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A Diapsid Reptile from the Pennsylvanian of Kansas

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A Diapsid Reptile from the Pennsylvanian of Kansas A DIAPSID REPTILE FROM THE PENNSYLVANIAN OF KANSAS ROBERT R. REISZ Department of Biology University of Toronto Erindale Campus Mississauga, Ontario Canada L5L 1C6 SPECIAL PUBLICATION OF THE MUSEUM OF NATURAL HISTORY, UNIVERSITY OF KANSAS NUMBER 7 1981 1^\ EmstWIayrLfcrary . / UNIVERSIT^BI^'XANSAS PUBLICATIONS ^ <) I ' ( MUSEUM OF NATURAL HISTORY Copies of publications may be obtained from the Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. A list of available Special Publications is provided on the inside back cover. University of Kansas Museum of Natural History Special Publication No. 7 February 18, 1981 A Diapsid Reptile From the Pennsylvanian of Kansas BY Robert R. Reisz Department of Biology University of Toronto Erindale Campus Mississauga, Ontario Canada L5L 1C6 University ok Kansas Lawrence 1981 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY Editor: Joseph T. Collins Editors for this issue: E. O. Wilev and Larry D. Martin h/[r'7 Special Publication No. 7 LIBRARY pp. 1-74; 26 figures 1 table Published February 18, 1981 II II o Q ?nnR - RD I U SiTY Copyrighted 1981 BY Museum of Natural History University of Kansas Lawrence, Kansas 66045 U.S.A. Printed by University of Kansas Printing Service Lawrence, Kansas ISBN: 0-S933S-011-3 This is the first of a series of Special Publications honoring the contributions of Dr. Theodore H. Eaton to the fields of vertebrate paleontology and zoology. The series will center around the Pennsyl\anian fauna of eastern Kansas, and will include papers on fishes, amphibians and reptiles, as befits the wide-ranging interests Dr. Eaton has shown in his research. E. O. Wiley Labry D. Martin 15 June 1980 Lawrence, Kansas CONTENTS INTRODUCTION - 1 ACKNOWLEDGEMENTTS - 2 DESCRIPTn'E AXD SYSTEMATIC HiSTORY 2 SYSTEMATIC DESCRIPTION 4 Family Petrolacosauridae Peabody, 1952 4 Genus Petrolacosauriis Lane, 1945 4 Pefrolacosmirus kansensis Lane, 1945 4 OSTEOLOGY - - 5 Skull 5 Dermal Bones of the Skull Roof 9 Dermal Bones of the Palate 21 Ossifications of the Palatoquadrate Cartilage 23 Ossifications of the Braincase 24 Mandible 26 Dentitiox 27 Sclerotic Plates — 28 Vertebrate 28 Cervical Vertebrae 29 Dorsal Vertebrae 33 Sacral Vertebrae 34 Caudal Vertebrae 35 Ribs 36 Pectoral Girdle 38 Pelvic Girdle 40 Limbs .-. 43 PHYLOGENETIC REL.4TIONSHIPS OF PETROLACOSAURUS 54 Hypotheses of Relatio.vships 57 Basic Taxa 58 Shared Derived Characters Testing the Hypothesis of Relationship Betv^^een Petrolacosaurus and Paleothyris 61 Shared Derived Char.acters Testing the Hypothesis of Relationship Between Petrolacosaurus and Youngina 62 Petrolacosaurus Compared with Aracoscelis 64 HEARING IN PETROLACOSAURUS AND OTHER EARLY REPTILES .. 66 CONCLUSIONS 69 SUMMARY 71 LITERATURE CITED 72 ) INTRODUCTION AH living reptiles can be grouped into Sauropsida, and therefore diapsids, could not four orders: Chelonia, Rhynchocephalia, be derived from captorhinomorphs. Squamata and Crocodilia. The latter three Romer (1966) rejected Watson's theory of can be associated with the largest assemblage the evolution of the ear and reiterated his of fossil reptiles, collectively called diapsid belief that all true reptiles can be derived reptiles. The diapsid condition refers to the from primitive captorhinomorphs. In a re- presence of two pairs of temporal openings view of early reptilian evolution, Romer on the skull roof behind the orbits. This con- (1967) suggested not only that the sauropsid- dition has been retained in a primitive form theropsid dichotomy, insofar as it exists, has in crocodilians and in the sole living rhyncho- little phylogenetic significance, but also that cephalian Sphenodon ptinctatum but has been the term Sauropsida has no systematic or modified in the squamates by the progressive phylogenetic meaning. Furthermore, he em- loss of dermal bones in the temporal region. phasized that the diapsids may not belong to Diapsid reptiles have an evolutionary his- a single phyletic unit. He argued that two tory long thought to extend only into the late discrete diapsid groups should be recognized, Permian. The early evolution of this assem- the Archosauria and the Lepidosauria. He blage is not well documented in the fossil proposed that the Lepidosauria (the squa- record. Consequently, the origins and phy- mates, rhynchocephalians and the ancestral letic relationships of member groups have eosuchians) on the one hand, and the Archo- been subject to dispute. sauria (the two dinosaur orders, pterosaurs, On several occasions, Watson ( 1951, 1954 crocodilians and the basic Triassic group, the reiterated his belief in Goodrich's concept of thecodonts) on the other, evolved separately a basic dichotomy among reptiles. According from the ancestral captorhinomoqjhs. It was to this theory, one group called the Therop- his suggestion that the miileretid skull pattern sida led to mammals and included the mam- represents an intennediate stage between the mal-like reptile orders Pelycosauria and The- ancestral captorhinomorphs and the lepido- rapsida. The other group, the Sauropsida, saurs. ' also called "true" reptiles, included the diap- In 1967 Reig set forth, and later (1970) sid reptiles and the turtles (Chelonia). Wat- elaborated, a radically different hypothesis for son argued for independent origins of the the origin of archosaurs. He proposed that Theropsida and Sauropsida from anthraco- the ancestors of archosaurs should be sought saurian amphibians via some unknown inter- among varanopsid pelycosaurs. Most of the mediate forms. He based this conclusion on comparable features cited by Reig as indica- his theory of the evolution of the ear (Wat- tive of phylogenetic relationships are, how- son, 1953). He discounted the possibility of ever, primitive characteristics found in the captorhinomorphs having given rise to the ancestral captorhinomorphs (Romer, 1971). Sauropsida and, hence, to the diapsid rep- Gow (1972), in his reconsideration of the tiles. He actually placed the Captorhino- milleretids, contended that this group gave morpha among the Theropsida. In 1957 Wat- rise directly to the Squamata. He felt on the son suggested that the diapsids evolved from other hand, that the Rhynchocephalia and the seymoriamorph anthracosaurs via the Millero- Archosauria had an entirely independent ori- sauria. gin. Vaughn ( 1955 ) also accepted the saurop- The above workers have based their theo- sid-theropsid dichotomy and felt that the ries of the origins and early evolution of the SPECIAL PUBLICATION MUSEUM OF NATURAL HISTORY NO. 7 diapsid reptiles mainly on the meager evi- primitive captorhinomorphs and to more ad- dence provided by the known Permian coty- vanced diapsid reptiles. The problem of hear- losaurs and on the Upper Permian and Trias- ing in Petrolacosaurus and other early reptiles sic diapsids. Thanks mainly to the labors of forms an integral part of this study. Robert L. Carroll and the late Frank E. Pea- body, the problem of the origins and early Acknowledgements evolution of diapsid reptiles can be reconsid- I am greatly indebted to T. H. Eaton of ered. Carroll, during a period of about ten the University of Kansas Natural History Mu- years, described and reconsidered all of the seum, at whose suggestion this study was available Pennsylvanian and Lower Permian undertaken and who kindly allowed me to specimens of primitive captorhinomorphs. One borrow the Petrolacosaurus specimens at his significant conclusion became evident: the disposal. I am particularly grateful to Dr. and described members of this group conform to Mrs. Eaton for courtesies extended to mc a single conservative morphological pattern during the time spent studying the Kansas suflBciently to indi- and remained generalized collection. cate that i^rimitixe captorhinomoqohs could Thanks are due also to R. L. Carroll whose be ancestral to most subsequent reptilian line- wise guidance, friendliness, and unrelenting ages (Carroll and Baird, 1972; Clark and Car- support were invaluable. I am very grateful roll, 1973). to P. L. Robinson of University College for In 1952, Peabody published a partial de- her many valuable and instructive sugges- scription of Petrokicosaiinis kansensis and tions, for numerous stimulating conversations, suggested that this reptile from the LTpper and for making the time spent studying in Pennsylvanian was a member of the most London so pleasant. I ha\'e profited also from primitive diapsid order, the "Eosuchia." The several interesting discussions with Donald fragmentary nature of the specimens available Baird of Princeton University and Malcolm for this study led others, however, question to Heaton of University of Toronto. Peabody's contention. Subsequently Peabody Thanks are due to Bobb Shaeffer and E. S. spent two grueling summers 1953 and 1954 — Gaffney, both of the American Museum of collecting more specimens but died before — Natural History, New York, for extended could undertake reconsideration of the he a loans of specimens. anatomy of Petrolacosatirus. This collection fonus basis of present These the the study. DESCRIPTIVE AND SYSTEMATIC show that Petrolacosaiirus is a specimens HISTORY diapsid reptile with affinities to both primitive captorhinomorphs and the earliest "eosuch- Lane first described Petrolacosaurus kan- iaiis." sensis in 1945 and referred the genus to the My interest in Petrolacosaiirus was aroused family Si)henacodontidae of the order Pely- during a visit to the Kansas University Mu- cosauria ( 1946) on the basis of a tarsus found seum of Natural
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