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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2882, pp. 1-19, figs. 1-14, table 1 June 24, 1987 The Triassic Reptile Prolacerta in Antarctica EDWIN H. COLBERT' ABSTRACT The prolacertid reptile, Prolacerta broomi, hith- skeleton, including hind limbs and some caudal erto known from the Lystrosaurus Zone of the vertebrae of prolacertilian relationships, likewise Middle Beaufort beds ofSouth Africa, is described would seem to represent an individual somewhat from specimens including two skulls with jaws, smaller than the South African species. The pres- other fragments with skulls and jaws, and various ence of Prolacerta broomi-in addition to a ly- postcranial elements, found in the Fremouw For- dekkerinid amphibian Cryobatrachus kitchingi, the mation ofthe Transantarctic Mountains. The Ant- dicynodont reptiles Lystrosaurus murrayi, Lystro- arctic fossils represent individuals generally some- saurus curvatus, Lystrosaurus mccaigi, and Myo- what smaller than those known from South Africa, saurus gracilis, the procolophonid Procolophon with slightly fewer teeth in the upper and lower trigoniceps, and the theriodonts Thrinaxodon lio- jaws, and with other minor differences. These vari- rhinus and Ericiolacerta parva-corroborates the ations are, however, probably attributable to age close relationships between Antarctica and South differences and perhaps geographic separation Africa and their probable connection during Early rather than to any specific distinctions. A partial Triassic time. INTRODUCTION The collection of fossil tetrapods secured were found during the second field season, from the Lower Triassic Fremouw Forma- and were collected in the vicinity ofthe junc- tion ofAntarctica during the austral summers tion between McGregor and Shackleton gla- of 1969-1970 and 1970-1971 contains var- ciers, at about latitude 85°13'S and longitude ious remains representative of the diapsid 174°30'E. The fossils were collected by Dr. reptiles, these being specifically prolacertili- James W. Kitching of the Bernard Price In- ans and thecodonts. All of these specimens stitute for Palaeontological Research, Uni- I Curator Emeritus, The American Museum of Natural History, New York City; Professor Emeritus, Columbia University in the City of New York; Honorary Curator of Vertebrate Paleontology, The Museum of Northern Arizona, Flagstaff. Copyright © American Museum of Natural History 1987 ISSN 0003-0082 / Price $2.25 2 AMERICAN MUSEUM NOVITATES NO. 2882 versity of the Witwatersrand, Johannesburg, are prevalent in the vicinity of McGregor and his associates, namely John Ruben, then and Shackleton glaciers. ofthe University ofCalifornia, Berkeley, and In the previous papers of this series it has Thomas Rich, then ofthe American Museum been shown that the fossil tetrapods from the of Natural History, New York. Fremouw Formation are closely related to At this point I wish to acknowledge advice those occurring in the Lower Triassic Lystro- and comments from Dr. Robert Carroll of saurus Zone, Middle Beaufort beds of the the Redpath Museum, McGill University, Karroo Series in southern Africa. Montreal; from Dr. Peter Galton, ofthe Pea- body Museum, Yale University, New Haven, DESCRIPTIONS OF FOSSILS Connecticut, and the University of Bridge- port, Bridgeport, Connecticut; and from the SYSTEMATICS late Professor Georg Haas, The Hebrew Uni- CLASS REPTILIA versity, Jerusalem, Israel. Consultations with these authorities were most helpful; it must SUBCLASS DIAPSIDA be emphasized, however, that all conclusions ORDER EOSUCHIA reached in this paper are solely the respon- sibility of the author. FAMILY PROLACERTIDAE The drawings were made by Miss Pamela Comments: The family Prolacertidae was estab- Lunge; the photographs by Mr. Mark Mid- lished by Parrington in 1935, in his original de- dleton, both ofthe Museum ofNorthern Ar- scription of Prolacerta broomi, new genus and izona. species. He said about the higher taxonomic re- This work has been supported in part by a lationships of the genus and species: grant from the National Science Foundation, Though most closely allied to the Eosuchia No. DPP 75-23126. Prolacerta cannot, I think, be included among them, since they are the most primitive diapsids and the basal group from which all later, spe- STRATIGRAPHIC RELATIONSHIPS cialized, forms may have evolved. It is excluded from the Squamata by the presence of part of For the benefit of those unfamiliar with the quadratojugal and the fact that the quadrate previous papers in this series describing is fixed. [But the quadrate is not fixed, as was Triassic tetrapods from Antarctica, a few re- shown by Camp in 1945 and by Gow in 1975.] marks will be included here concerning the The proper place seems to be in a new family stratigraphic relationships of the Fremouw of the Thecodontia, for which the name Pro- Formation. This, the lowest unit in the Ant- lacertidae is proposed. (Parrington, 1935: 205) arctic Triassic sequence, overlies the Permian In his paper describing in detail the skull of Buckley Formation, from which it is sepa- Prolacerta, Camp (1945: 95) stated that for Pro- rated by a disconformity, and in turn it is lacerta the "concert of characters seems to place it in the Protorosauria (=Eosuchia), and within overlain by the Falla Formation and above the family Protorosauridae." that the Prebble Formation, both of Triassic The family Prolacertidae was placed within the age. The Prebble Formation is separated by order Eosuchia by Romer in 1956 and 1966, and a disconformity from overlying Jurassic vol- by Piveteau in 1955. It was included within the canic rocks, the Ferrar Dolerite, which may Lacertilia by Tatarinov (in Rozhdestvenskii and also be intruded into the Triassic sediments. Tatarinov) in 1964. In 1956 von Huene included The Fremouw Formation in the Transantarc- Prolacerta in his underorder Prolacertilia, but did tic Mountains may reach thicknesses of 650 not designate a family to contain the genus. m. Kuhn-Schnyder (1962: 132) described a skull of The sediments ofthe Fremouw Formation Macrocnemus and showed its close similarities to are cyclic in nature, ranging from the skull ofProlacerta. He felt, however, that the coarse, con- relationships of the higher taxa of lepidosaurians glomeratic channel deposits at the base ofthe were too uncertain to warrant a definitive position cycle to fine floodplain deposits at the top. for Prolacerta and its relatives. "The relationships The specimens described in this paper were between Eosuchians, Archosaurians and Prolacer- collected from the fine-grained sediments that tilians are by no means clear. Therefore we at pres- 1987 COLBERT: PROLACERTA 3 ent classify only the Prolacertilians with the Lep- temporal arch incomplete, the jugal termi- idosaurians." nating posteriorly in a point that does not Pamela Robinson, in her paper of 1967 on "The reach the quadrate. Quadratojugal persisting, Evolution of the Lacertilia," placed the origin of but very slender. Supratemporal fenestra this order among eosuchian diapsids such as Pro- lacerta. However she did not attempt formal des- rather large. Nasals extensive, overlapped by ignations of the higher taxa of these reptiles. maxillae. Numerous palatal teeth on ptery- In his definitive study ofProlacerta, Gow (1975: goid and palatine. Lower jaw slender, prob- 1 18) established a new order, Parathecodontia, for ably with a ligamentous symphysis. Teeth the family Prolacertidae-probably to be included pleurothecodont. Five premaxillary teeth, within the Lepidosauria. His justification for set- more than 20 maxillary teeth extending to a ting up this new order of reptiles was as follows: position beneath front of orbit; more than 20 "It is thus clear that Prolacerta is representative dentary teeth. Vertebrae amphicoelous. Post- of an as yet poorly known group of early Triassic cranial skeleton slender; bones frequently prearchosaurian thecodonts which in all proba- hollow or internally cancellous. Pectoral gir- bility gave rise to the later Macrocnemus and Tanystropheus. These can then be regarded as a dle with expanded, very thin scapulocora- sterile group which paralleled the Squamata in the coid, heavy clavicle and long interclavicle; loss of the lower temporal arcade." pelvic girdle with broad puboischiatic plate. Although the Prolacertidae probably do repre- Limbs slender; hind limb much longer than sent a group separate from both the eosuchians forelimb. Ankle with large astragalus-calca- and the lacertilians in the strict sense of the two neum. "Hooked" fifth metatarsal, with fifth terms, it seems premature to set up a new reptilian digit offset from the other digits. Typical rep- order for the three genera involved. Since Prola- tilian phalangeal formula. certa seemingly is not directly ancestral to the true lizards, perhaps the best course at the present time is to retain the family Prolacertidae in the Eosu- Prolacerta broomi Parrington chia, realizing that a satisfactory definition of the Eosuchia is not now possible. Prolacerta broomiParrington, 1 935: 197-205, figs. This is substantially the procedure followed by 1-3, pl. 9. Evans (1980). She recognized the subclass Diap- TYPE: A skull in the Museum of Zoology, sida to contain three infraclasses, namely Eosu- Cambridge University, from Harrismith, Or- chia, Archosauria and Squamata, and she placed ange Free State, South Africa. the Prolacertidae along with 14 other families AND Lower Trias- within the Eosuchia.
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