveolar surface of the maxillae guished by having the borders of the verte- broad and usually with one to three rather bral and costal scutes raised into sharp- distinct ridges, never with a ridge at the crested ridges on the plates and by having symphysis of the alveolar surfaces of the the humerus markedly compressed in the premaxillae, palatines and vomers forming a plane of the head and with a deep medial deep trough, cervical vertebrae rather slender, pit for the latissimus dorsi. carapace never hinged, centrum of first dorsal ADDITIONAL CHARACTERS: Size comparable vertebra not elongate. to that of Gal6apagos species. Shell rather elongate, thin and weak in central portions tTestudo cubensis Leidy, 1868 of carapace and plastron. Areas underlying Testudo cubensis LEIDY, 1868, Proc. Acad. Nat. the marginal, gular, and anal scutes more or Sci. Philadelphia, vol. 20, p. 179. less thickened. Neural and pleural plates TYPE: Part of a first right pleural plate, probably partly discontinuous with vacuities Academy of Natural Sciences of Philadelphia at the sutural junctions. Nuchal scute ab- No. 8923. sent, the first marginal scutes in broad con- HORIZON: Pleistocene? (see below). tact. Marginal scutes often, and plastral TYPE LOCALITY: The Chapepote spring at scutes mostly, with impressed borders. Pos- Bafios de Ciego Montero, Santa Clara Prov- terior free margin of the carapace strongly ince, Cuba. recurved. Anterior rim moderately recurved. ADDITIONAL LoCALITIES: Casimba de Jati- Free margins markedly thin, dentate.