Museum of Comparative Zoology Breviora 13

B R E V 11 R A I seuin of Comparative Zoology HARVARD UNIVERSITY Numbers 464-487 1981-1986 MC2 LIBRARY NOV 1986 HARVARD UN CAMBRIDGE, MASSACHUSETTS 1986 02138 U.S.A BREVIORA Museum of Comparative Zoology CONTENTS Numbers 464-487 1981 No 464 The Origin of the Crocodiloid Tarsi and the Inter- relationships of Thecodontian Archosaurs Brinkman 23 pp December No 465 By Donald 30 The Structure and Relationships of the Dromasaurs (Reptilia: Therapsida) By Donald Brinkman 34 pp December 30 1982 No 466 Systematics of the Mexicana Species Group of the Colubrid Genus Lampropeltis, with an Hypothesis Mimicry By William R Garstka 35 pp June No 467 30 Three New Species of the Anolis Punctatus Complex from Amazonian and Inter-Andean Colombia, with Comments on the Eastern Members of the Punctatus Species Group By Ernest E Williams 38 pp June 30 No 468 A New Forest Skink from Ponape By A Ross Kiester 10 pp No 469 June 30 Catalog of the Primary Types of Bombyliidae (Diptera) in the Entomological Collections of the Museum of Comparative Zoology, with Designations of Lectotypes By Neal L Evenhuis 23 pp June 30 No 470 Systematic Implications of Innervation Patterns in Teleost Myotomes By Quentin Bone and R Dana Ono 23 pp June 30 No 471 Arthur Loveridge —A Life in Retrospect By Ernest E Williams 12 pp June 30 No 472 Fishes of the Suborder Labroidei (Pisces: Perciformes): Phylogeny, Ecology, and Evolutionary Significance By Leslie S Kaufman and Karel F Liem 19 pp June 30 1983 No 473 The Interrelationships of Pelycosaurs By Donald Brinkman and David E Eberth 35 pp April 29 No 474 Ruthiromia Elcobriensis, A New Pelycosaur from El Cobre Canyon, New Mexico By David A Eberth and Donald Brinkman 26 pp April 29 No 475 New or Problematic Anolis from Colombia New I Anolis from the Cloud Forest of Western Colombia By Stephen Ayala, Dennis Harris, and Ernest E Williams 1 pp April 29 Calimae, No 476 Species, Townsend's Unmapped North Atlantic Right Whales (Eubalaena Glacialis) By William E Schevill and Karen E Moore April 29 1984 No 477 New or Problematic Anolis from Colombia Propinquus, Another New II Anolis Species from the Cloud Forest of Western Colombia By Ernest E Williams pp No 478 September New or Problematic Anolis from Colombia III Two New Semiaquatic Anoles from Antioquia and Choco, Colombia By Ernest E Williams 22 pp September No 479 Agonistic and Courtship Displays of Male Anolis Sagrei By Michelle P Scott 22 pp September 1985 No 480 Three New Lizards of the Genus Emoia (Scincidae) from Southern New Guinea By Walter C Brown and Fred Parker 12 pp June 21 No 481 A New Anolis of the Lionotus Group from Northwest- ern Ecuador and Southwestern Colombia (Suaria: Iguanidae) By Kenneth Miyata June 21 No 482 New or Problematic Anolis from Colombia IV Anolis New Species of the Anolis Eulaemus Subgroup from Western Colombia By Ernest E Antioquiae, Williams pp June 21 No 483 Notes on Pristidactylus (Sqamata: Iguanidae) By Richard Etheridge and Ernest E Williams 18 pp June 21 No 484 Male Aggressive Behavior in a Pair of Sympatric Sibling Species By Jonathan B Losos 30 pp June 21 1986 No 485 The Anatomy and Relationships of Stereophallodon and Baldwinonus (Reptilia, Pelycosauria) By Donald Brinkman and David A Eberth 36 pp August 30 No 486 Thelodus "Macintoshi" Stetson 1928, The Largest (Agnatha: Thelodonti) By Susan Turner 20 pp August 30 Known Thelodont No 487 The Identity of Larval Parasudis (Teleostei, Chloroph- thalmidae), with Notes on the Relationships of Auli- poform J Fishes By Karsten Stiassny 24 pp August E Hartel 30 and Melanie L BREVIORA Museum of Comparative Zoology Index of Authors Numbers 464-487 1981-1986 Ayala, Stephen 475 Bone, Quentin 470 Brinkman, Donald Brown, Walter C 464, 465, 473, 474, 485 , Eberth, David E 480 473, 474, 485 Etheridge, Richard 483 Evenhuis, Neal L 469 Garstka, William R 466 Hartel, Karsten E 487 Harris, Dennis 475 Kaufman, Leslie S 472 Jonathon B 484 Losos, Liem, Karel F 472 Miyata, Kenneth 481 Moore, Karen E 476 Ono, R Dana 470 Parker, Fred 480 Scott, Michele P 479 Shevill, William E 476 Stiassny, Melanie L J 487 Turner, Susan 486 Williams, Ernest E 467, 471, 475, 477, 478, 482, 483 ORA B R W;X°1 Museum of Comparative Zoology I Cambridge, Mass 30 S ISSN 0006 %9K December Number 464 1981 THE ORIGIN OF THE CROCODILOID TARSI AND THE INTERRELATIONSHIPS OF THECODONTIAN ARCHOSAURS Donald Brinkman Abstract The tarsus of the proterosuchian Chasmatosaurus represents the primitive archosaur tarsus This kind of tarsus is also present in rhynchosaurids, trilophosaurids, prolacertids, and Protorosaurus, and suggests that these reptiles are members of a single radiation Two distinct kinds of crocodiloid tarsi are present in thecodonts, a crocodile-normal tarsus and a crocodile-reversed tarsus The crocodilereversed tarsus could have originated from the crocodile-normal tarsus, but the reverse relationship is not plausible Gracilisuchus, the only "ornithosuchid" with a crocodile-normal tarsus, shows features of the skull that are not consistent with placement in the postcranial characters, ancestral to is a ornithosuchid ancestor but could not be plausible pseudosuchian a its Ornithosuchidae Euparkeria on the basis of both cranial and with a crocodile-normal tarsus The tarsus of Erythosuchus neither contradicts nor supports a relationship between Erythrosuchus and rauisuchids INTRODUCTION In years, recent it has been recognized that a number of and an complex is structurally distinct kinds of tarsi are present in archosaurs, understanding of the evolution of this structural necessary for an understanding of the interrelationships of the group In the tarsus of crocodiles and typical pseudosuchians, the ankle joint passes between the astragalus and calcaneum, the astragalus being locked to the tibia and the calcaneum integrated with the pes astragalus 'Museum 02138 In dinosaurs, the ankle joint passes distal to the and calcaneum Krebs (1963, 1973) argued that this of Comparative Zoology, Harvard University, Cambridge, Massachsuetts BREVIORA difference derivation precludes No 464 of dinosaurs from any known pseudosuchian Recently, two additional kinds of tarsi have been recognized in thecodonts Proterosuchian thecodonts of the family Proterosuchidae have a distinct tarsus that primitive (Cruickshank, 1972; Carroll, in many ways Bonaparte (1971) is 1976) recognized that two distinct kinds of crocodilelike in pseudosuchians, one astragalus has a peg that like fits that fits in a crocodiles in a socket seen in advanced ornithosuchids in process that tarsi are present in which the on the calcaneum, and one which the calcaneum has a of socket on the astragalus Chatterjee (1978) termed these the crocodile-normal and crocodile-reversed tarsus respectively Also, two pseudoschians with mesotarsal ankle joints have been described (Romer, 1971) One of these, Lagerpeton, has a fully developed mesotarsal joint The second, Lagosuchus, retains a and a complex articulation between the astragalus and calcaneum (Bonaparte, 1975a) The evolution of these tarsal patterns was recently discussed by Cruickshank (1979) Cruickshank showed that the proterosuchian tarsus is an excellent structural ancestor to the crocodile-normal tarsus and argued that the two kinds of crocodile tarsi can be used to separate pseudosuchians into two groups Based on this, Cruickshank suggested that Gracilisuchus, which has a crocodilenormal tarsus, be removed from the Ornithosuchidae, all other members of which have a crocodile-reversed tarsus However, the origin of the crocodile-reversed tarsus remains unknown If the crocodile-normal tarsus was ancestral to the crocodile-reversed tarsus, then a crocodile-normal tarsus could have been present in primitive ornithosuchids, and the presence of a crocodile-normal tarsus would not bar Gracilisuchus from the Ornithosuchidae Thus, in order to use the structure of the tarsus as a basis for posteriorly directed calcaneal tuber interpreting the interrelationships of archosaurs, it is necessary to obtain a more precise understanding of both the origin of the crocodile-reversed tarsus and the phylogenetic position of Gracili- suchus THE CROCODILE-NORMAL TARSUS In extant crocodiles, five elements are present in the tarsus: the astragalus, the calcaneum, and the second to fourth distal tarsals LARVAL PARASUDIS RELATIONSHIPS 1986 more attenuated body form Halliday (1968: Fig nile that The shows in the recently 1) illustrates initial a 99.0 mm transformed juvenile SL transformed juve- body deepening internal pigmentation of the anterior viscera is present in transforming specimens Apparently during transformation, pig- ment migrates from the visceral peritoneum onto the parietal peritoneum, and the pigmentation of the caudad extension of the mesentery is greatly reduced In fully transformed juveniles the parietal peritoneum is strongly pigmented throughout the abdominal cavity but little pigment is associated with the visceral peritoneum We have examined a number of transforming Chlorophthalmus specimens from both the North Atlantic Ocean and the Coral Sea In each of these specimens, tranformation occurs at a considerably smaller size (ca 35 to 40 mm SL) than that observed in our Para- sudis specimens DISTRIBUTION The western Atlantic specimens we have examined come from areas off the Brazilian coast to areas east of the southern tip of Scotia (Fig specimen 5) (MCZ larva and its Included in our study Nova one eastern Atlantic 62402) Preliminary morphometric analysis of this is eastern Atlantic locality indicate that it may represent Parasudis fraser-brunneri (Poll 1953) However, due to a lack of additional material for detailed study and the close morphological resemblance of 1966:184), we P fraser-brunneri to P truculenta (see Mead are unable to confirm the specific identity of the eastern Atlantic specimen midwater trawls Speciopening and closing nets (MOC 10 and 20) indicate that the larvae are most commonly taken above 150 m (five m; to 100 m; 70 to 140m; to 203 m; 70 to 150 collections: 70 to m) Information from non-closing nets indicate similar distributions However, other collections taken with non-closing nets indicate that the larvae may approach the surface rather closely, since Parasudis larvae are found in collections taken above 50 m Except All of the larval Parasudis were taken in mens from the WHOI for a few larvae from the specimens were taken 4934 m deep relatively shallow Straits of Florida, all in trawls over ocean bottoms from 1937 to BREVIORA 10 O HO 40* 60 No 487 0" ( *C o-r?^^y r iO *y £o wq ( 40 6° ,0 ) o ) o o « =£>