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S. Associate in Vertebrate Palaeontology Ph.THE OSTEOLOGY OF THE REPTILES BY SAMUEL WENDELL WILLISTON M.D.D. American Museum of Natural History..D. Ph..A. Late Professor of Palaeontology in the University of Chicago ARRANGED AND EDITED BY WILLIAM KING GREGORY. New York Professor of Vertebrate Palaeontology Columbia University CAMBRIDGE HARVARD UNIVERSITY PRESS 1925 .D.. N. Sc.


i8s2-igiS. Vol." Journal of Geology. The American Museum of article. comprehensive survey of the evolution of the Reptilia as a whole.^ Death overtook him before the final revision and completion series of of this finished the greater part of the text work. 1918. repeatedly found itself unable to accept this one notwithstanding its good will." The new drawings that Williston made for this book have been supplemented by earlier works. xxvi.FOREWORD book we have the exploration and research In this chief results of Williston's half-century of in the field of vertebrate palaeontology. and through his good offices the Harvard University Press now has the honor of publishing the "Osteology of the Reptiles. and private publishers proposed conditions that were not acceptable. but happily not before he had and had made for it with his own pen a large new and excellent line-drawings. Here we find the gist of his earlier researches upon the mosasaurs. many other illustrations. plesiosaurs. After much unsuccessful correspondence in vari- ous directions. either to the Williston Memorial Committee. which had published Williston's earlier books. while others many of these the authors to 1 whom have been copied from the original pubHcations of they are credited. The University of Chicago Press. For the long delay since 191 8 there have been too many causes to be profitably set forth in detail. 673-689. and the epitome of his last. mostly from Williston's illustrate the present text. . The writing of this book was thus the culminating effort and achieve- ment of his inspiring career. which were needed to The University of Chicago Press has courteously loaned cuts. pp. the sad plight of Williston's unpublished work of came to the notice of Professor Thomas Barbour Harvard Uni- versity. " For an excellent account of Williston's life and work see Henry Fairfield Osborn's Samuel Wendell Williston. and pterosaurs of the marine Cretaceous of Kansas. or still to Professor Williston's family. In accordance with Williston's wishes the writer undertook to put his last work in shape for the publisher and to see it through the press. the substance of his later and fundamental discoveries among the primitive reptiles of the Permian of Texas.

vi FOREWORD Natural History. Every tions effort has been made to keep intact the spirit and letter of the original text. has at all times given indispensable support in the work of making ready book for the press. Meadowcroft of the Museum. Fink and Mrs. and in the few places where corrections or emenda- have seemed necessary they have been placed in square brackets. K. . C. Noblcj Curator of Herpetology. as also the footnotes that record some of the more conspicuous discoveries and advances since 191 8. E. H. with the cordial approval of President Osborn. has supplied critical notes on the this sections dealing with recent reptiles. P. W. Special acknowledgment is due to Mrs. G. Dr. G. K.

Squamata LacertUia or Sauria Ophidia or Serpentes Rhynchocephalia Pseudosuchia Pelycosimia Phytosauria Crocodilia 65 66 72 Dinosaurs (Saurischia. External Appearance. Excrescences. Ornithischia) Pterosauria 74 77 77 79 83 87 88 CHAPTER The Vertebrae II 90 CHAPTER The Ribs and Sternum p-a X'\^ III 112 .CONTENTS PART I THE SKELETON OF REPTILES INTRODUCTION PAGE The Primitive Skeleton of Reptiles The Primitive Skull of the ReptHia 3 6 7 The Primitive Postcranial Skeleton CHAPTER The Skull of Reptiles I g . Chief Openings The SkuU Elements The Mandible The Skull of the Cotylosauria Chelonia 9 13 27 33 Theromorpha Therapsida Nothosauria Plesiosauria 44 46 52 56 56 59 Placodontia Ichthyosauria Protorosauria 60 62 .

VIU CONTENTS CHAPTER The Pectoral and Pelvic Girdles The The Pectoral or Shoulder Girdle Pelvic or IV 124 1 24 142 Hip Girdle CHAPTER V The Limbs The The The The Propodials Epipodials 155 158 165 169 195 Mesopodials Metapodials and Phalanges PART II THE CLASSIFICATION AND RANGE OF REPTILES CHAPTER The Problem of Classification VI 205 CHAPTER A VII 210 Synoptic Classification of the Reptilia CHAPTER Subclass Anapslda Order Cotylosauria [Suborder Seymouriamorpha] Family Seymouriidae . VIII 215 215 217 [Suborder uncertain] Family Sauravidae [Suborder uncertain] 217 217 218 Family Gymnarthridae Suborder Diadectosauria Family Diadectidae Bolosauridae Suborder Labidosauria Family Captorhinidae Pariotichidae 218 218 218 218 218 218 218 218 Stephanospondylidae Genera incertae sedis [Suborder uncertain] Family Limnoscelidae Suborder Pantylosauria Family Pantylidae Suborder Pariasauria Family Pariasauridae 220 220 220 220 .

CONTENTS Suborder Procolophonia Family Procolophonidae [Suborder uncertain] IX 220 220 221 221 Family Elginiidae Order Eunotosauria Order Testudinata or Chelonia Suborder Amphichelydia Family Proganochelydidae Pleurosternidae 222 Baenidae Suborder Pleurodira Family Pelomedusidae Chelyidae Miolanidae Suborder Cryptodira Family Thalassemyidae Toxochelyidae Desmatochelyidae Protostegidae Cheloniidae 223 223 223 224 [Dermochelyidae] Chelydridae Dermatemyidae Emydidae Testudinidae Suborder Trionychoidea Family Plastomenidae Trionychidae 224 224 224 224 224 225 225 225 225 225 226 226 226 226 227 227 227 227 CHAPTER IX Subclass Synapsida Order Theromorpha Suborder Pelycosauria Family Sphenacodontidae Suborder Edaphosauria Family Edaphosauridae Suborder Poliosauria Family Poliosauridae Ophiacodontidae Suborder Caseasauria Family Caseidae Suborder uncertain Family Paleohatteriidae Genera incertae sedis Order Therapsida Suborder Dinocephalia Family Tapinocephalidae Deuterosauridae 228 228 233 233 233 233 233 233 233 233 233 236 236 236 236 237 238 239 .

X CONTENTS Rhopalodontidae Titanosuchidae Suborder Dromasauria Family Galechiridae Galeopidae Macroscelesauridae Suborder Anomodontia Family Dicynodontidae Suborder Theriodontia Tribe Gorgonopsia Family Gorgonopsidae Ictidorhinidae Burnetidae Tribe Bauriasauria Therocephalia Family Scylacosauridae Ictidosuchidae 239 239 239 239 239 239 239 240 240 242 242 242 242 242 243 243 243 243 243 Lycosuchidae Scaloposauridae Alopecopsidae Whaitsidae 243 243 243 Genera incertae sedis Tribe Cynodontia Family Nythosauridae Cynosuchidae Cynognathidae Diademodontidae . Theriodontia (?) incertae sedis 244 244 245 245 245 245 CHAPTER X Subclass Synaptosauria Order Sauropterygia Suborder Nothosauria Family Nothosauridae Suborder Plesiosauria Family Plesiosauridae Pliosauridae 246 246 247 247 248 248 248 250 250 250 251 251 Cryptocleididae Elasmosauridae Polycotylidae Brachaucheniidae Genera incertae sedis Order Placodontia Family Placodontidae 251 252 CHAPTER XI Subclass Parapsida Order Proganosauria 253 253 .

.' Order Squamata Suborder Lacertilia (Sauria) Tribe Kionocrania . Family Geckonidae Euposauridae Agamidae Iguanidae Anguinidae Helodermatidae Lacertidae Tejidae Scincidae Tribe Platynota 264 265 266 266 266 266 267 267 267 268 268 268 Family Varanidae Dolichosauridae Aigialosauridae Tribe Pythonomorpha (Mosasauria) Family Mosasauridae Globidentidae Tribe Amphisbaenia 269 269 269 270 272 273 273 -. Family Amphisbaenidae Tribe Rhiptoglossa 273 274 Family Chameleontidae Genera incertae sedis Suborder Ophidia (Serpentes) Family Typhlopidae Boidae (Pythonidae) Paleophidae Viperidae Elapidae Colubridae 274 274 274 275 276 276 276 276 277 277 CHAPTER Subclass Diapsida ? Order Proterosuchia XII 278 278 .CONTENTS Family Mesosauridae Order Ichthyosauria Family Mixosauridae Shastosauridae Ichthyosauridae XI 255 255 256 256 256 258 Ophthalmosauridae Order Omphalosauria Family Omphalosauridae Order Protorosauria Family Araeoscelidae ? 258 258 259 259 261 Protorosauridae Saphaeosauridae Pleurosauridae 262 262 < .

xii CONTENTS Order "Eosuchia" Family Younginidae 278 278 279 279 279 281 Superorder Diaptosauria Order Rhynchocephalia Suborder Rhynchosauria Sphenodontia (Rhynchocephalia vera) ? Choristodera Suborder Thalattosauria Superorder Archosauria Order Parasuchia Suborder Pseudosuchia Family Aetosauridae Ornithosuchidae Scleromechlidae Suborder Pelycosimia Suborder Phytosauria Family Phytosauridae Stagonolepidae [Suborder Desmatosuchia] 283 283 284 284 284 285 285 285 285 286 286 286 286 287 Order Crocodilia [Loricata] Suborder Eusuchia Family Teleosauridae Pholidosauridae Atoposauridae Goniopholididae 288 288 288 288 . Predentata] Suborder Ornithopoda (Predentata) Family Nanosauridae Hypsilophodontidae Iguanodontidae 293 293 293 294 294 294 294 295 . 289 Dyrosauridae Hylaeochampsidae Gavialidae Tomistomidae Crocodilidae 289 289 289 289 290 Genus incertae sedis Suborder Thalattosuchia Family Metriorhynchidae [Dinosauria] 290 290 291 291 291 Order Saurischia Suborder Theropoda Family Plateosauridae Anchisauridae 291 291 292 292 Suborder Sauropoda (Opisthocoelia. Cetiosauria) Family Cetiosauridae Camarasauridae Atlantosauridae Diplodocidae 293 293 Genera incertae sedis Order Ornithischia [Orthopoda.

CONTENTS Trachodontidae (Hadrosauridae) Suborder Stegosauria Ceratopsia xiii 295 295 295 296 Order Pterosauria Suborder Pterodermata (Rhamphorhynchoidea) Family Rhamphorhynchidae Suborder Pterodactyloidea Family Pterodactylidae Ornithocheiridae 296 298 298 Pteranodontidae Nyctosauridae Genera incertae sedis 298 298 298 298 298 .




the loss of gills in their youth and entire emancipation from the water. and their evolution was more rapid. and more spe- cifically assured. for the most part. retaining in in this interval. Both the reptiles and the amphibians had changed of millions of years. frogs. the beginning of their stock. Characters that are common to many amphibians became more and more rare among the reptiles. salamanders. At tion of Eosauravus that time there was a considerable diversity of known forms. The modern toads. varying degrees their ancestral characters. became more progressive than the amphibians. were restricted more and more to subordinate roles. and only a few of the more progressive continued to develop. They.INTRODUCTION THE PRIMITIVE SKELETON OF REPTILES That the reptiles were evolved from the Amphibia. of which. The reptiles. while losing or adding others in various by the acquirement of a new mode of life. and the amphibians. and by the close of Triassic times had become restricted to habits and habitats no longer invaded by them. more earli- from which we may very properly conclude that their est ancestors. belong- ing to at least four well-differentiated groups and twenty or families. lost those characters and adaptations that brought them into immediate competition with the reptiles. and blindworms differ far more from the higher amphibians of Paleozoic times than did the latter from their con- temporary reptiles. and very prob- ably in Lower Carboniferous times. . an interval perhaps ways. know belong With the excep- from the middle Pennsylvanian of Ohio. our knowledge of them goes no further back than the late Carboniferous and early Permian. unfortunately. the skull is unknown. from that order known as the Temnospondyli. lived much earlier. to find in the rocks of the We therefore never can expect Permian any real connecting link between the two classes. handicapped by inherited habits. cer- tainly at the beginning of the Upper Carboniferous. seems now The earliest as also the most primitive reptiles that we to the order called the Cotylosauria.

. from nearly A little less than one third natural size. Primitive Cotylosaur Seymouria. from above. I. complete specimen.Fig.

upon the widely open palate of the Amphibia as a final distinguishing charthe beginning of their evolution into acter of their class. but earliest tiles. except in the and feet. Seymouria (Cotylosauria). while all reptiles had acquired a reduced embolomerous form with one disk. A. many tiles bones in the digits no amphibians with as as the early reptiles possessed. Fig. We relied. from side. In general literature the Amthe phibia are distinguished from Reptilia by the possession of two occipital condyles. with a considerable gap in the structure of their ver- tebrae. but only a very few of their stock persisted as late as the Per- mian. In the vertebral column there was a general change among the Temnospondyli from the embolomerous to the rhachitomous from the more simply divided centrum of two disks to the tripartite centrum composed of wedges. an inheritance their ancestral fishes know from had the we Lower Carboniferous that — — of which only one known dea. retained the single condyle until mammals.INTRODUCTION Nevertheless. the intercentrum. there were still so many inherited characters among both the amphibians and atlas reptiles of early Permian times that nothing distinctive of either class can be found in the skeleton. how- natural size. ever. of the amphibians had a is [closed] palate like that of the [earliest] rep- but of these none at the beginning of Permian times. until recently. when they too developed a double condyle. The all earliest amfrom phibians doubtless had a all single occipital condyle. In other words. a single condyle and a closed palate are more primitive characters of the tetrapods than those we had assumed as characteristic of the Amphibia. One third scendant with that character survived to the Permian. The reptiles. and one t3^e. wedge. We know with as many bones in the tarsus as the early . and no repamphibians had. all Doubtless amphibians of Lower Carbonif- erous times had embolomerous vertebrae. from above. we now know known that some. that is. the centrum. B. if not all.

eight pairs of cranial bones. four pairs of palatal bones. . the supraorbitals of various Squa- mata and the predentary and doubtless the last two. with five openings in roof: A. and not are simply dermal bones came temporarily attached thus. Median parietal (pineal) foramen.. — A. They may be summarized as follows: The Primitive Skull of the Reptilia Rugose. Premaxillae (px) Maxillae (mx) Septomaxillae (sx) p.2) from the Lower Permian of Texas. in front of the middle of the skull]. The girdles which beand limbs of the two classes are distinguishable only by minor characters. Paired Bones 1 3. And there is no bone in the skeleton of reptiles that is interfrontal of a very few. Paired orbits beyond middle [i. one unpaired palatal bone seventy-eight in all. B. from the what were the primitive characters of the reptilian skeleton in almost every detail. Nasals (no) Frontals (fr) Parietals (pa) Interparietals (ip) Median roof bones . 7. C. 5 6. And of the we do not know from these later rocks. in a single genus Seymouria (Figs. except the rostral all. except the been found in the skull of early reptiles. 1. eight pairs of mandibular bones. three unpaired cranial bones. Seventeen pairs of roof bones. for the ear [the "otic notch"].6 THE OSTEOLOGY OF THE REPTH^ES feet of the earliest reptiles feet of the but doubtless when we discover the shall find we them not different from the contemporary am- phibians. while to the skull. and probably all never shall from rocks later than the Lower Carboniferous. 1. / Nasal bones 4. if bones of certain dinosaurs. terminal nares. preparietal of the Anomodontia. indeed. 2. divided. and all. between tabular and squamosal.e. characters common to the two classes in comparison of all it is not difficult to decide any one animal. An emargination of the occipital border. Every known bone in the skull of the Temnospondyli. has not known in these same amphibians. Paired.

Quadrates (^m) Articulation of mandible Exoccipitals (eo) Paroccipitals (poc) Prootics (pc) Postoptics (al) \ Cranial bones Stapes (sip) Epipterygoids (ep) Ethmoids (se) AA. Supratemporals (st) Tabulars (/) Squamosals (sq) Quadratojugals (qj) Dentaries (d) Coronoids (cor) Splenials {sp) Temporal bones 20. 2. 37. 32. Membrane Bones. 27. 39. 34. 24. Basisphenoid (bs) The Primitive Postcranial Skeleton A. cleithra and elongate interclavicle 3. 26. Dermal plates or scutes. 19. 35. 16.INTRODUCTION 8. 21. 13. Paired clavicles. Prefrontals (pr) Postfrontals (pf) Postorbitals (po) 10. 11. 22. 1. Postsplenials (psp) Angulars (an) Prearticulars (pa) Mandible 23. 28. Surrounding orbits Jugals (j) Intertemporals (it) 14. 25. 29. 31. Unpaired Bones Palate Cranial bones Parasphenoid (ps) Supraoccipital (so) Basioccipital (bo) |. 40. 17. Surangulars (sa) Articulars (art) Prevomers (pv) Palatines (pi) Palatal bones. . Lacrimals (la) 9. 38. 12. dentigerous (ec) Pterygoids (pi) Ectopterygoids 30. 18. 36. 15. Exoskeletal Sclerotic plates in orbits. 41. 33.

atlas embolomerous. 8. about twenty-three presacral. ^ [But see footnote i. 3. the fourth finger strongest and longest. 4. all vertebrae to tenth or twelfth caudal with free. 10. a supracoracoid and a supraglenoid foramen. Hand pentadactylate. fused in adult life. 5. Cartilage Bones. procoracoids and metacoracoids.THE OSTEOLOGY OF THE REPTILES AA. two (three?) in middle row. all well ossified. holocephalous ribs. 5. 6. the fourth toe strongest and longest. closed.] . the three forming the glenoid socket. phalangeal formula 2. ^ [But see footnote on page 187 below. Notochordal vertebrae. 4. 1. two or three cervicals. Carpus with four bones in proximal row. — Ed. Endoskeletal 2. 4. Legs short and stout. .] — Ed. 1 1 12. five in distal row all well ossified. Slender and numerous parasternal ribs. 4. 7. No sternum. 3. on page 122 below. 3. Intercentra between all vertebrae. Scapular girdle composed of paired scapulae. . 3. Tarsus composed of nine bones two in first row ^ two in second five in distal row. articulating continuously with intercentrum and diapophysis. : . with entepicondylar foramen. proatlas. 5. 9. Feet pentadactylate. acetabulum formed by the three bones. Humerus dilated at extremities. phalangeal formula 2. Pelvis plate-like with small obturator foramen only. one sacral. moderately long tail.

offensive and defensive habits. Excrescences or horns on the skull have been developed in not a few. The earliest known is that of the cotylosaurian Chilonyx. and the theromorph berances. in- sectivorous reptiles may have a more slender skull (Fig. and others have united or formed new associations. 45) is more or less elongate. 70 a) the a median facial and supraorbital rugosities. Even some skull. with large protu- excrescences. not only on the face Perhaps of few other but also along the posterior margin of the greatly extended skull. like Elginia. while those reptiles using the jaws to crush invertebrates always have a short and powerful skull (Fig.CHAPTER I THE SKULL OF REPTILES External Appearance. turtles. horned lizards and moloch have many sharp protuberances and horny excrescences. 33. as in the modern gavials. with Tetraceratops. 49). reptiles (Fig. It has not a few bones in various forms. like the southern Miolania. and Chief Openings The lost skull of reptiles. sometimes very long (Fig. to such an extent. A few carnivorous dinosaurs have (Fig. as of other vertebrates. indeed. which. though not a lizards. Excrescences. of horns or spines. 55 d). though a few have been skull of carnivorous added temporarily from the exoskeleton. that there are several in later reptiles about whose homologies there has been and solid yet is dispute. The reptiles (Figs. fish-eating always long. has undergone many changes in adaptation to food. as though for the support In the Ceratopsia development of horns and spines was carried to a remarkable degree. like the horns (Fig. like that of a wolf. 67). It has developed excrescences or horns for defense or offense. 52 b). had horny protuberances at the back part. all reptiles none has surpassed some of the modern chafacial meleons in the development of lizards. but it has gained permanently no new bones. have horns upon the facial Usually the median unpaired horn is borne by the . 58) is The face of aquatic. or has been covered at times with a armor of skin bones. Some of the later Cotylosauria. were size of they magnified to the dinosaurs would be equally imposing.

thalattosaurs (Fig. A partial secondary palate. Primitively located near the extremity of the face (Figs. are located far They back from the extremity in the volant pterodactyls (Figs. as in the chameleons it is formed by the maxillae. 46 a). and the lacrimals have lost relations with them since Permian times. to the posterior border of the palatines. The internal nares. but in the nasals.. 55). Figs. formed by the undergrowth of the maxillae and palatines as a secondary palate.lO THE OSTEOLOGY OF THE REPTILES mammals. in the later crocodiles even .3. A similar respiratory canal. In the plesiosaurs the internal nares are also {e. 50). 68) and Choristodera (Fig.g. In the very slender-faced amphibious Crocodilia (Fig. 31. normally situated almost immedi- by a respiratory canal. life Doubtless such horns or protuberances were covered in {external nares) with a horny sheath. to a greater or less extent in the Cynodontia and ately below the external (Fig. they are often confluent in later ones (Figs. In most aquatic reptiles they have receded toward the orbits. chameleons (Fig. In those reptiles in which the external nares are situated posteriorly. there may be a partial reversion of the respiratory . 67). wherever located. the supraoccipital in pteroall into enormous saurs. 68). or rather the face has grown away from them. however. plesiosaurs (Fig. 2. 59. is character- the Phytosauria (Figs. The paired facial horns are borne by the prefrontals or The frontals and parietals are sometimes developed crests in the dinosaurs. and they almost always have the same relations with the first three of these bones. into the pterygoids.). 46). 69) in the former and in the early kinds of the latter. 66). 32. nasal and lacrimal. or choanae. with the opening way back. 61). are carried back Crocodilia (Fig. often for a long distance. the all nasals are often excluded from them. probably separated from the cavity istic of of the mouth by a membrane only. 72) as in most birds. formed by the union only a little of the palatines or maxillae. 63). each was surrounded by the premaxilla. 55).66). maxilla. the nostrils retain their primitive position at the extremity of the face. etc. only (Fig. 71. Well separated by the premaxillae and nasals in the older reptiles. 66. and phytosaurs (Fig. postorbitals. as in the ichthyosaurs (Fig. They by the are surrounded by the maxillae in the nasals in the phytosaurs (Fig. external nostrils The vary greatly in position. 61. 67). proganosaurs. occurs in some Chelonia and Anomodontia.

2. ternal nares. 71c) lie between the prevomers and palatines. etc. Sauropterygia (Fig. 62 d). but absent in some true lizards. The 32). very large in certain shell-eating cotylosaurs (Fig. orbits. the postorbital in Araeoscelis (Fig. 59). leaving not a single element invari- ably associated with the orbit. in all the Theromorpha (Figs. all the b. only the prefrontal and postorbital may be left. primitively (Figs. 43) a separate bone. 65 30. 59). 33-42). postfrontal. maxillae and palatines. 55). but usually lacrimal. jugal. known. a small unpaired element of doubtful homologies. 46.. In snakes (Fig. may sometimes (Figs. 47) divided The parietal or pineal foramen. occurring in (Figs. 70 a). 55. 48). 67 a) all phytosaurs and true pseudosuchians (Fig. and all snakes (Fig. 31. 59). 45. and most lizards (Figs. the chameleons. but with these possible exceptions appears to be absent in Archosauria (Figs. 50). and therapsids (Fig. 44. absent in other reptiles. the preparietal. 52) and Hyperodapedon (Fig. most Saurischia (Fig. and The frontal usually forms a part of the upper margin. 43. is 55 A. 66 b. It has been reported in certain doubtful Pseudosuchia and more or less doubtfully in a few phytosaurs and dinosaurs. 56). 49. 21. had It is present. (Figs. 3. 68. laterally. The prefrontal is excluded in some dinosaurs. 71) and some Crocodilia. 70 a. the Diaptosauria (Figs. 22. b) and Pterosauria (Fig. so far as become inconstant even in that order. 30. 65) surrounded by the prefrontal. T^T. 49. 23. directed upward sometimes were primitively in aquatic animals (Fig. with the internal opening in front of the external. 45 d). 66 b. d). 32). it 43.THE SKULL OF REPTILES canal. Usually there is but one.).) as also the Chelonia (Figs. in the Proganosauria. 6. . in front of or surrounding the foramen. may occur between the frontals pos- teriorly (Fig. (Figs. but there may be two or even three on each side in certain Theropoda (Fig. 22). the maxillae sometimes below (Figs. 44. 48. II The in- by the prevomers and surrounded by the premaxillae. there is In the Anomodontia and Gorgonopsia (Fig. 59). Antorhital or preorhital vacuities are very characteristic of the Archosauria. Usually located between the parietals anteriorly (Figs. 65 b. Sometimes the postorbital bar is incomplete in lizards (Fig. 22. Ichthyosauria (Fig. 56). snakes (Fig. 56. postorbital. 33. etc. 60-62). and in the Therapsida with the exception of a few forms. 43-45).

Hes between the postorbito-squamosal arch and the parietal. or parietal and squamosal. as in Squamata. the jugal is excluded from the lower margin fail to by the union of the squamosal and postorbital. 54). the squamosal may meet the postorbital above the opening. 49) have the single opening bounded below by the squamosal and postorbital. 19.12 THE OSTEOLOGY OF THE REPTILES the skull roof. It is (Figs. it is like the upper one of those reptiles with two temporal openings. that is. the supratemporal. 44 b). 48) and Placo- dontia (Fig. and until its certain nature will not be determined more is known of their terrestrial antecedents. paired subtemporal openings occur they are bounded medially by the pterygoids and the basis cranii. 22) and Chelonia. It is usually considered to be what it really appears to be. Either the upper or the lower opening may occur independently. 53 c). The lower or lateral temporal opening ap- peared primitively jugal. permitting the parietal to form the upper boundary in part. the quadratojugal (Figs. are character2. 47. In not a few of the Therapsida. but has. 64. In the Cynodontia (Fig. a vacuity between the parietal. S3.) is The post-temporal opening situated on the occipital aspect of the skull. 62. the Dinocephalia especially (Fig. 44 f. below by the jugal. an additional bone helping to form its posterior or outer border. The intertemporal vacuity. 55. b). 1 [In addition to the openings noted in all reptiles in the palatal aspect of the skull. or both together. 45) and some other Theriodontia. 65. 46. to which was added.] dermal covering of the temporal region. and this is the condition in mammals. It is present in some Cotylosauria by the author. save the Cotylosauria (Figs. above by the sides of the parietal. 4. (See pages 61-69 below. 50). in some of the double-arched forms. when complete. arose primitively by the separation of the postorbito-squamosal bar (Fig. and certain other forms here grouped under the Parapsida. for there its real is doubt as to homology. the Ichthyosauria (Fig. and the paroccipital on each side. or by the Ed. ^3 a) from the parietal (Fig. — . in most if not all. laterally by the lower temporal bar. The upper opening. if present. All known forms of the Sauropterygia (Figs. with a lower temporal opening (Figs. The single temporal opening of the Squamata (Figs. 70 A. 45 d). 53 a) between the squamosal and the bounded above by the postorbito-squamosal arch. the upper temporal opening only. the supratemporal or tabular.^ Openings through istic of all reptiles back of the orbits.

54 c). 700. some lizards (Fig. 4. 3. the chameleon lizards. articulating directly with the frontals. anomodonts (Fig. 46. 29) or They do not occur in the Cotylosauria (Figs. 24. strictly aquatic reptiles (Figs. where the dentigerous border is greatly elongated. most reptiles Theromorpha. 72 a). the posterior process forming a partial division between the nasal chambers. 46. and many snakes. 7. Theromorpha (Figs. 6. 45 g). The cranial region thus exposed by these various openings has been exposed to a greater or less degree in most Chelonia (Figs. 3032) in a different way: by the emargination of the roof bones from behind or from behind and below.THE SKULL OF REPTILES (Fig. and is generally present in later reptiles though absent or vestigial in the Crocodilia. five teeth in each. (Fig. and sometimes united with the nasals (Fig. 63. 47) Elongate in the and in the Pterosauria.3. 40 c. The dentigerous part is short in the long-faced plesiosaurs (Fig. and thalattosaurs (Fig. when present. 1 and Theromorpha (Fig. Edentulous in the chameleon lizards they take no part in the boundary of the nares. some dromasaurians. though there are as many as twenty. Primitively short (Figs. 66. They are also edentulous in the turtles (Figs. They are situated between the palatines and maxillae posteriorly. though present 21 a. The Skull Elements The primitive relations of the skull elements may be discussed seri43)- atim. They are often fused Teeth. 30-32). (Fig. 72) since the goids. articulating with maxillae. separating the frontals. with their chief modifications in later reptiles. are in a single row and rarely exceed five or six in number in each. a median pro- longation separates the nasals. terepenes. 72). 44 c). They form Four or the anterior boundary of the external and internal nares. 42 c).0). the later pterodactyls (Fig. most ornithischians (Fig.three in some phytosaurs (Fig. 55). in many Therapsida. 21 b). 19. 69. and are usually also bounded in part by the ectopteryPosterior palatine or suborbital openings occur in (Figs. but are absent in some turtles. in the first group (Fig. the whole temporal roof until. 66) and even more in ichthyosaurs. 47). . Premaxillae (px). 72). 2. 61). 55. nasals and prevomers. 56). In the plesiosaurs (Figs. in some forms like the is lost. 22. 48 c). 46) sometimes directly with the parietals. pterosaurs (Fig.

69). 28-32) and Crocodilia (Figs. single or etc. later c). 68. 44) in the crocodiles (Fig. 3. 28-32) either absent . In most reptiles since Permian times they also articulate with the nasals above {e. side. 43.) Present probably in Nasals (na). and lacrimals. Except in most Cotylosauria (Figs. 4. and in most Chelonia (Figs. Pterosauria (Fig. 69 b) with each other on the palate. F. 43). some Anomodontia (Fig. they also articulate with the maxillae on the sides. They are edentulous in the Chelonia (Figs. 3. prefrontals. 72). some Theromorpha and Therapsida. 2 b. 44 Ophidia. from natural size. g. located partly within the nasal chamber. (Figs. odontia. etc. above with septomaxillae and lacrimals. 59). they are absent in the reptiles. 33. Articulating with premaxillae. frontals..14 THE OSTEOLOGY OF THE REPTILES Primitively (Figs. 22. 21. as also in many Anomodontia and Theri. Maxillae (mx). A. the so-called turbinals of reptiles. 30-32). They are absent in many Chelonia (Figs. posteriorly with the jugals. 6. forming the outer boundary of both external and internal nares in part. 19. Saphaeosaurus. and post- orbitals internally. 3. Fig. Figs. 23. Small bones. The teeth may be in numerous rows. Ornithomimus. In some Dromasauria they extend back on the face to meet the lacrimals. ectopterygoids. B. articulating anteriorly with premaxillae. Pantylus (Cotylosauria). 22. Little can be said about them in other extinct reptiles. but appearing more or less on the outer 44 side at the back part of the external nares. t. . Three fourths Septomaxillae {sx).). forming more or less of the partition between the ex- ternal nares.^. Dromosauria. 45. 2. 33. all the earhest and most early Squamata (Fig.

also directly with the pterygoids. d. Interparietals (ip). 68. etc. 67). supratemporals. etc. 30) and Crocodilia. epipterygoids. often formin their relations. prefrontals. they are situated on the occipital surface and are usually unpaired. 22. supposed to be these bones. 69. but surround them in the Phytosauria Very large in the Ichthyosauria (Fig. 9). Unknown (la). 2.. 33. They are often fused in the midline. 42 d). Primitively (Figs. and interparietals. Figs. 44. originally named dermosupraoccipitals. 2 b. In the absence poral and supratemporals. In the later Cotylosauria (Fig. articulating with pariand supraoccipital.g. 56). They do not help form any part of the cereparietals etals. they also articulate with the large postfrontals. and some lizards (Fig. postfrontals. they 56). 44 a. and jugals.) large. A vestige. ex- tending from orbit to nares. 2. Lacrimals Primitively (Figs. pterosaurs (Fig. Fused in most late reptiles (e. 46). 22. Often fused in midline (Figs. some Therapsida (Figs. 54 c) . 46) They do not (Fig. articulating with prefrontals. {e. 66. 4.) of the intertem- and prootics. 22) back of the on the superior surface of the skull. and in some of the former are separated externally in the middle. intertemporal.. septomaxillae. 72). . Figs. g). or doubtful in other reptiles. 23. 1 sepa- rated by the premaxillae in the plesiosaurs. Frontals (fr).THE SKULL OF REPTILES or fused with the premaxillae in the Mosasauria (Fig. Rhiptoglossa (Figs. 55 d). Primitively (Figs. 23. 72). occurs in some Crocodilia. 2 a. ^^. 31 b. parietals. Parietals (pa). postfrontals. the parietals articulate directly with the squamosals and postorbitals. and some lizards. bral wall. 45. pterodactyls (Fig. in the Chelonia (Fig. Primitively (Figs.) articulating with nasals. articulate directly with the premaxillae (Fig. 3. etc. below with the supraoccipital. 4) articulating with frontals. In the latest Cotylosauria {Procolophon) . and ethmoids. g. postoptics. most Theromorpha . ing the middle of the upper margin of the orbits. Always present and not varying much In the plesiosaurs (Fig. tabulars. most if not all Theromorpha (Figs. and probably absent in some of the former enter into the formation of the nares in the (Fig. maxillae. nasals. tabulars. 72). 69). 4. 50).

and and by a descending process with the palatines. with the palatines and pterygoids in the Crocodiha (Fig. 70 a). articulating with lacrimals. which postnasal and adlacrimal have been proposed. 2. postfrontals. been urged by for Jaekel and Gaupp that these bones are not the homologues of the mammalian lacrimal. they are excluded from the narial margin. 131-135. xlii. 22) at the upper anterior border of the orbits. 4.1 THE OSTEOLOGY OF THE REPTILES (Fig. articulating posteriorly with the postorbitals. — Ed. 30 b). frontals. though much reduced and excluded from the orbital margin in the Theropoda (Fig. from above. 99. Excluded from the frontals in the 1 [The cumulative evidence against the views of Gaupp and Jaekel. S3) and Therapsida (Figs. They are small or vestigial in the Squamata. they articulate with the prevomers in the Chelonia (Fig. They are of extraordinary size in some Theropoda It has (Fig. and all other reptiles. 70 c) they articulate with the postorbitals or postfronto-orbitals frontals are absent as discrete bones. 4. nasals. with regard to the reptilian homologue of the of the mammalian lacrimal. Sometimes (Fig. has been set forth in the Bulletin American Museum of Natural History. vol. Primitively (Figs. and should be called by another name. Pantylus. Prefrontals (pr). 43. 70 a). 45). Three fourths natural size. when the post- Below. 60 a). and absent in most Chelonia and in Sphenodon (Fig.] . 44. 69 d).^ Fig. Never absent. pp.

71. In the absence of the intertemporal and supratemporal. 32 a). and squamosals. p««/y/«. 70 a) in the Theropoda. but not in most other reptiles. Crocodilia (Figs. 50). postorbitals which take their place. jugals. and the snakes 70). 5.1 • 1 /T^' 5. quadratojugal. articulating in front with lacrimals ["Maxillae" — a lapsus calami for "parietal"? Ed. 28. 5. 49 a) help form the anterior boundary of the upper temporal opening. 2.] . many mosasaurs (Fig. the latter also in the Ichthyosauria (Fig. forming the under boundary 1 of the orbits. It extends far and may arand squamosal. 31 b) and most snakes (Fig. 22) at the upper back part of the orbits. Primitively (Figs. 4.f. supporting the paired horns of the Ceratopsia (Fig. Rarely in the lizards (Fig. 22. 59). Postfrontals (pf).THE SKULL OF REPTILES 1 Ichthyosauria (Fig. left . In the absence of the its postfrontal it takes place. 50). part of the temporal roof. 45 b). Jugals ijii). 30 a. They extend forward Postorbitals (po). Thought by some to be the homologues of the mammahan lacrimals.4. the Pterosauria (Figs. b. 2 b. 30. Dinosauria (Fig. 45). 33) large. 70 d). and so called. 59). articulating with postfrontals. 1 . Fig. 72). articulating with prefrontals. articulating with the quadratojugal. 68. jugal. ticulate with frontal. 30. forming a large back in the Chelonia (Figs. 31 69). parietals^ the intertemporal or supratemporals postorbitals. 3. Cotylosaurskull: temporal region. also A descending process articulates with the maxillae or ectopterygoids. 56) it it is does not meet the squamosal. when present. 70). parietal. 50). the postorbital . 2. and with the In the Chelonia (Figs. 43). Primitively (Figs. 54). • also articulates with the parietal (I IgS. 69) large. lizards (Fig. It still retains its connection with the maxillae in certain Chelonia (Fig. frontals. 44 d. Three fourths natural size. 33. . and many Therapsida (Figs. from without. Primitively (Figs. c. 31 a). often extending forward to meet the [sic] ^ prefrontal (Figs. 22) at the upper posterior border of the orbits. It is extensive also in some of the Dinosauria (Fig. they are absent or fused with the Sometimes they (Fig. to the nasals in the Ichthyosauria (Fig. In the Crocodilia (Fig. 56) (Fig. or even the lacrimal (Fig. 30. 46 c. 3.

An amphibian bone known only in Seymouria between the post19) articulating at 19) of the Cotylosauria. 56). Absent in the more specialized Cotylosauria. 4. are probably present in most Theromorpha (Figs. as in the Amphibia. The tabular has been identified by the author as the posterior bone of the arch in the Squamata (p. 59 b) and some lizards (Fig. Absent in some Ophidia {it) (Fig. In the lizards (Fig. In later reptiles the squamosal has undergone many changes. on the inner side perhaps with the ectopterygoids. 62) and Squamata (p. 22) on the dorsal surface of the skull in the Cotylosauria. and articulating in front with jugals and is postorbitals. and maxillae. 33 a). 4. 65 c). above with by an extensive over- lapping suture. Primitively (Figs. 62). 55) they Intertemporals (Figs. Squamosals lars (sq). at the outer side of the interparietals. and. [But see tabular below. Supratemporals {st). 55 a. 44 g). Below. though sometimes vestigial in the Chelonia. side. in the absence of the latter directly with the parietals (Figs. probably all Theromorpha. with the upper end of the quadrates and the outer end of whence the name ''paroccipital plates" given to them by Baur. usually and perhaps correctly called the supratemporal. they cover the posterior part of the temporal region. 53 a). 65). Primitively (Figs. articulating with the squamosals and supra- temporals. . It is unknown in other reptiles.1 THE OSTEOLOGY OF THE REPTILES postorbitals. intercalated frontal. but always present. and some " Pseudosuchia (Fig. extending back of the quadrate to articulate with the pterygoids (Fig. They are known to be absent in but a single genus of Cotylosauria. . 4) articulating with tabu- and supratemporals above. with the squamosals and quadrate jugals. 2. 2. 2. 42 d) and Therapsida (Fig. 2. the paroccipitals. the tabulars behind. 19). overlapping extensively the quadra- tojugals on the sides (Fig. supratemporal. and all other reptiles save possibly the Ichthyosauria (p. the sides of the parietals with the postfrontal and postorbital anteriorly.] Tabulars (/). Generally known as the posterior bone of the arch in the Squamata (Fig. to). may not articulate with the squamosals. parietal. and Ichthyosauria (p. interrupted by the otic and the squamosals on the outer notch in Seymouria (Figs. 33 B. 62). Primitively (Figs. 33 a. 7). and postorbital. 2.

perhaps. Theropoda (Fig. — Ed. Paired bones on the palatal surface. and Squamata. behind with the quadrates. 69). 22. On the other in hand. 40 c). 46 b). Only in the Chelonia (Fig. 42 d). Its relations with the quadratojugal are also inconstant. They (Fig. 30 a). 70 c). some Theromorpha. 52. 69 c) and Predentata (Fig. but is confined chiefly to the posterior side of the quadrate (Fig. as Crocodiha (Fig. dentigerous (Figs. Phytosauria (Fig. and some Predentata c). 63).THE SKULL OF REPTILES 19 Squamata. 69 c). and Theriodontia (Figs. It is absent in most Therapsida. the pterygoids and palatines behind separating the internal nares. 66 b). the head of the quadrate. articu- lating in front narrowly with the jugals. Pterosauria. 70 Dj. 21 b). though sometimes fused in the Rhiptoglossa. articulating with the premaxillae in front. and Ichthyosauria. prevomers single.'' Diaptosauria (Fig. Theropoda. Only in the Cotylosauria and some Theromorpha does it articulate with the pterygoids. Pseudosuchia (Fig. articulating with the postorbitals. are relatively large in the primitive skull. 33 a). region (Figs. (Fig. 55 a). It mandible often In the single-arched skull the quadratojugal tends to is probably present in all Theromorpha. jugal. as most recent authors identify the squamosal. also the case in the Prevomers (pv). are edentulous in all known reptiles except the Cotylosauria 6). it may extend forward to unite with the maxillae some In the Squamata (Fig. 65). p. overlapped by the squamosals. lost in the Crocodilia (Fig. 44 b). called the supratemporal it) and the postorbital (rarely excluded from posterior side of the temporal and usually with the Quadratojugals (gj). of the articular surface for the sometimes forming a part disappear. In later forms it articulates with the paroccipital to a limited extent. it may articulate with the postorbital below as well as above the temporal opening. 3. 2 b. ^ certain Poste- [But see footnote. It enters into the boundary of the lower temporal opening only in the Crocodilia (Fig.^ the Sauropterygia and the Squamata. 44 e). 32 b) are the 6. but not in the more primitive Theromorpha (Fig. At the outer t^t^). "Pseudosuchia.] . supporting In many Therapsida (Figs. It ver>' large in is some Chelonia (Fig. as in all excluded in many Predentata (Fig. it articulates with the bones usually plesiosaurs (Fig. The quadratojugals (Fig. 70 is the other double-arched reptiles. 43 c. 70 a). 43.

31 a.20 riorly in the THE OSTEOLOGY OF THE REPTILES Squamata (Figs. gerous. size. Dentigerous. 21. They may join in the middle in the Chelonia (Figs. Palatines {pi). 32 b) and Plesiosauria (Fig.) Bones of the posterior part with the prevomers in front. 62 e. 55 c. but do articulate with the parietals in many Chelonia. 32 b) and in the Crocodilia (Fig. of the palate. the palatines and ectopterygoids laterally. (Figs. 42 c) and some Therapsida. 6. Cotylosaur skull: from below. the quadrates and squamosals posteriorly. 63 b) and most Squamata (Fig. articulating 40 c. but are lost in other reptiles except the Rhynchocephaha (Figs. More or less denti- Fig. As stated above they do not articulate with the prevomers anteriorly in the Squamata and many Chelonia and Plesiosauria. 56 b) they articulate with the palatines only. Their connection . 54 b). Pantylus. 31 b. 7. the maxillae on their outer. 46 b). and with the descending process of the prefrontals above. Primitively (Figs. Generally believed not to represent the un- paired vomer of the mammals. Pterygoids (pt). the basisphenoids on the inner sides. 69 b) below the prevomers. articulating with the prevomers and pterygoids on their inner sides. 40 c. 6. Three fourths natural Teeth are still present in the Theromorpha (Figs. as also in some Chelonia (Figs. 6. 21) forming the posterior boundary of the internal nares.

in the early reptiles (Figs. trum 47 f). many Chelonia. and are probably generally present in reptiles. and Therapsida. Notho- 47 e). as in the Plesiosauria (Fig. 72 c) of the Pterosauria. (Fig. connecting the pterygoids with the posterior end of the maxillae. is 21 (Fig. where also with the jugals in the they pass above the palatines to unite with the pterygoid.] . sometimes Squamata (Fig.^ Ecto pterygoids (ec). 55 c). Theromorpha. and Squamata.THE SKULL OF REPTILES with the prevomers Rhiptoglossa. Pantylus. Rhynchocephalia." are 24 c. Enlarged one half. though perhaps present. In some Therapsida sauria (Fig. though absent in the Crocodilia. The ectopterygoids are believed by some to be the homologues of the Epipterygoids (ep). various genera of the Cotylosauria. etc. They have been ob- served in the Temnospondyli. as in most modern but are fixed in the Therapsida (Fig. lost in some Cynodontia 43 c) and Teeth are generally present in the Theromorpha. with section of mandible through condyle. above with the parietals (Fig. they unite along the middle line. — Ed. 6. There is an interpterythem partly filled with the parasphenoid ros(Fig. Rhynchocephalia. Placodontia whole 49 b). 21 a). most Squamata (Fig. 43 c). the ' [Watson has shown that this view is untenable. and in some Therapsida and ''Pseudosuchia. A pair of bones articulating below with the pterygoids. Fig. 40 c) they more or less loosely articulated with the basipterygoid process of reptiles. There is a pterygoids are -4A theory that the the real homologues of the mamecto- malian alisphenoids. In the early reptiles (Figs. pterygoid process of the alisphenoid of the mammals. 6. 21. 8). back of the palatines. quadrate region. they are certainly absent in some of the left Temnospondyli. 43 c) goidal space between and not a few others. the basisphenoid. The pterygoids (transpalatines) have not yet been certainly demon- strated in the early Cotylosauria. cept the Ichthyosauria and Chelonia. recognized in all They have been other orders ex- Cotylosaur skull: 7. 55 c). Most remarkable are their relations in Pteranodon (Fig.

articulating with the parietals and interparietals above. the exoccipitals.22 THE OSTEOLOGY OF THE REPTILES In the Ophidia. 30 b) they have been identified with a plate of bone and the basisphenoid of many forms. Cotylosaur skull: A. (Figs. Cotylosaur skull: from behind. right quadrate from Lalidosaurus hamatus Cox>e. Rhiptoglossa of the Squamata. ful. Their real homologues are yet doubt- FiG. C. 21 b. 42 d. the same from above. paroccipitals. Amphisbaena. 9. by some they have been mammals. but which has never been demonstrated in any air-breathing . Fig. posterior basicranial bones from above. 9.) Unpaired. Labidosaurus hamatus Cope. and including a part of the semicircular canals formerly believed to be in a separate bone called the epiotic. intercalated between the descending plate of the parietal Chelonia (Fig. 8. and prootics. identified with the alisphenoid of the Supraoccipital {so). B. below.

occipital and quadrates. reptiles. with the supraand prootics. intercalated between the supraoccipital. ninth. 30. as in the Chelonia (Fig. forming the larger part of the boundary of the foramen magnum. op). and so-called supratemporal. the double condyles of the Cynodontia and mammals. 10. (Figs. 51). it by the exIn most less into Ophidia and Crocodilia. supraoccipital. Primitively (Figs. tabulars. and. The prootics (Figs. Exoccipitals (eo). 59. basioccipital.) tively Only in the Cotylosauria primi- do the paroccipitals exist as distinct bones in the adult. the postoptics. Among stapes. enters more or the boundary of the foramen magnum. and supporting the head of the quadrate. so far as known. 56 b) . On the inner side they help form. exoccipitals. modern they separate bones in the adult. or even the whole. closely articulated with the to each other Primitively the exoccipitals took but little basioccipital only. 31 b). Prootics (pc). only of with the exoccipitals. the otic capsule. squamosal. 21 b. Paroccipitals (po). approximated both above and below. part in the formation of the occipital condyle. 11. of hearing. suturally or loosely articulated (Fig. and containing a part of the internal Their relations are yet poorly known in the primitive for the pas- They usually have foramina perforating them sage of the third and sixth nerves. In the Theromorpha. articu- lating with the exoccipitals. other reptiles they are known to be free only in the Ichthythey are indis- osauria (Fig. are In the Chelonia supraoccipital. They form a large part of the brain-case exteriorly in . and tenth nerves. 21 b. the paroccipitals are fused with the supraoccipital. when organ present. posteriorly for the eighth. tension of the large foramen reptiles save the magnum to the parietal roof. but in many later reptiles they form a large part. as in the Am- phisbaenia (Fig. paroccipitals. 31 B.THE SKULL OF REPTILES animal. In other reptiles tinguishably fused with the exoccipitals in the adult. articulating with the basioccipital-. or. prootics. prootics. 23 Primitively more or less excluded from the margin of the foramen magnum. 69) are a con- spicuous part of the brain-case. Only in certain plesiosaurs is the supraoccipital paired. by the recession of the basioccipital. 9. 8. 42 d) small. intercalated between the exoccipitals. stapes. and form' the posterior boundary of the foramen for the fifth nerve. reptiles. basisphenoid.

al). articulating with the so-called supratemporals. D. po. ps. 59) and amphisbaenian lizards. paroccipital. bo. Edaphosaurus. basioccipital. the tabulars. 57) their outer extremity extends to the outer extremity of the Fig. stapes. the same from side. parasphenoid. otosphenoids. paroccipital. Temnospondyl skull: basicranial bones. exoccipital. B. prootic. from without and within. A. F. basisphenoid. E. longitudinal and transverse sections. H. as the author believes. basisphenoid. the same from above. genus indet. from below. so. apparently not . natural size. Posto plies (as. (laterosphenoids. ep. Eryops. G. eo. epipterygoid. C. " alisphenoids"). Variable and yet doubtful bones in the reptiles. in the mosasaurs (Fig. 10. or. supraoccipital. Theromorph skull: occipital complex. st. natural size. pc. bs.24 THE OSTEOLOGY OF THE REPTH^ES the snakes (Fig.

They form the lateral brain-case in the Crocodilia. F. B. Fig. 25 mammaKan alisphenoid. the same from in front. Eryops. Eryops.THE SKULL OF REPTILES homologous with the called. A. articulating as in the Crocodilia (Fig. specimens. They are present in and snakes. G. postoptics given to back of the optic foramen. they have been recog- nized in the Temnospondyli. section through parasphenoidal rostrum. 69) below with the basisphenoid. section through parasphenoidal rostrum near base. above with the is parietal. II. B. the same from above. but are absent in the Chelonia lizards they are imperfectly ossified. D. Pelycosaur skull: occipital complex: A. D. In the and are usually lost in prepared the Rhynchocephalia and most other reptiles. Between them and the prootics the foramen for the fifth nerve. . section through parasphenoidal rostrum at middle of orbit. though usually so Imperfectly known in the early reptiles. Dimetrodon. whence the name them by Cope. and Theromorpha. obliquely from behind. C. C. Temno- ' spondyl skulls: E. near front part of orbit. Cotylosauria. Cacops. from below.

) (ps). 69 d). C. — G. 56). but in most other without a perforating foramen. Paras phenoid 21 A. outline of same at extreme front end of parietals. which closely fused with its is under side. It is perforated near its proximal end by the foramen perforating artery. B. d. between the paroccipital and basisphenoid. 12. 30. F. or foramen ovale. the Ichthyosauria It is for a also large in and Plesiosauria. This seems to be the rule in the early reptiles.^ It extends far forward as a narrow rostrum in the internal cranial bones. prevomer. basioccipital.) Forming the floor of the brain-case in front of the basioccipital. 69 d). Parity lus. basisphenoid. pituitary front the fossa or depression for the body (st). ar- ticulating in front above with the postoptics (Fig. 11 c. Parity lus.) is The all early reptiles a large and mandible: immature from below. firmly fused in under side and Cotylosaur skull: Natural size. ii d. 6. 30. II c). though in ' some {Labidosaurus sp) it [It is separate in at least some geckos. section of rhinencephalic chamber opposite hind margin of orbits. K. bone. section at front of basisphenoid. and externally below It lodges in with the pterygoids. d. and extending toward or touching the quadrate. 12). E. Natural size. 6. temnospondyls (Fig. It is stout and short in the Amphisbaenia (Fig. articulating over the auditory opening. 690). A. (Figs. continuous with the para- sphenoid in front (Fig. pal. behind above with the prootics (Figs. 13. D. Fig. A membranous. Stapes stapes in Fig. Basisphenoid (Figs. (Fig.] . lo. Cotylosaur skull animal. forming the floor to the ethmoidal cavity. 1 1 e-g) and some a) quite to the cotylosaurs (Figs. the adult with the of the basisphenoid. unpaired bone. (Fig. 10 c.26 THE OSTEOLOGY OF THE REPTILES (bs). never a separate bone. externally above with the stapes (Fig. N. same in front of orbits. 21 prevomers. reptiles is slender.

differing from that of their temnospon- dyl antecedents (Figs. Transverse. II e-g). opening into the nasal and paranasal cavities Similar bones have been observed in various in front of the orbits. All of these. Otosphenoid. = Petrosal. There was no median ossified interorbital septum in these reptiles. 13) and theromorphs. In the temnospondyl amphibians between the orbits and in front of the optic foramina. Skull Elements Ectopterygoid Interparietal — Synonyms Postoptic Paroccipital = = = = Transpalatine. they have no immediate relation to the sphenoid. though never in the same reptile. cotylosaurs (Fig. or Sphencthmoid. 14. Dermooccipital. possibly nine. It has been homolo- by Broom with the median vomer {eth) of mammals. the precoronoid and intercoronoid. "Alisphenoid. = Goniale. the accompanying [Possibly "paroccipital plate" was intended. They have been called sphenethmoids." Opisthotic. in and some but five. there is a pair of thin bones lying closely below the frontals and united with the parasphenoid below. — Ed. and are probably constant among early reptiles. Squamosal Tabular = "Epiotic.THE SKULL OF REPTILES may gized 2 J not extend in front of the basisphenoid. Postsplenial = Opercular. have persisted to all modern times. the so-called vomers. except the postsplenial. the paired bones in front. 15) only in the loss of one or two slender bones along the inner margin of the teeth. and not probably any median septum. = Prosquamosal. separate bones. Postparietal. 18). Laterosphenoid. The relations of ^ none having more than these bones will be seen six. known in a single cotylosaur (Fig. Splenial Supratemporal = Supramastoid. Prearticular = Preangular. whence the name prevomers for Ethmoid (Fig. A cartilaginous interorbital septum is present in most modern reptiles but is ossified in none. Dermosupraoccipital." Postparietal [sic] ^ Prootic The Mandible The mandible of reptiles was composed primitively of eight.] . though Probably the median ethmoidal plate arose from the fusion of these bones. enclosing an undivided cavity for the olfactory lobes. Suprasquamosal.

skull and mandible. psp. intercoronoid. etc. and the late Pterosauria (Fig. The den- tary (d) is always present and dentigerous. some Anomodontia (Fig. E. Temnospondyl skull and mandible: A. Saphaeosaurus. extending along the inner margin of the teeth to the hind from near the symphysis end of the dentary on the inner Trimerorhachis alleni Case. In the Dinosauria. {e. or some of them at least. cor. the intercoronoid . and may in some cases be In the Plesiosauria (Fig. postsplenial. 18). Fig. has been restricted to a place at the posterior end of the dentary in later forms absent. icor. e). Fig. 55 b). C. some Theropoda. right mansame.. B. inner side. may perhaps be a distinct bone in front. even entering into the symphysis in some cases. left side. g. F. dible. or there it also extends far forward. 31 B.28 figures (1 THE OSTEOLOGY OF THE REPTILES 6-1 8) and do not require a detailed description. outer side. D. coronoid. The coronoid {cor). 25 a) it still retains its ancient character. 44 c) and Dromasauria. sections of mandible as designated. pcor. precoronoid. except in all Chelonia (Fig. 71 e). 14. possibly composed of two bones in some of the earliest reptiles (Fig. the side.

B. The angular {an). on the inferior border posteriorly. articular. c. forms the upper margin mandible back of the coronoid. the prearticular. articulating with the dentary in front. though sometimes indistinguishably fused with the articular. many temnospondyls. as in in later reptiles. MeckeHan orifice (Figs. In some it may take part in the articular surface for the quadrate. and extending to the hind angle of the jaw. 29 Primitively (Fig. right ramus from below. and the outer margin of the Fig. but no such teeth are known The surangular of the (sa). and surangular behind.THE SKULL OF REPTILES or precoronoid. d) it helps form the inner . Trimerorhachis insignis. (Figs. 69 border of the Meckelian orifice. 15-18). 15-18). is always present In the crocodiles (Fig. 18) it often bore teeth. the same from above. Temnospondyl mandible: A. always present in reptiles. 15.

. Pelycosaur: left mandible. Pelycosaur: right mandible. E. outer side. A. Labidosaurus hamatus incisivus Cope. inner side. Dimetrodon D. Cotylosaur: right mandible. C. Cotylosaur: right mandible. outer side. Fig.B. Labidosaurus hamatus Cope. Dimetrodon incisivus Cope. Dimetrodon incisivus Cope. inner side. Cope. Pelycosaur: left mandible. i6. inner side.

15-18). Diadectes {Nothodon). One half natural size. entering into the tively (Figs. from within. extending 31 Meckelian symphysis primiMeckel- back to the posterior inferior ian foramen. In Pantylus (Fig. and angular. Fig. B. so far as known. Pantylus. sections corresponding to letters. prearticular.THE SKULL OF REPTILES The splenial (sp). 18) only. C. right mandible from inner side. Cotylosaur: A. 17. the postsplenial corresponds to its posterior part as in the known Stegocephalia (Fig. is not infrequently absent. 15). Fig. It [the splenial] never bears . the same from below. the same from above. D. 18. articulating with the dentary. Cotylosaur: left mandible. coronoid.

and dentary. and doubtless of the posterior inferior ian orifice. 60). but in all late reptiles it restricted to the inner side. 16-18). 15-18) it name opercular often given to it. ensheathed by the united angular and coronoid. only cartilage bone of the mandible. in whole or part. is still present in the crocodilians (Fig. Plesiosauria (Fig. 25 a). appearing is on its outer face. It is fused with the articular in the Squamata extending far forward in the mosasaurs (Fig. for the passage of a track of the chorda tympani nerve. and splenial. It many 58). It is a thin bone and forms the cover to the Meckelian groove. between the angular. 69 c). between the prearticular and angular. 69). It was present in the Dinosaurs. and remains a separate bone in the modern turtles (Fig. is very characteristic of most Crocodiha (Fig. there are in the early reptiles one or two smaller openings through the inner wall: the posterior one just in front of and below the anterior end of the large orifice. splenial and is dentary. probably an acquired character. Ichthyosauria. 70 a). 31 e) and young Sphenodon (Fig. Openings in the mandible Aside from the large opening for the entrance of nerves and bloodvessels [and dible (Figs. e). prearticular. (Fig. the coronoid and splenial in front. Primitively formed a large part of the inferior border of the man- dible anteriorly. angular. between the coronoid. recognized only within recent years. is a thin bone. and Pseudosuchia (Fig. It enters into the mandibular sym- physis in most long-jawed reptiles. the apparently wholly absent in the Crocodiha (Fig. Theropoda (Fig. Believed to be the malleus of the mam- malian ear.32 teeth and is THE OSTEOLOGY OF THE REPTU^ES more or less inconstant. articulating with the articular behind. 55 b). is . 69 d). the angular below. It The articular. jaw muscles] at the posterior upper part of the man16-18). may be indistinguishably fused with the pre- articular or surangular. A foramen posteriorly. Phytosauria. where it was first named. it early (Figs. absent in Sphenodon and most Chelonia. a large perforation of the outer wall of the mandible. other extinct reptiles. 65 a. surangular. 16-18) and many later reptiles. The prearticular (goniale) {pa). forms the Distinct in all cotylus. forming the hind border foramen and the lower margin of the Meckelwas present in all early reptiles (Figs. whence the (Figs.

poral. pj. The parietal foramen is absent in one or two . maxilla. 20. supratemporal. 17-24. E.THE SKULL OF REPTILES The Skull of the Cotylosauria (Figs. Seymouria haylorensis. jugal. qj. prefrontal. do. 13. St. quadratojugal. /. clavicle. sq. j. One ic. 25 B. dermoocci- pital. One half natural size. postfrontal. 26-29) 33 CotyloFig. /. 19. po. I-Q. «. C. 16 D. pf. nasal. /r. frontal. m. clavicle. lacrimal. cl. prefrontal. half natural size. c. /'/. pm. tabulate. intertempqf. premaxilla. sc. Fig. coracoid. Seymouria haylorensis. Skull and pectoral girdle: from the side. n. scapula. saur skull: from above. 12. postorbital. lacrimal. /. inter- But few modifications of the primitive characters of the skull are known in this order. nasal. squamosal.

in the Diadectidae the posterior ones are transversely molariform. known The inter- not reaching the nares. in some genera they are obtuse and cuspidate. postsplenial occurs in but one genus [Pantylus]. The lacrimal in Procolophon is small. in no known forms are they sectorial. Teeth occur on the coronoids. parietals are reduced and posteriorly placed in a few. Doubtless with future discoveries other modifications of the primitive structure will be found. . the supratemporals and tabulars in a few others. The teeth are usually conical throughout. as also the intertemporal [Seymouria]. and in the latest genera may be vestigial.34 THE OSTEOLOGY OF THE REPTILES may The genera.

After Watson.Qu. from below. QuJ. B. occipital view. Two thirds natural size. Cotylosaur skull: Seymouria. A. Fig. 21. 35 .0cfEN. Pf ftR0cFxOcB.0v. Two thirds natural size.

One half natural size. * ". •' y -J.v'-^ 36 .^'-3 •*. Cotylosaur skull: Diadecles^ from the side and above.Fig. 22. .

jugal. 23. yy. lacrimal. y. pqf. Cotylosaur skull: A. from above. nasa]. po. /. angular. pf. ang. 37 . from the side. sur. /. pm. B. pa. «. dentary. premaxilla. do. q. postfrontal. parietal. j^. maxilla. d. dermooccipital.Fig. quadrate. squamosal. surangular. /. Limnoscelis paludis. quadrato- jugal. tabulare. prefrontal. m. frontal. postorbital.

pa. st. Limnoscelis paludis. stapes Two (?). fifths 38 . splenial.Fig. sp. Cotylosaur skull: from below. 24. natural size. prearticular.

surangular. prearticular. 27 Fig. Two fifths natural Fig. Trinacromerum osborni Williston. Cotylosaur: left mandible. A. after Williston. coracoid. from within. One half natural size. sp. Limnoscelis paludis. B. pa. size. 25. Plesiosaur: left mandible to symphysis. Labidosaurus hamatus. 9. 39 . ang. Limnoscelis paludis. angular. Cotylosaur. A. 26. J«r. B. quadrate. articular. the same. Cotylosaur. C. 25 Fig. «r/. inner side. Fig. Captorhinus agtiti Cope. 1903. Inner side of mandibles. splenial. 27. Outline of back of skull. cor.Fig. outer side.

Cotylosaur skull: from above. 40 . Two thirds natural size.Fig. Labidosaurus hamatus Cope. 28.

an. st. ep. a. articular. postparietal. angular. from below. from the side. pt. stapes. exoccipital. bs. quadrate. q. pp. 41 . B. epiotic.v. Labidosaurus hamatus Cope. 29. Two thirds natural size. basisphenoid. I'lG. pterygoid. ^.B Cotylosaur skull: A.

Fig. Pleurodiran skull: Macrochelys. 42 . One half natural size. from the side and hemisection. 30.

from the inner side. size.Cryptodiran skull. One half natural size. from the side. Cryptodiran skull. D. from below. from above. C. One half E.One Fig. 31. Colpochelys. Three halves natural natural size. B. occiput. the same. 43 . half natural size. Pleurodiran skull: Podocnemis. A. Trionychoid mandible Piatypeltis. T/w/^jjofA^/yj.

and the parietal men. The prefrontals are large. The palatines often meet in the middle line between the pre- . The parain certain sphenoid has been forms. proatlas.44 THE OSTEOLOGY OF THE REPTILES The Skull of the Chelonia (Figs. postfrontals. 32. The nasals are usually absent. except in Dermo- by descending plates from the parietals to the pterygoids. meeting in The prevomers are single and sometimes obsolete. is lately recognized as a distinct its bone no postoptic. from above. C. though kinds. It relatively slender in some of the more predaceous always fora- lacks the septomaxillae. but place is taken. The quadrate may or may not reach the basisphenoid. however. Trionychoid skull: Platypeltis. the middle line. ethmoids. dermosupraoccipitals. A. Natural size. There chelys. from below. tabulars. Note abnormal only in some ancient forms. sometimes with an intercalated epipterygoid. atlas of same from the side. B. supratemporals. and the lacrimals are present Fig. ectopterygoids. they usually articulate above with the prefrontals. is usually absent. 30-32) The skull of the Testudinata or Chelonia is never elongate. which.

It remains cartilaginous in the Dermochelyidae. as in some cotylosaurs. . The stapes is slender. Both upper and lower jaws are encased in a cutting horny sheath.THE SKULL OF REPTILES vomer and the pterygoids. The mandibles have a large. usually but incorin the terapenes. the splenial is rarely present (Emydura. roofed over. rectly called the splenial. and are without teeth. 45 The pterygoids also usually meet in the middle. forming a rudimentary secondary palate. quadratojugal. The paroccipital remains free throughout life. of the roof sals Usually from behind or below. it squamosal. by the large postorbital. in some wholly so. a Triassic genus. The palatines also often meet for a short distance below and in front of the internal nares. etc. and yet is. though separated in the Trionychoidea by the basisphenoid. and is more or less exposed by the emargination sides. The temporal region primitively was wholly in some marine turtles. bones are known to occur only in Stegochelys. Small teeth on the pulatal Toxochelys. The condyle is largely formed by the exoccipitals.) . its ear-cavity sometimes wholly surrounded by bone. or from both and the squamo- and quadratojugals may even become vestigial in the process as The quadrate is always large. free prearticular.

or compressed and sectorial. often markedly anisodont. lateral view. quadratojugal. 32-42) More modifications of the skull structure are found in this order than in the Cotylosauria. conical. ^/. postorbital. bounded by the jugal. obtuse. as would be expected. and they are always confined to the occipital surface when present. l6 A-C. There is a lower temporal opening. the former usually. . postorbital. The lacrimals seldom extend to the nares. The supratemporals are doubtfully present in any. 9. /)«. /)o. unpaired. parietal. Fig. >. lacrimal.prefrontal. In the Edaphosauridae only may it possibly extend to the parietal. The teeth of jaws and mandibles are more variable. and squamosal. The quadratojugal is smaller and never extends far forward. 32 its. The teeth are often want- ing on the prevomers and are sometimes present on the coronoids.46 THE OSTEOLOGY OF THE REPTILES The Skull of the Theromorpha (Figs. Theromorph skull: Op/iiacotio>i mirus Marsh. II A-D. /. lO A-D.jugal. quadrate. yj. The interparietals and tabulars are probably sometimes absent. if not always.

from above. A. 47 . Theromorph skull: Sphenacodon. 33. B. from the side.Fig. One third natural size.

35. 34. 36. Fig. Natural size. Theromorph skull: Varanosaurus brevirostris Williston. from the side. from the side. Two thirds natural . Theromorph skull: Glaucosaiirus megalops. Theromorph skull: Mycterosaurus.Fig. Fig. Natural size. from the side and from above. size.

Fig. 39. 37. After Broili. from above. from above. Theromorph Theromorph skull: Varanosaurus acutirostris Mycterosaurus. size. skull: Fig. Two Two thirds natural thirds natural Fig. Natural 49 . Fig. Broili. size. 37 Fig. from above. size. 38 Theromorph skull: Varanosaurus brevirostris Williston. 38.

the same from below. from the side. B. Naosaurus daviger. 40. Two fifths natural size. C. SO . skull from above. D. A.Fig. from inner side. Theromorph skull: right mandible.

D. po. from the side. quadrate. surangular. exoccipital. postorbital. palatine. 42. prearticular. skull. sa. splenial. pa. /)w. dentary. basisphenoid. pt. ds. ang. from behind. c. d. from above. Three fifths natural size. angular. Natural size. C. Fig. from below.Fig. pa. maxilla. Casea broilii Williston: A. sp. stapes. dermo- supraoccipital. from inner side. 41. Theromorph skull: Casea broilii Williston. eo. q. paroccipital. poc. prew. coronoid. basioccipital. art. bo. pterygoid. bs. 51 . skull. B. j. articular. maxilla. left mandible of same. so. supraoccipital. st. jugal.

Only in some of the is essentially dicondylar. The parietals may be united in some of the Bauriasauria. The parietal foramen is usually absent. a well-formed secondary of the maxillae in palate in the Bauriasauria and Cynodontia. In the Cynodontia the condyle be absent or present. The supratemporals are never present. it is generally present. Some Dromasauria and the females of some Anomodontia are Other anomodonts may have a single caniniform tooth in each jaw. The pterygoids and palatines meet in the middle line in the Dinocephalia. or canines and molars. 43-45) Many more modifications of the skull are found among the reptiles grouped under the name Therapsida or Anomodontia sens.the postorbitals and squamosals not meeting. There is a separate bone in front or surrounding the parietal foramen in the small. edentulous. There is a partial false or front of the nares in the secondary palate formed by the union Anomodontia. the molars sectorial or cuspidate. Sollas. The pterygoids do not meet the small quadrates in the Cynodontia. and Cynodontia. The postf rontals are often absent the quadratojugals are present only in the Dinocephalia and are Dromasauria do the lacrimals and septomaxillae exclude the maxillae from union with the nasals. reaches the parietal in theTherocephaha andTheriodontia. Bauriasauria. The Therocephaha have anisodont sectorial teeth. The ectopterygoids may Therocephaha are there teeth on the palatal bones. as also the tabulars. The vomers are fused into a single bone in the Gorgonopsia. — Ed. The interparietal or dermosupraoccipital is always on the occipital surface of the bone and is unpaired. 1 [Broom. Watson. the Cynodontia with real heterodont dentition. usually bounded above as in the Theromorpha. lat.] .52 THE OSTEOLOGY OF THE REPTILES The Skull of the Therapsida (Figs. and von Huene have observed a distinct quadratojugal in various Therapsida. ^ Only in some of the Anomodontia and Gorgonopsia. The temporal foramen.

from the size. After Broom.Fig. 53 . side. from below. from above. 43. One half natural A. Gorgonopsian skull: Scylacops capensis. C. B.

54 .Fig. occiput. Watson. B. Cistecephalus (Anomodontia). One half natural size. Scylacosaurus below. Dicynodon (Anomodontia) from the side. One half natural size. After Watson. from the side. Mormosauriis (Dinocephalia) from above and from side. Therapsid skulls: A. After Broom. Two sevenths After Broom. E. G. One fourth natural size. 44. C. from F. D. One half natural size. Gorgonops (Gorgonopsia). One twelfth natural size. After Broom. from above. (Therocephalia). After natural size. After Watson. Diademodon (Cynodontia).

Cynognathus crateronotus. 45. upper teeth. One third natural size. 55 . After Broom. the same. Cynognathus platyceps (CynoAfter Broom. dontia).Therapsid skulls: A. D. from the side. One half natural size. from above. Bauria (Bauriasauria). from above. B. C. from the side. Fig.

as the front part of the orbit. 4^-48. The brain-case is more or less open in front on the sides. dividing . sometimes very slender in the short-necked types. There is a pineal foramen. join each other back of the parietals on the upper surface of the skull. The bounded below by the postorbitals which are more or less elemiddle in a crest. which may The paroccipitals are always fused with the exoccipitals.56 THE OSTEOLOGY OF THE REPTILES The Skull of the Nothosauria (Fig. the bone. the postoptics either reduced or absent. bounded below by the jugals and maxillae. above by the parietals. The orbits The postfrontals are probably present in all. or supratemporals. The stapes is large and stout. extending back at least as far slender. is border also [short?] shut. extend to the roof. in some at least. 47) [No manuscript. tabulars. The vated in the squamosals. which extend as far forward as the external nares. 246. There are no interparietals. they appear to be absent in some. The nasals have never been certainly determined. 25 a) The extreme aquatic adaptations certain modifications in the structure skull that are unique of the plesiosaurs have caused and relations of the bones of the among reptiles. On less the occipital surface the supraoccipital is excavated more or by the foramen magnum. There are no teeth in the palate. above or between the frontals. in the long-faced kinds even to articulate with the parietals. possibly they are fused with the frontals. always shorter in the long-necked forms. The general shape of the skull seems to bear a definite relation to the length of the neck. the lacrimals have been identified in a few forms only. 247] The Skull of the Plesiosauria (Figs. The premaxillae are always greatly elongated. The are all quadratojugals are conceded to be absent in single large temporal opening is plesiosaurs. and squamosal. See pages 211. forming the inner border. The maxillae are The alveolar much more elongate than in the icthyosaurs and phytosaurs. The prefrontals are small.

C. One sixth One sixth After Andrews. Muraenosaurus. fourth natural size. 57 The anterior nares are situated either between the prevomers and the maxillae. B. B. from below. After Jaekel. 46. After Fraas. One size. . One fourth natural size. Sauropterygian skulls: A. One size. C. Nothosaurus. Sauropterygian skulls: A. After Andrews. Fig. natural size. 47. After Linder. Simosaurus. Plesiosaurus. Thaumatosaurus. Peloneustes. One fourth natural from below. Fig. from above.THE SKULL OF REPTILES Very great also. are the modifications of the palate. ninth natural natural size.

y. ternal nares are in front. are always a remarkable posterior interpterygoidal vacuity.58 or between small. angular. There Fig. as also posterior palatine and in- other openings in the palate. from the side: ang. C. Plesiosaurus. may or may not be present. extending along the sides of the teeth internally and meeting each other in some forms in a median symphysis. po. sixth natural size. sttr. THE OSTEOLOGY OF THE REPTILES them and the is palatines. tary. Elasmosaurus. splenials As usual in long-faced forms. 48. hke the external ones. An anterior interpterygoidal vacuity. postorbital. The sometimes very much in front. po. surangular. maxilla. Trinacromerum. divided in the middle throughout by the large parasphenoid. from the side. jugal. postorbital. premaxilla. Plesiosaur skulls: A. from the s\Ae: pm. the meet in a median symphysis. d. One B. ternal nares. the pterygoids meeting in front of and to a slight extent behind them. of the ex- The coronoids are elongate bones. and. pm. jugal. . _/. denw?. premaxilla.

After Broili. Placodus. Fig. flat. Placochelys. 49. Placodont skulls: A-D.49) 59 The skull of the Placodontia is almost unique among reptiles for crushing teeth upon the the extraordinary development of large.THE SKULL OF REPTILES The Skull of the Placodontl\ (Fig. . After Jaekel. One fourth natural E. One half natural size.

postfrontal. jugal. possibly is the prevomers are also single. unlike those of other aquatic reptiles.6o THE OSTEOLOGY OF THE REPTILES all told. the palatines are very large. In consequence. The very large orbits have the usual bound- ing bones. and lacrimal. articulating in front not only with the whole extent of the frontals but also with the nasals and prefrontals. the maxillae are short. posteriorly with the so-called supratcmporals. bears numerous teeth. The frontals are small. tabulars have been found. and possibly has arisen is in the bar below the opening identified Broili cannot find. in Placodus as few as twenty chelys still in Placo- fewer. has been highly and peculiarly modified in many ways. which cephalia. it is seen. is broadly separated above by the very large nasal. jaws and palatines. Except for the postfrontals. 50. are very small to the posterior part of and the pterygoids are restricted the palate. and All bones are paired. 51) The The skull of the ichthyosaurs. both views are probably incorrect. which is in Placodus the more probable. The postorbitals are long bones forming nearly the whole posterior . teeth. widely separated from the pre- The massive cranium has a large temporal opening bounded above by the parietal. but their relations are somewhat changed. So. The largest skulls of Placodus are about ten inches long. the structure here. ing each other throughout as do the pterygoids in the median The ectopterygoids vomers. with the postfrontal entering into the anterior border. also. below by the united postorbitals and squamosal. while retaining not a few primitive characters. postorbital. meetline. prefrontal. There a large epipterygoid. The stout lateral by Jaekel in Plachochelys as composed of the squamosal and quadratojugal. is The nasal only of the roof bones unpaired in Placodus. greatly elongated premaxilla. is like that of the Dino- same way. the postfrontals are extraordinarily large. The Skull of the Ichthyosauria (Figs. by Huene as the supratemporal and squamosal. Huene believes there is an interparietal. No each with three incisor-like The premaxillae in Placodus are large. The prefrontals are long. since Broili finds only the squamosal.

foramen is at the front sometimes be- tween the frontals. more probably a primitive. The large end of the parietals. is Most characteristic of the ichthyosaur skull the structure of the temporal region. . 50. quadratojugals. and lacrimals. about which there has been dispute from the time of . side. maxillae. unlike all other reptiles since The is the primitive Cotylosauria. from bones. On the occiput the paroccipitals. character. an ectopterygoid has not been recognized and probably absent. stout bone. Ichthyosaur skull: Eaptanodon {Ophthalmosaurus) from the above. After Gilmore. 6i The jugals are long. stapes a short. save the Chelonia. possibly an acquired. articulating with postorbitals. parietal There are no dermosupraoccipitals. there are no teeth on the palatal Fig. are separate. and from below. with their usual articulations. The is relations of the bones of the palate and the boundaries of the nares are primitive.THE SKULL OF REPTILES boundary of the orbits.

/r. was re- tained in any reptiles after the If. Owen The large temporal vacuity is admittedly the upper one. bs. The upper. was later tive skull. or tabular. from the rear. and doubtless is recognized in the skull of the Squamata. and closed it is quite improbable that a preexistent one orbit. posterior bone completing the upper border of the tem- poral vacuity. frontal. there can be no question less And un- we accept the wholly improbable theory that new bones have been developed in the temporal region of the ichthyosaurs. of identity. and quadrate. the other two must be homologized with the supratemporal. angular. must be the squamosal. basisphenoid. and quadratojugal. having as it does the same relations with in the paroccipital. as a com- parison of the cotylosaur skull will make evident. rarely mosasaurs extend- ing forward to articulate with the postorbital. d. The outer bone. dentary. as some recent writers assert. the oldest cer- . as in the primi- three. by the encroachment of the About This region. postorbital. however. the supratemporal was persistent '"g in the ichthyosaurs instead of the tabular. 53 c-e) In the order here provisionally called the Protorosauria the skull is completely known in none. and there is no certain evidence yet that it forthcoming cotylosaurs. the postfrontal. There is no lateral foramen. parietal. After Gilmore. on the outer by the postfrontal and the so-called supratemporal. 51. Ang. The Skull of the Protorosauria (Figs. bounded on the inner side by the parietal. sometimes obsolete in ichthyosaurs. has five bones. by no possibility can Ichthyosaur skull: Baptanodon {Ophthalmosaurus). The supratemporal bone was the first to be lost in the primitive skull.62 THE OSTEOLOGY OF THE REPTILES to the present. it be the bone on the outer side of the squamosal. the author prefers to believe is the tabular and not homologous with the bone so the supratemporal. but best in Araeoscelis. and the squamosal. 52. We cannot conceive of its being anything else. FiG.

In Araeoscelis. and ^rohahXy Protorosaurus there . is a parietal foramen. of the reptilian skull. Araeoscelis. Twice natural size. In is Araeoscelis the temporal opening bounded externally by three Fig. and two bones posteriorly. Aside from the tabular. size. Plenrosaurus. so far as known. The lacrimal is small or vestigial in The ably postfrontal is present in Araeoscelis. and quadratojugal have been applied in almost all possible combinations. supratemporal. responding bones in the lizard which have been the subject of more controversy than any others poral region. Pleurosaurus and the Squamata. and only in this genus are there indications of the presence of the dermosupraoccipitals. from the side. Pleurosaurus. the postorbital in front. all. supramastoid. to . Natural B. from the side. Only two of these are present in Araeoscelis. Proball have teeth on the palatal bones. 52. about which there is doubt because of their evident identity with the corskull. to which the names mastoid. but none in Saphaeosaurus. roof bones are all paired in all. lies in Their chief interest the structure of the temporal region. there are three recognized bones of the primitive temall present in the Cotylosauria and Ichthyosauria. squamosal. prosquamosal. bones. temporal. Parapsid skulls: A. suprasquamosal.THE SKULL OF REPTILES The 63 tainly-known reptile with a single typical upper temporal vacuity.

from above. Mycterosaurus (Theromorpha). size. calling the posterior . B. from the side and from above. Natural size. from Natural Squamata arch is the supratemporal. however. some.64 which all THE OSTEOLOGY OF THE REPTILES of these names have been given by is different authors. The and more general opinion that the posterior one of the Araeoscelis Fig. etc. 53. D. reverse these names.. E. One half natural size. C. from Natural size. skulls: A. Araeoscelis (Protorosauria). Sauranodon (Protorosauria). the anterior and outer one the squamosal. Pleurosaurus (Protorosauria). above. above. Parapsid.

the true supratemporal. pres- ence has never been positively determined in the Theromorpha and however. paroccipital. in the Ichthyosauria (and a distinct bone between it Saphaeosaurus) there at present is and the temporal opening that must be either the squamosal or supratemporal. however. or the quadra- tojugals. seems improbable. It is a fact. There is no certain solution of the problem. The exoccipitals and paroccipitals . less of present in only a very few of the Therapsida as more or a vestige.THE SKULL OF REPTILES one the supratemporal or quadratojugal. temporal have disappeared in these and that the squamosal has usurped their position and functions. the an- one the squamosal. however. streptostylic quadrate. the prosquamosal. all single-arched reptiles otherwise. The Skull of the Squamata (Figs. are always fused parietals the pterygoids never reach the vomers the inter- and either the supratemporals or tabulars. secondarily or less fixed in . and Therapsida. 54-59) The more skull of the Squamata is at once distinguished from that of all other reptiles by the movable. squamosal. and has wholly disappeared in the Sauropterygia. that both the tabular and suprareptiles. to form part of the articular surface for upper end. as was formerly believed and yet is by some. according to the identification. its 65 both in the past and the present the posterior bone terior synonym. that the quadratojugal is a very inconstant bone in in the skull of the lizards. has no relations with the quadrate as has the bone so called If so. are absent. The supratemporal is the first bone its to disappear in the temporal region of the Cotylosauria. some forms. That it should lose its original position at the lower outer side of its the quadrate. It is quite possible. Yet others call the outer anterior bone the his reasons for believing that The author has given is none of these but the tabular instead. ? Furthermore. the quadratojugal absent. is He believes that the posterior the tabular because its it occupies the primitive position of that bone in relations to the interparietal. and quadrate. is It is very small in the Thero- morpha. The teeth are . the bone articulating with it in front and forming the outer boundary of the temporal opening may be the quadratojugal.

enclosing the auditory meatus. supramastoid. and there time. and rarely in certain degraded burrowing lizards (Fig. and quadrate. and a third bone whose homology the supratemporal. to). 56) the quadrate articulates above nor- mally with three bones. the mosasaurs there On the inner side the c). Below. 56) by a close sutural joint. In some lizards the tabular has suffered reduction or has become indistinguishably fused with the squamosal. the squamosal. in some. as here identified. squamosal. which tinct orders. articulates with the squamosal. the quadrate articulates with the pterygoid on the inner by a rather free joint in most lizards. supratemporal. The reasons for their identification as the tabular and squamosal will be found side in the discussion of the skull of the Protorosauria. is always Other characters are very variable in this sometimes divided into two or three dis- Sauria or Lacertilia In the lizards (Figs. all is these bones. 55 a. less fixed the quadrate lying against the brain-case and more or pterygoid. extending nearly or quite to the semicircular canals. there has been great dif- by no means unanimity at the present The tabular. but which is usually called The squamosal may be absent in those lizards without a temporal arch. 54 c) only. by the The paroccipital usually but not always helps support the quadrate. As regards the identity of ference of opinion. usually slender stapes abuts against the quadrate (Fig. 55. supratemporal. the prearticular of the mandibles fused with the articular. extensive order. In the mosasaurs (Fig. and even the paroccipital (opisthotic). forming more or less of the boundary of the temporal opening. and prosquamosal. The squamosal as here considered has been called the quadratojugal. at the distal and under side of the parietal process. is yet disputed. 56) the "supratemporal" may also be absent. paraquadrate. 55 is an elongated suprastapedial process arching backward and often extending to the lower end. paroccipital. firmly wedged in between the paroccipital and prootic.66 THE OSTEOLOGY OF THE REPTILES is acrodont or pleurodont. The tabular (Fig. like the AmIn phisbaenia (Fig. paroccipital. or supratemporal. it has a long internal process. . has been called the squamosal. as in some turtles.

vestigial in the Heloder- matidae. 54. premaxilla. parietal. from the side. pterygoid. lower figure. sq. pj. jugal. maxilla. oc. very rarely. basisphenoid. frontal. m. . Tylosatirus. splenial. /. po. coronoid. nasal. occipital condyle. ep. and Amphisbaenidae. epipterygoid. Anniella. c. the jugal. as in Uromastix. q. by a slender prolongation with the tip of Fig. an. postorbital. mal. angular. Platecarpus. anteriorly with the postorbital. from above. with the jugal only. pm. middle figure. transverse. /r. Clidastes. squamosal. pt.THE SKULL OF REPTILES The squamosal normally articulates with the tabular 67 and quadboth in rate posteriorly. prefrontal. sp. lacri- na. pa. bs. the lizards and mosasaurs. tr. from below. quadrate. j. vomer. In some lizards it has suffered reduction. palatine. v. Mosasaur skulls: Upper figure. and is absent in the Geck- onidae. and often.

Chameleon. Varanus. from the side. same. . Lacertilian skulls: A. D. Three fourths natural size. 55.B. C. from above. E. and the upper end of quadrate. left mandible. F. Natural size. the postoptic. from below. Conolophus. Natural size. the Fig.

The premaxillae may be paired 54 c) the or fused. squamosal. B.THE SKULL OF REPTILES 69 parietal. or the vestigial. united bone is fused with the nasals posteriorly. side view of atlas and axis. completely Fig. and postorbital. below. Five roofed over by dermal bones.^w^tf^««. D. The temporal fossa. 56. . C. 55 a) bounded above by the below by the tabular. from above. and the size. side. or obliterated by the union of the tem- poral arch with the parietal. as in the 'Amphisbaenidae (Fig. y/w/). in the mosasaurs (Fig. 56 a). latter may be absent or The lacrimals are always small. normally (Fig. may be wholly absent. Lacertilian skull: halves natural A.

the sides with the maxillae and more They sometimes bear teeth. articulates normally with the premaxilla. — G. 54 a).] . are intercalated between the prevomers and pterygoids. They usually bear teeth. The pterygoids have is the normal articu- lations except that in front they articulate with the palatines only. posteriorly with the palatines only. The epipterygoid. below with The is postorbito-jugal and the postorbito-squamosal arch lizards. The prefrontals are always tering into the formation of the nares in the Varanidae and Mosa- sauridae (Fig. 55 d). usually paired. The posterior palatine opening usually large. ectopterygoid. may be vestigial or even entirely absent The maxilla ectopterygoid. They articulate with the palatine by a descending process. articulating on or less with the ectopterygoids. usually. 55 d) ending forward over the orbit to meet the prefrontal. palatine. the two bones are indistinguishably fused or the postfrontal is absent. however. N. It in lizards. laterally usually with the maxillae. They very rarely bear small teeth.^ articulating 1 in a [Also Dibamid«. prevomer. Posteriorly the postorbitals articulate as usual with the squamosal the jugals. On the palate the prevomers are paired or partially fused in the Rhiptoglossa. is present so far as known in all lizards except the Amphisbaenia and Rhiptoglossa. postorbital. a slender rod. unlike those of most other reptiles. with the jugal. they are separated from the nares in the Rhiptoglossa (Fig. en- sometimes vestigial or absent. The postfrontal and postorbital are not rarely found united by suture in the mosasaurs. They articulate in front (Fig. lizards The postfrontals are rarely large in and are often absent. and often with the tip of the squamosal. 55 c). are sometimes fused with the premaxillae or with each other. 55 c) with the pre- maxillae. It articu- with the maxilla. when absent the postorbitals take their place. It always bears a single row of acrodont or pleuro- dont. pointed or obtuse teeth. may be absent in various terrestrial jugal The lates a slender bone bordering the orbit below and extend- ing forward to meet the lacrimal when that bone is present.70 THE OSTEOLOGY OF THE REPTILES large. sometimes with the nasal and prefrontal. The nasals. K. The palatines (Fig. . sometimes (Fig.

] . exocci57. cor. articular. — G. exoccipital. The by brain-case of lizards. The and the and parietals always are fused in the midis line (Fig. and some others have descending processes of which meet ber. itive in the middle below. quadrate. articular. and postoptics. 56 c) and Mosasauria the sides of the parietals are partially decurved. strengthening the peculiar The mandibles noid. usually present. usually lost in macer- AMAi-i-#^^ Fig. ang. pa. Helodermatidae. prearticular. st. surangular. pitals. postfrontal. Piatecarpus. membranous in front on the are small ossifications in the wall The postoptics (Fig. 55 a). occipital. the supraoccipital. with a long fused prearticular. q. 58) are basisphenoid. sur. Mosasaur mandible: Clidastes. forming incomplete cerebral (Fig. angular. 55 b. al) membrane. is as of other reptiles. pterygoid. 55 d. In the Amphisbaenia (Fig. is prootics. to the but they do not reach.^ The parietal foramen.has\eo. coronoid. stapes. surangular.THE SKULL OF REPTILES pit 71 on the upper side of the pterygoids and extending to or toward frontals usually the parietals. N. basisphenoid. in the Rhiptoglossa. 59 b). art. which in the mosasaurs is more or less ensheathed by the union of the coronoid and angular. ation. absent in many terrestrial lizards parietals may the be either paired or unpaired. basioccipital. formed FiG. K. 1 [Some geckos have them separate. en- closing a rhinencephalic champrim- very much like the one of the early reptiles. angular. i'0. p/. walls. paroccipitals. occ\p\ta\ view. as in the snakes (Figs. 58. frontals The frontals and The frontals in the Varanidae. pt. composed of the dentary coroand splenial. but less more or sides. inner side of right mandible.

Only the dentary is freely articulated in the mandible. 59 e) have but one functional tooth attached to the maxilla. enters into conical teeth of the premaxillae. epiptery golds. a similar joint. the The premaxillae are small and The pterygoids bear long teeth. as usual in with a long median symphysis of the mandibles. and the prootics are largely exposed on the side of the skull. which may also be absent. and there is no jugal. The vipers (Fig. the symphysis.72 joint THE OSTEOLOGY OF THE REPTILES between the angular and splenial. and palatines usually the maxillae below. Ophidia or Serpentes (Figs. the quadrate articulates proximally with the tabular only. . but in the and dentaries more specialized types are rather loosely lodged in grooves. The postorbitals may meet two mandibles are usually united in front by ligament only. The skull of snakes differs from that of lizards. the maxillae rarely edentulous. there are many variations in the skull of the more than in many other groups of reptiles called orders. rarely. no ectopterygoid. usually united in front by suture but are ligamentously connected in the mosasaurs and some land lizards. reptiles The long splenial. There is no ossified interorbital septum. the posterior bones closely fused. and. As is seen. especially in the complete closure of the brain cavity in front by descending plates from the parietals and frontals. with an opening at its base and another near its apex for the passage of venom. there is no parietal foramen. There no temporal arch. often edentulous. the former always meeting the basisphenoid below. and squamosals. in the constant absence of the postoptics. except that the coronoid appears to be absent or fused. It is hollow. 59 a-e) lizards. the latter sometimes interrupted by the coalesced optic foramina. The parietals are is always fused. the bone usually so called being clearly the prearticular. The primitively were inserted in sockets. though less The mandibles are is found in the monitor lizards. well developed. maxillae. The mandible of Ophidia has the primitive structure.

and palatine bone with teeth. Crotalus. B.Fig. Natural size. C. from above. Ophidian skulls: A. Python. E. Natural size. 73 . 59. occiput. D. and from the side. from the side.

quadrate. squamosal. 5y. 213. prf. postfrontal. Jy. angular. prearticFig. y. Some 25. dentary. 61. Thalattosaur skull: Thalaltosaurus.74 THE OSTEOLOGY OF THE REPTn. from the side and above. /. nasal. Fig.ES The Skull of the Rhynchocephalia (Figs. . premaxilla. One eighth natural size. 279. wj. postorbital. pm. frontal. maxilla. jugal. f. an.] skull characters are noted on pages 20. 21. from above and from the After Merriam. articular. ular. 60. 60-63) [No manuscript. pj. «. /)o. coronoid. side. surangular. y. d. jugal. prefrontal.quadrato/). /)«. ^r/. Rhynchocephalian skull: Sphenodon (Tuatera). ja. parietal.

One fourth natural size. C. from below. 75 . Stenometopon. After Huxley. from above. E. After Burckhardt.Fig. Hyperodapedon. B. D. One fourth natural size. After Boulenger. the same. from the side. After Boulenger. from above and from the side. 62. Rhynchosaur skulls: A.

• . 63.Fig. 76 s-n •/-' . from above and from below. After Brown. Choristoderan skull: Champsosaurus. One half natural size.

The Skull of the Pseudosuchia
(Fig. 65 a-e)



skull of the typical




much like

that of the

Pelycosimia (Fig. 64), in structure. All the bones of the skull roof are present except the dermosupraoccipital, tabular, and supratemporal; the lacrimal

small; there


palate bones have the primitive relations.
occipital and tabular {Youngina, Fig. 64 vomers and pterygoids (Proterosuchus).

no parietal foramen; and the Other forms, however,

referred to this group provisionally, have both the dermosupra-

and teeth on the prelateral

The upper and

temporal openings, a large antorbital vacuity and one in the mandible, are like those of the Parasuchia.

large, as are also the orbits.

The antorbital foramen is The supra temporal foramen is large

and never posterior

in position.

The Skull

of the Pelycosimia
(Fig. 64)

skull of the Pelycosimia differs

chiefly in the position of the external

from that of the Phytosauria and internal nostrils near the

extremity of the face, and at a considerable distance in front both of
the orbits and antorbital openings.


short in front of the




no respiratory channel back of the internal

nostrils, so characteristic of the





carnivorous in type.

Fig. 64.

Pelycosimian skull: Erythrosuchus, from above.

After von Huene.


sixth natural size.


The Skull of the Phytosaurla.
(Figs. 6s, 66, 67)


The skull of


Phy tosauria is nearly uniform

in general structure,

characterized especially by the elongated face and posterior location
of the external nostrils.

No bones are

fused in the midline, and none,

save the primitive dermosupraoccipital, tabulars, and supratemporals are missing.


paroccipitals, as usual, are firmly fused with

the exoccipitals.


no parietal foramen. The supratemporal
depressed below the level of the parietals

openings are more or


but retain their primitive boundaries. The well-developed quadratojugals enter into the formation of the lateral temporal openings



a primitive quadrate foramen between the

quadratojugal and the quadrate.
large antorbital foramen








bounded by the

maxilla, nasal, lacrimal,



greatly elongated face


chiefly of the premaxillae,

which extend back

to the anterior

ends of the nares, with the septo-

maxillae intervening, in the middle.


nostrils are

surrounded by

the large nasals and are elevated to or above the superior plane of the skull.

The bones

of the palate retain their primitive relations,

and there

are small posterior palatine vacuities, larger in the

more primitive
forming the

The pterygoids meet broadly
some almost forming an

in the



roof of a deep respiratory channel between the heavy, underarching
palatines, in

incipient secondary palate, in

the phytosaurs, as in the crocodiles, doubtless caused by the large



interpterygoidal opening

and parasphenoid are


The elongate prearticular of the mandible is fused with the articuAs usual in slender-jawed reptiles with a long symphysis, the splenial participates in it, an acquired character. The condition of

the coronoid


not yet definitely determined, but

it is


present, though small.

A large foramen, so generally characteristic of

the Archosauria,


constant in the outer wall of the mandible be-

tween the surangular, angular, and dentary. The teeth are numerous, set in deep sockets and confined, as


other archosaurians, to the premaxillae, maxillae, and dentaries,

Fig. 6s.

Pseudosuchian skulls: A, B, Euparkeria, from the side and from above. After Three Five sixths natural size. C, Youngina, from above. After Broom D, E, Ornithosuchus, from above and from the side. After Bouhalves natural size. lenger. One half natural size.



either cylindrical throughout or partly or chiefly flattened


and dentiabove^




two or three teeth on each

side, especially

Fig. 66.



Machaeroprosopus, from above and from below.

are cylindrical and



The dentulous portion

of the



long, with

twenty or more teeth on each


chief differences in the skull structure of the various


of the order are


in the relative position of the external nares^

Fig. 67. Phytosaur skull: Mystriosuchus. />W2, premaxilla; na, nasal; /, frontal; p, prefrontal; /, lacwj, maxilla; rimal; pj, postfrontal; po, postorbital; pa, parietal; sq, squamosal; qj, quadratojugal; pi, palatine; /, transverse;
in, internal nares; en, external nares;

basisphenoid; eo, exoccipital.

pt, pterygoid; After McGregor.



The Skull of the Crocodilia (Figs. coelian types. In the broader-faced amphicoelian and in it is the procoelian types a cylindrical bar with a considerable space be(Fig. basisphenoid. is There is no ossified interorbital septum. exoparoccipi- and pterygoids. in the median line between the basioccipital and basisphenoid. excluding the prevomers . postoptics. 68. squamosals. laginous septum in face. The quadratojugals take part in the formation of the lateral temporal opening. and the paroccipital parietals is is fused with the exoccipital. more slender and cylindrical jaws. parietals. and parietal foramen. epipterygoids. ao) are fully ossi- tween fied. The most important modifications of the crocodilian skull are found in the palate. The lateral temporal opening is separated in the teleo- saurs from the orbits by an unmodified postorbital bar immediately all below the skin. An antorbital vacuity often present but only rarely has been found in the early pronasals may or may not separate the external nares. and the shape of the teeth. a triangular bone. The and f rentals are fused in the midline. distinguishing these reptiles from all others. and almost [or entirely] obsotals. an extensive connection. small in the later ones. connecting with the premaxillae they are divided by a cartiin the teleosaurs. postorbitals. 69 d. The postoptics extending from the basisphenoid it to the frontals. in those with slender more flattened and compressed in those with a compressed and elevated face. excluded from the foramen The supraoccipital magnum. septomaxillae. and the skin. The nares are always at the extremity of the it no matter how long and slender may be. The maxillae meet broadly in the middle line. There is a eusta- chian canal connecting with the otic sinuses. life. tabulars. The quadrates are firmly wedged in between the quadratojugals. The supratemporal openings are large in the early forms. the more general carnivorous habits of the latter.V THE SKULL OF REPTILES 83 the extent of the elevated facial carina in front of the nares. 69) The skull of the Crocodilia invariably lacks the postfrontals. narrow bones between the quadrates and jugals extending forward to meet the postorbitals. supratemporals. The . doubtless because of the more dominant fish-eating habits of the former. prootics. lete in some.

One C.-TUX Crocodilian skulls: A. After Fraas. above. Onehalf natural E. After Lortet. After Andrews. 68. One half natural size. from above. B. ozz\\)\iX. AlUgatorellus.. Fig. from below. Hylaeochampsa. About one fifth natural size. Alligator. size. Teleosaurus. from above. from twelfth natural size. T). . Dakosaurus.

. 69.Fig. Alligator skull: one half natural size.

The pterygoids also meet in the middle line. 69 d) the splenials meet in a median symphysis in slender jaws. which may or may not be divided by a median meeting below the nasal tubes in front of them. or in front of the pterygoids in the tomistomids. In the goniophilids the openings are surrounded by both partition. tween the angular. and forming the the respiratory canals. and do not often appear on the palatal surface. The palatines also meet in the middle line in all. in Hylaeochampsa. they are separate as usual. There is a large mandibular foramen beside. In the early teleosaurs these openings were at the posterior border of the palatines. absent or fused with the angular. in all procoelian forms completely surrounding the internal nares. and. The prevomers form a pair of tubes above the maxillae and palatines. possibly representing the epipterygoids. The prearticular is apparently wholly absent. The pterygoids articulate posteriorly and externally with the postoptics by a narrow pillar. and dentary on the outer in the Thalattosuchia. an additional opening in the palate between the ectopterygoid and maxilla.86 THE OSTEOLOGY OF THE REPTILES of the premaxillae from the palatal surface. palatines and pterygoids. sending up procfloor of esses for articulation with the postorbital. In the mandible (Fig. except rarely just back in the caimans. . articulating posteriorly with a suprapalatal prolongation of the pterygoids. surangular. There are large posterior palatine vacuities at the sides of the palatines.

70) [No manuscript.] Skull characters noted. 28. . Ornithischia) (Fig. pages 17.The Skull of the Dinosaurs (Saurischm. 214. 291-296. 32.

After von Huene. 17. 72) Skull characters noted. pages 11. Ornithodesmus. One half natural size. B. 71. from Fig. After Hooley. Pterosaur skulls: A. 19. Rhamphorhynchus . 214. Pteranodon. E. 71. C. palate. About one fourteenth natural size . front part of the side. 14. 28. 21. Rhamphorhynchus.] of the Pterosaurl\ (Figs. from the side.The Skull (No manuscript. 296-298. One half natural size. Campylognathus . end of beak. D. After Plieninger. One fourth natural size.

from below. Nyctosaurus.Fig. from above. C. the same from below. Pteranodon. 72. after Eaton. B. Pterosaur skulls: A. About one fourth natural size. 89 . About five eighths natural size.

column or backbone of reptiles. are [For the modern embryological viewpoint of the composition of reptilian vertebrae see Schauinsland. anterior dorsal from the side and front. Sphenodon (Rhynchocephalia). Projections from the vertebrae. or centrum. etc. after Eaton. 73 b) is composed of a body. 87 B. c). Ophidia. and an arch.^ Fig. in Hertwig's tieren. anterior dorsal from the side and front. C.. D. and are often pairs of proc- Two in front and one behind. for articulation with adjacent vertebrae. or neurapophysis.] Handhuch der Entwickeluiigsgesc/ii elite der Wirbel- . as of all other air-breathis made up of a variable number of separate segments called vertebrae. the neurocentral sutures more persisspinal tent than in most mammals. F. 73. young or aquatic reptiles always (Fig. called processes or apophyses. serve for the attachment of muscles or ligaments. adult land reptiles often showing them. Iguana (Lacertilia). — Ed. 1906.CHAPTER II THE VERTEBRAE The ing vertebrates. E. each ossifying separately and uniting at variable times. anterior dorsal from behind and in front. or for the support of characteristically different in different reptiles. esses springing from the arch. A vertebra (Fig. Pter- anodon (Pterosauria). cervical from the side. one ' ribs. Anterior dorsal and cervical vertebrae: A. B. G.

only in reptiles with a long. have additional articulations. fitting into a cavity. flat The pair in front. the posterior end of the next preceding vertebra. f) known as zygosphenes and zygantra (Fig. The in the posterior part zygapophyses of the may be obsolete or even absent column of aquatic reptiles. as is seen. in that the wedge-shaped median process. 9r as zygapophyses. 73 e). The vertebrae of all snakes. 74. where they were first recognized and described. the posterior pair. that is. called exapophyses (Fig. especially characteristic of certain cotylosaurs. 73 D. The zygosphene is a wedge-shaped process at the anterior end of the arch. toward the dorsal side. Dorsal vertebra: Diadectes (Cotylosauria) from behind. slender vertebral columns. posphen*". flexuous. toward the ventral side.THE VERTEBRAE known that is. They strengthen the articulations. above and be- tween the zygapophyses. the prezygapophyses {az). extensions of the zygapophysial articulations about their inner ends. or rather. (Fig. They occur. though restricting vertical flexure. postzygapophyses. of the next succeeding vertebra. is turned downward. always has the or concave articular surface directed upward. showing diapophyses. placodonts. 74) is below and between the inner ends of the postzygapophyses. — Ed. or downward and outward.] . their articulating surfaces facing in opposite directions to those of the zygapophyses above them. which corresponding cavity. and some mosasaurs. and hy- In certain other reptiles is this arrangement reversed. or upward and inward while that of . The later pterodactyls have another pair of articulating processes. Zygosphenes and zygantra strength- en the articulations. called hypo- sphene (Fig. fits into a at the zygantrum. [Exceptions to this rule occur in recent lizards. the postzygapophyses (pz). flexible vertebral in column/ and are absent in those mosasaurs which the column is less elongate and Zygosphenes are also known to Fig. but limit torsion. at each end of the cervical vertebrae on the ventral side. the hypantrum. and are substitutes. at the front end Hyposphenes and hypantra are and dinosaurs. some lizards. occur in certain aquatic Stegocephalia with long. 73 g).

^ — Ed. and simons). Paired hypapophyses (lymphapophyses) are characteristic of the caudal vertebrae of snakes. A like process or facet on the side of the is centrum anteriorly for . often as far back as Hypapophyses the tail. is properly called a hypapophysis (Figs.92 THE OSTEOLOGY OF THE REPTILES for the peculiar saddle-shaped articulations of the cervical vertebrae of birds. 1 [The eggs are cut. is the spine or neura- pophysis. A longer or shorter process on the sides of the arch for the support in part or wholly of the ribs 75). — Ed. 73 d-f) The and are never long or slender in aquatic reptiles. of which probably the anterior ones. As a rule. is known is as a diapophysis (Fig.^ They also often occur on the cervical vertebrae of lizards. at least. in the middle. 73 f) Either is commonly called a transverse process. 74) If the cavities are deeply concave. in all cases are coossified ribs. the vertebrae are . where they replace the absent all chevrons. 75 a). and most amphibians.] . spines of most sauropod dinosaurs in front of the sacrum are broadly divided. the vertebrae are called notochordal.^ When reptiles. on the under side of the vertebrae. and the same term merely often applied to a like process on the sides of the caudal vertebrae. And this was the primitive condition found in the Cotylosauria (Fig. A process. of extremely variable size and length.] of in the oesophagus. 73 e. The spines are always short in legless or slender crawling reptiles (Fig. the ends of the centra are concave. are characteristic of snakes. in front at least. communicating with each other through the centrum. articulation of the head of the rib called a parapophysis (Fig. the notochord was continuous in life. doubtless for the lodgment of stout muscles and ligaments used in controlling the long neck. as they are in early nearly all fishes. not the stomach (Fitzin the [The distinctions between lymphapophyses and hypapophyses break down embryology modern lizards. for the attachment of muscles and ligaments controlling the neck and head. that is. V-shaped. not crushed. On the dorsal side of the arch. 73 b. the spines are longest and stoutest at the beginning of the dorsal series. paired or single. in some masticatory instances they are developed to serve as a sort of apparatus for the crushing of eggs in the stomach. sometimes rudi- mentary. sometimes very long. and turtles. crocodiles. known as amphicoelous (Fig.

sl = Amyda) living river-turtle. amphicoelian types. Platypeltis spinifera. have the convexity centrum at the front end. plano-concave. otherwise known caudal vertebra of the procoelian crocodiles. dating possibly from early Jurassic times. and Sphenodon. but most others have an extraordinary combination of types. in early Cretaceous times {Hylaeochampsa) . occur. are not frogs having them.THE VERTEBRAE 74) 93 to the present time and Triassic Ichthyosauria and continuous in the living gecko lizards. Procoelous vertebrae. f). as also the cervical verte- many theropod and orthopod dinosaurs. and are doubtfully known and caudals fishes in other reptiles. save the cervicals of certain turtles. and in some modern and many modern sala- Most remarkable are the cervical vertebrae of the Chelonia. restricted to reptiles. all amphicoelous. procoelous. More flat usually. are procoelous. so far as we yet know. They do manders. however. posthe Crocodilia Procoelous vertebrae appeared among . Eocene times. A ball-and-socket joint appears at time. in only the first ( plano-convex. called procoelous. with the concavity in front. opisthocoelous. among lizards. bowl. graduall reptiles became the prevailing one. The presacral vertebrae brae of of the of the sauropod. some modern They doubtand less arose in terrrestrial crawling reptiles with a flexuous column. Possibly the ptero- dactyls acquired the ball-and-socket articulations after the attain- ment of flight. 73 e. as are also the presacral vertebrae of the Pterosauria. the turtles. or almost {platycoelous) or even quite {amphiplatyan) all Until after the middle Jurassic times the vertebrae of reptiles known that were amphicoelous. since early except the geckos sessing them. Such vertebrae have been called opisthocoelous. ally This kind of vertebra. The earliest that we know had amphicoelous vertebrae throughout the column. dating from Lower Cretaceous. persisting until early Eocene (Dyrosaurus) of all The vertebrae known snakes (Fig. The caudal vertebrae of some turtles are procoelous. has opisthocoelous cervical vertebrae. . however. aigialosaurs and it was doubtless from such ancestors that the mosasaurs. aquatic reptiles. in the cervical region of certain Triassic Stegocephalia. inherited them. since middle Permian flat times the cavities are shallow. the ball or convexity behind. just the reverse of procoelous.or saucer-like. and even biconvex. however.

supposed that of this hollow- ness and lightness the cervical and dorsal vertebrae. Eosauravus. One tenth natural FiG. dorsal. twenty-two to twenty-five dorsals. one or two Trimerorhachis. and sometimes cervical lumbar region mammals. The rated pleurocoelous (Fig. between the shoulder the sacral that which sup- ports the hip girdle. sacral. Except in the snakes and legless lizards. sacrals. subaquatic in habit. but the arch curiously strengthened tresses. sepa- by a median an oval or round is orifice at each Not only the centrum thus lightis ened in these dinosaurs. We may tiles.94 THE OSTEOLOGY OF THE REPTILES turtles and certain pleurodiral no other order have saddle-shaped articulations. the spinal column of reptiles is divisible into cervical. been an increase in number in some. dinosaurs are peculiar in having large cavities in the centra. tail. After Hatcher. size. the so-called atlas. had at least twenty-four or twenty-five pre- . 75. In of reptiles are the variations so great as in this. served to keep the body erect in water. and the caudal that of tail. correlated with the other- wise solid or cancellous skeleton. has thirty-one precaudal vertebrae and no differen- The earliest reptile that we know. hardly venture to guess as to the primitive number of vertebrae in rep- We are quite sure that there has decrease in others. the dorsal that and hip the Dorsal vertebra of Diplodocus (Saurischia). tiles by plates and but- Certain other South African rep- {Tamboeria) also have pleurocoelous It is vertebrae. an aquatic and a short or Lower Permian temnospondyl. the caudal and precaudal. and caudal also. with side. their natural habitat in wading or swimming. The that in front of the shoulder- girdle. a The land temnospondylous amphibians that real cervical vertebra. girdles. where but two regions are recognized. we know have but one moderately long tiated sacrals. 75) presacral vertebrae of the sauropod partition. as in is regions.

two sacrals. and we may omit the very variable terrestrial reptiles.THE VERTEBRAE caudals. and not more than sixty caudals. The smallest number of presacral vertebrae known in any reptile sixteen is recorded for Brooksia. In all probability the earliest reptiles were seen. and they had not less legitimately presumable that than twenty-three nor more than twenty-six verte- brae in front of the sacrum. and a long tail. as against thirty-five in modern turtles and four hundred and fifty in some modern snakes. the largest number found in any early reptile. a single sacral. a recent cha- — — meleon lizard. is the number of vertebrae known. six it is our comparisons. The numbers of presacral and sacral 95 In no embolomerous amphib- ian vertebrae in reptiles Presacral may Sacral be tabulated as follows: Cotylosauria 23-26 18 1-3 2-3 2-3 Chelonia Theromorpha Therapsida Nothosauria Plesiosauria 23-27 25-28 2-7 2 Proganosauria Ichthyosauria Sauranodon (Saphaeosaurus) Kionocrania (Lacertilia) Rhiptoglossa Dolichosauria Mosasauria Rhynchocephalia 40~42 40-105 29-34 40-65 22-23 22-74 16 29 3-4 2 o 2 0-2 2 2 29-42 25 o 2 2 Rhynchosauria Choristodera Pseudosuchia Phytosauria Eusuchia Thalattosuchia 23-24 23-26 23-26 26 2 2 2 23-24 25 23 2 2 Theropoda Sauropoda Stegosauria 2-5 26 27 Trachodontia Iguanodontia Ceratopsia 30-34 24-28 24 4-5 3-4 8-9 4-5 7 The earliest reptiles had functional tail in ribs and a sacrum. The majority of have between twenty-three and twentypresacral vertebrae. . it is lowland and crawling in habit. or altogether between eighty and ninety vertebrae in the whole column.

They been found in the anterior vertebrae of some plesiosaurs. though not by all. in early Permian times. where they are usually Double intercentra have also been observed vertebrae of Procolophon. often in their normal places between the centra but frequently shifted it for- ward on the preceding centrum. It was Cope the identity of the parts and his views are who first recognized now generally accepted. The former are known in only a few amphibians. THE OSTEOLOGY OF THE REPTILES The earliest reptiles probably all have a small or more or less wedge-shaped bone intercalated between the adjacent ventral margins of the centra throughout the column. though the centra are deeply biconcave. that is. In the Ichthyosauria. from the Mississippian. They occur in many procoelous lizards throughout the neck. but have remained to modern times in the They have persisted in nearly all reptiles in the tail as the chevrons. in the more primitive vertebrae. a few other reptiles.96 Ifiler centra. to which Professor Cope in 1878 gave the name inter centrum (Fig. and never occur in procoelian. and in the young of certain plesiosaurs.. It is now universally believed that the undivided or holospondylous vertebrae of reptiles were evolved from divided or temnospondylous have also vertebrae of the Stegocephalia. Intercentra had previously long been known as "intervertebral" or " sub vertebral wedge-shaped bones. The intercentrum of the first vertebra has. Penn- . vestigial. having been entirely lost as simple intercentra only in the crocodiles and or middle gecko lizards and in Sphenodon. amphicoelian. either loosely attached or coossified with an exogenous outgrowth. 76 e). first two to four vertebrae. or opisthocoelian reptiles. only two to four intercentra have been observed. 76 a-c) and rhachitomous (Fig." but their significance was ill understood. called by Cope embolomerous (Fig. 76 d). pophysis. and more or less in the neck. a cotylosaur. all Amniota as the basal piece or "body" of Intercentra are characteristic of deeply amphicoelous or noto- chordal dorsal vertebrae. forming with a functional hypa- Where they occur between the centra they is may be elon- gated into false hypapophyses. With the more complete ossification of the ver- tebral centra they began to disappear in the dorsal region. A similar condition known in some in the anterior Chelonia on the paired. Temnospondylous vertebrae are of two kinds. remained functional in the atlas.

from Illinois An embolomerous vertebra is composed of two subequal. while the pleurocentra behind are paired. 76 d) differs in that the intercentrum or hypocentrum is more or less wedge-shaped. lates is borne by all The head of the ribs articuwith the intercentrum. that is. and probably an intermediate condition between the embolomerous and rhachitomous types. Rhachitomous vertebrae are much more widely known in numerous forms from the Pennsylvanian and Permian of various parts of the world. bones. This reversion of the pleurocentrum to a more primitive ontoit genetic condition is is the chief objection to this theory. rhachitomous and holospondylous vertebrae appearing later. We have seen that the more primitive phylo- genetic condition of the intercentra persists longest in the neck and d). as was . The earliest known amphibian vertebrae are embolomerous. 76 is other rhachitomous amphibians. resting upon both the intercentrum and pleurocentrum. the anterior one the intercentrum. while the intercentrum bearing tail. there . its apex not reaching the dorsal side. The artic- head or capitulum of the ribs is chiefly on the intercentrum. or hypocentrum. The three neurocentrum. with its base on rib. so far as our present knowledge goes. as in the embolomerous forms.THE VERTEBRAE sylvanian. but chiefly the ular surface for the latter. with the basal side above and their apices reaching the ventral line only narrowly or not at all. 76 a-c) and Texas. disk-like and perforated for the notochord. A rhachitomous vertebra (Fig. bearing the exogenous chevron in the tail. and the arch or neurocentrum. in which the single pleurocentrum is typically embolomerous. noto- chordal disks. givfirst it ing origin directly to the reptilian holospondylous type. suggested by Cope that the rhachitomous type was derived from by the loss of the of the pleurocentrum upper part of the intercentrum and the lower part and the division of the latter into two lateral parts. the ventral line. nevertheless the more probable. and 97 Lower Permian. but best in Cricotus (Fig. And this is one of the reasons why it would seem that the embolomerous type is the more primitive. but chiefly by the pleurocentra. the surface for the articulation of the tubercle of the on either the arch or diapophysis. the posterior one the pleurocenlrum. the tubercle with the diapophysis of the neurocentrum. In the caudal vertebrae of Eryops (Fig.

and median caudal. caudal. E. the most amphibian-like.98 its THE OSTEOLOGY OF THE REPTILES exogenous chevron is typically rhachitomous. D. The intercentrum is here remarkably large for a reptile. from the side. 76. 76 e). Dimetrodon (Pelycosauria). Trimerorhachis (Temnospondyli). dorsal. Additional evidence fishes. from the side and front. basal caudal. in the is fur- nished by the fact that while truly embolomerous vertebrae occur in modern Amia. sauria). real rhachitomous vertebrae i . G. Eryops (Temnospondyli). 79). as will be shown in the discussion of that bone. Vertebrae: A. the atlas (Fig. nearly half as long as the notochordal centrum or pleurocentrum. F. Cr. in that it is wedge- shaped. And this very probably represents the real intermediate condition between the embolomerous and holospondylous vertebrae. of all known reptiles (Fig. C. for instance. from the side. Evidence that reptilian vertebrae arose in dorsal vertebrae of a this way is also seen in the young Seymouria. dorsal intercentrum from behind and below. Fig.Vo/Mj (Temnospondyli). otherwise. intercentrum from side and below. Seymouria (Cotylomedian dorsal. B. And it is also almost the condition found in the first vertebra of primitive reptiles.

Certain it is cormus). the rocks. have but one or two vertebrae which may properly be called cervical. first rib the attached to distinction it is definitely determines the beginning of the thorax. tail true. the other with it the undivided pleurocentrum. The evolution. is paralleled by the separation of the primitively single intercentrum into pairs. the changes in their shapes. and a its final loss everywhere in the column save . the intercentrum. and but little motion of the head in a lateral direction. may indicate approximately the beginning of the dorsal however. In such. The retrogression of the disk-like pleurocentrum into the paired pleurocentra of the Rhachitomi. observed in Procolophon. as in the Sauropterygia reptiles and Archaeosauria. But the early had no sternum. and chevrons of the tail synonym of the earlier and thus the term hypocentrum becomes purely term intercentrum. like the Temnospondyli. more so than that of the dorsal verteThe Cotylosauria. of the holospondylous reptilian vertebra from : the temnospondylous amphibian vertebra seems clear by the simple increase in size of the notochordal centrum and the progressive dein the atlas crease of the intercentrum to a wedge-shaped. like their immediate ancestors. and some plesiosaurs. with dorsal embolomerous vertebrae. the its one with base below. composed of two half-disks. found in the position of the pectoral girdle as found in brae. then. had practically no neck. The number is very variable. with other modifications. subvertebral bone. Primitive reptiles. then.THE VERTEBRAE are 99 ancient fishes {Eury- known only among amphibians. since the pectoral girdle is almost invariably found lying immediately back of the skull. have in the above. is series. pseudo-rhachitomous vertebrae. Better evidence. and free ribs were continuous from the atlas in their to the sacrum without change mode of articulation. . Cervical Vertebrae (Figs. In those reptiles having a sternum. indeed. many turtles. the Stegocephalia. between the angle of the jaws. 77-81) The number is of vertebrae in the neck or cervical region of reptiles not always easily determinable. is The almost as definite in those in which there a change in the articulation of the rib from the centrum to the arch. the front end of the interclavicle.

the latter in the tail-propelling aquatic types. arch of atlas. primitive theromorph: pa. that of in some cases these numbers upon the interpretation different observers. nine. eight. true lizards. the monitor lizards. Theropoda. are those given for the Therapsida. . and mored efinitely by the position of the scapula and clavicles as observed in various specimens. a Fig. odontoid. These numbers. Crocodilia. by the diapophyses. proatlas. the mammals. the dolichosaur lizards. 77. Modern chameleons is the have but five. ax.lOO THE OSTEOLOGY OF THE REPTH^ES longer neck. the Pseudosuchia. and Ophiacodon. from thirteen to as probably also the increase in number among the trachodont and sauropod dinosaurs may be attributed to water many as seventy-six. Iguanodontia. and seven sistently in their number that has remained so perdescendants. as many as fifteen. the plesiosaurs. are only approximately correct. among strictly amphibious or aquatic reptiles there has been an increase or decrease in the number. as shown by the lengths of the ribs. dependent what constitutes a cervical vertebra by On the other hand. the Pterosauria and Phytosauria. with intercentra. eight or nine. o. six or seven. however. with at least six and prob- The Theromorpha have a ably seven vertebrae (Fig. 77).two. The ancient proganosaurs have ten or eleven. It must be remembered. the Trachodontia and Sauropoda. the nothosaurs. Ceratopsia. Notochordal cervical vertebrae. thirteen. of axis. sixteen to twenty-one or twenty. an. as this order is imperfectly known. eight to ten. the Chelonia and Rhynchocephalia.

having seven. usually 80 D. B. the is first of our usual nomenclature. had practically no neck. the same proatlas. like the mosasaurs and aigialosaurs. c. Theromorph vertebrae: A. by some. as the arch of a vertebra whose centrum is represented by the anterior end of the odontoid. the atlas. 79. by others. intercentrum. present. atlas and axis. lOI tail. l) is a small. homologous with the so-called Amphibia. The following cervical vertebrae. or epistropheus. o. paired. 79 more or less vestigial neural arch between the arch of the atlas and the occiput. is in- constant and vestigial. are differentiated from the dorsal series more or by their erect or shorter spines. inner side. the proatlas. Another theory. the most specialized of all aquatic reptiles. D. or the slenderness rib articulation. Fig. /. of a vertebra formerly intercalated be- tween the atlas and the atlas of the skull. The first two or three of the cervical vertebrae are markedly differentiated in all reptiles. C. neurocentrum of atlas. crocodile. The first of these. pleurocentrum (odontoid). The proatlas (Figs. with a fin-Hke two. 91). Ophiacodon. Proatlas. and has not been included in the numbers above given. It is believed to be the arch Fig. the axis. representative of a vertebra fused with of the atlas. and mode of The cervicals of the ar- later pterodactyls have additional ticulations on their ventral sides. The more is second. from the front. as merely the separated spine by others. as has been described above Proatlas. neurocen- trum (arch). from the side. Pleurosaums probably had but five and the ichthyosaurs. The or less closely united with the atlas. third. «. lost The marine . as the Dimetrodon. Sphenacodon. (p. 78. namely. is that of Jaekel. by Baur. transverse processes. as in the higher animals. less to which has commend it. the same atlas. that the . axis. when less less and ribs. the occiput in the reptiles.THE VERTEBRAE habits.

in the way by zygapophyses. 79) of the former group. A roof-shaped. but no complete specimen has yet been discovered in . Fig. 78.I02 centrum THE OSTEOLOGY OF THE REPTILES of the proatlas is the so-called intercentrum of the atlas. It all is present many if not forms of the Theromorpha and Therapsida. the arch and inter- . wedgeupon a single large. of the predentate dinosaurs. In Iguanodon (Fig. or it may have is entirely disappeared. occurring as far back as Jurassic times. from which he concluded that the real body of this vertebra had become permanently fused with the basioccipital. it is doubtless present throughout the order. It has also been reported in the cha- meleon abnormal element and the mammals Erinaceus and Macacus. 80 d) also articulating with the atlas. that composed upon a large. atlas. axis. In its mammals. a small bone on each side. unpaired proatlas has been described in Rhampho- rhynchus. . notochordal pleurocentrum. As an it was also found by Baur in a trionychoid turtle {Platypeltis spinifer. reptiles (Fig. 80 l). and articulating with the arch of the atlas in the primitive way. is Atlas (Figs. partially fused with the occiput. arising from paired cartilages. which can be homologized with the is By some the so-called atlas of the amphibians thought to be represented by In the earliest is. embolomerous. 79). In it is the Crocodilia. as modern Sphenodon (Fig. roof-shaped. These articular surfaces appear to be present in known genera. a single bone in the adult. all of them loosely connected with the of a paired arch resting in part in part shaped intercentrum. a Jurassic pterosaur. the atlas temnospondylous in structure. in the it is paired. and the Triassic Plateosaurus of the Theropoda. 78) and Dimetrodon (Fig. Probably it will be evenlizards tually discovered in many other extinct reptiles. 80). is necessitating the view that the axial intercentrum merely an acfill cessory or provisional bone developed below the odontoid to out what would otherwise be an unoccupied space! Positive evidence of the proatlas has been discovered in several genera of the Cotylosauria. articulating in front by a facet on the exoccipital. 79. the arch of the atlas or neurocentrum articulating in the usual highest development. There ians no vertebra in the known amphibatlas of reptiles. behind with an anterior zygapophysis on the arch of the less both surfaces looking more or all downward. the proatlas. as also in several genera of the Sauro- poda. In it Ophiacodon is (Fig. 32).

D. C. N. Trinacromerum (Plesiosauria). J. Biplodocus (Dinosauria). Camptosanriis (Dinosauria). I. M. Platecarpus (Mosasauria). .Champsosaurus (Choristodera). Enaliosuchus (Crocodilia). L. Iguana (Lacertiha). Cymbospondylus (Ichthyosauria). after Marsh. Trinacromerum (Plesiosauria). C'. Apatosaurus (Dinosauria). V.Fig. and ribs: A. Nyctosaurus (Pterosauria). B. E. after Brown. H. K. 8o. after Riggs. ¥ . G. after Merriam. Chrysemys (Chelonia). Iguanodon (Dinosauria). after Dollo. after Gilmore. after Gilmore. Gavialis (Crocodilia). axis. 6'/)/^f«oa'o« (Rhynchocephalia). O. after Jaekel. Atlas. Baptanodon (Ichthyosauria).

centra have become contiguous below it. 78) ^^^ odoutoid. and the Squamata (Figs. After Holland. and Dimetrodon (Fig. 80 d). l). 80 f). that is. and the large odontoid is perforated for the noto- chord./j(Saurischia). This lack of most reptiles at was compensated for joint by the ball-and-socket cand the mals. In few is there any degree of roit. reaching the vertebra. still 80 g) and Chelonia (Fig. but the first and second inter- trum or centrum. large . the odontoid. however. which has no independent motion whatever. a small. and is not united with the axis. 81. tooth-shaped. atlas. 80 m) the pleurocentrum retains its . but most nearly approached in the Theriodonts. completed in the middle by the frOUt end of (Fig. The arch bears a rib upon its diapophysis.I04 THE OSTEOLOGY OF THE REPTILES ring. One fourth natural size. The pleurocen- and notochordal primitively. In no reptile did the atlas attain the spe- cialization of the mammals. tation about not more than between the torsion. as in the embryonic cartilage of mammals. In the Crocodilia (Fig. In very few do the two bones of the arch fuse with the intercentrum into a complete arch ring. and Phytosauria it is yet largely visible from the side. In the Rhynchocephalia (Fig. the pleurocentrum or true centrum. Atlas and axis of D//)/o^o<:. centrum are fused into a which revolves about its pleurocen- trum. 80 b. 79) the condylar cup is formed by the intercentrum and arch. grew progressively smaller till it finally ventral side of the disappeared wholly from side view in the Pterosauria (Fig. axis and the fol- lowing vertebra. Long ago. even approximately. or spout-shaped bone firmly fused with the axis in front and usually described as a part of it. lost in between the single condyle mam- In the primitive Ophiaco- don Fig. 80 e) most Dinosauria. the odontoid was recognized by Cuvier it as really the body it of the axis. in least. Choristodera (Fig. or does the pleurocentrum unite with the axis as a real odontoid.

is In the marine crocodiles (Fig. 81) its rib has migrated forward. In the short-necked Ichthyosauria the atlas earlier and in axis show a progressive fusion from the forms (Fig. Among the crocodiles anomodonts. but is is to a greater or less much reduced. its usually and stouter persistent notochord continuous through the notochordal odontoid. In both the plesi- osaurs and pterodactyls the atlas and axis are fused. articulating with the the odontoid At other times more or less united with the axis. (Fig. 80 g) and the dinosaurs is (Fig. h). primitively larger than the following intercentra. which a complete disk represents the atlas. The axial intercentrum may be paired or single. Its prezygapophyses are small and turned outward at the base of the spine. 78-81). doubtless because of the loss. to those in which the bodies of atlas and axis are imperfectly or indistinguishably fused (Fig. 80 c). The axial intercentrum nearly always present. 105 reaching the ventral side. and some lizards it has disappeared or is represented by the merest vestige. serving for the attachment of neck muscles.THE VERTEBRAE primitively large size. with no motion be- tween it and the ring of the atlas. and is intercalated between the bodies of the atlas and axis in the usual way. indistinguish- ably so in the adult. chelonians. . It is small in the dinosaurs and 80 G. 80 h) the pleurocentrum atlas axis. and platycoelian vertebrae the front end Its is flattened for sutural or ligamentous union with the odontoid. Axis (Figs. forming pseudo-hypa- pophyses. When paired they are more or less elongated. the in its broader spine. fusion. though in the modern cocodiles (Fig. opisthocoelian. and in its relations with the atlas. The axis differs from the following vertebrae more elongated centrum. fused with the odontoid or apparently absent. or great decrease in the size of the axial inter- centrum. In the Plesiosauria (Fig. is centrum usually longer and usually bears a rib. Among the Chelonia the is may fuse into an independent vertebra. both are slender-necked animals with small or vestigial cervical ribs. more reduced. 80 c). In procoelian. In the Cotylosauria and Theromorpha the front end of its centrum is deeply concave. 80 a) the odontoid extent visible from the side.

In the lat- FiG. for the support of the pectoral arch. One half natural ter order three or more of the anterior ones may be more or less immovably united notarium. In the Chelonia they are fused throughout in the caralargest The number having a in the sacrum. In the chameleon lizards there are as few as eleven.io6 THE OSTEOLOGY OF THE REPTILES Dorsal Vertebrae (Fig. forty-one or forty-two. Ophiacodon mirus Marsh (Theromorpha). in the pterodactyls about twelve. twentieth vertebrae. found in Pleurosaurus a slender. aquatic Jurassic reptile. invariably ten. 82) The smallest number of dorsal vertebrae known in reptiles is that of the Chelonia. 82. Seventh to size. two or twenty-three and In reptiles lack- . from the side. is About is thirty the usual number In terrestrial reptiles the number never exceeds twentyusually about eighteen. plesiosaurs. pace. forming the of dorsal vertebrae in reptiles is .

stout ribs for the support of the pelvis. 84). i). Rarely among the am- phibians are there more than one certain temnospondyls ern urodeles^ are two. A second vertebra (Fig. It is known and modto have quite certain. Sacrum and caudal vertebrae of ^ [Also some frogs. that reptiles began their career with but a single rib-bearing sacral vertebra. that is. thirty-five in the mosasaurs. And this numseries ribs to Fig.THE VERTEBRAE ing a sacrum the the girdles 107 number between may be much greater. vertebrae in front of the sacrum not bearing ribs of any kind. legless lizards. . either fused to- gether or separate. There however. as many as seventy-four in and some Sri terrestrial. however. however. seen from below. there is not same distinction between thoracic and lumbar vertebrae that there is in mammals. 164) of lumbar vertebrae. — Ed. even in the Coty- are. was soon added from the basal caudal by the enlargement of the come in contact with the ilium on each side. inasmuch as Seymouria of the Cotylosauria is known to have no more (Fig. true Sacral Vertebrae (Fig. bearing short.] Macrochelys (Chelonia). examples (Fig. often the As has been said. 83) The sacrum is of land vertebrates composed of from one to four or five vertebrae. 83. losauria.

dJ .

Usually. swimming rep- such reptiles move sinuously through the water. another. Caudal Vertebrae (Figs.tailed lizards (Trachysaurus). however. part. the zygapophyses remain visible and are some- times functional. Those with propelling on the other hand. swift-moving. of use only as a steering organ. but they are not true sacral vertebrae. animals with propelling legs.THE VERTEBRAE ber. are present between the two sacrals. for the (Fig. rule. even the zygosphene and zygantrum The sacrum is lost. for instance. while movements upon The spines of the caudal vertebrae in land reptiles are seldom long. However. crawling reptiles have a long and slender short-tailed reptiles are invariably slow in their land. Stumpy. have three or four From one to three additional vertebrae have been fused with the sacrum in front in the Pterosauria (Fig. certain chameleon lizards and the basilisc lizard are exceptions . sauropod dinosaurs. the . and some Dinosauria. Invariably it is long in tail-propelling. 83-85) The Ohio most tail of the earliest known reptile. Not only may arches and spines the sacral vertebrae be closely fused. in in most and even most the latest mammals dal series. to the present A third vertebra. was long and slender. where hind legs have lost locomotive functions. from the cau- was early united Still many Theromorpha and The Cotylosauria. 76. not only and legless lizards. has 109 reptiles remained persistent time. while other skinks have a very long and slender one. two. have a broader and flatter body and short tail. but their may become almost indistinguishably united. though supposed to be exclusively water animals. from the Coal Measures of had. only a moderately long with not more than sixty vertebrae. 118 d). were joined in the Plesiosauria. have practically no tail. use of the legs as propelling organs. The length it of the tail. in the snakes In Iguana. but also in the mosasaurs and late ichthyosaurs. Dinocephalia and Anomodontia. 84). and possibly two. have a very long slender tail. depends so much upon habits that may be extremely variable even in members of the same order. however. The Cotylosauria tail. preventing much legs. and more or less whiplash-like at the end. tiles. purely aquatic sacrals. As a tail.

causing the easy loss of the distal This septum was once supposed to represent the division of the centrum. from one to six in number. is the widening and elongation of the caudal at first [anteriorly] spines throughout. The cloaca in the living animal occupies the space below them. sometimes with rudimentary chevron-like processes. [less] is and then more distally until a terminal fin developed with the end of the column in the lower lobe (Fig. at least in those rep- with long tails. most lizards. not occurring in the early Chevrons. It is between the primitive component parts embryo. now thought to be an acquired character. are called pygals. of the first indications of tiles habits. out chevrons but with They have the ordinary intercentra in those reptiles in which they [intercentra] are persistent throughout. along which part. the Proganosauria Saphaeosaurus and Sphenodon. and coracoid {cor) oi Geosaitrus (Thalattosuchia). two. 85. Tail. those withribs. 85). scapula (sc). really outgrowths from the inter- centra (Fig.no THE OSTEOLOGY OF THE REPTH^ES One swimming vertebrae distally are more slender and the zygapophyses weak. 84) for the protection of the vessels on the under side of the tail. After Fraas. Fig. or haemapophyses (Fig. The number is more or less reduced in modern reptiles. it readily breaks. occur below and between the caudal centra in . There is an unossified vertical septum through each caudal centrum in many lizards. The basal caudal vertebrae. 76 d). the Crocodilia have but one.

a condition found in the proximal chevrons of Sphenodon. as in the temnospondyl amphibians. may broadly divergent branches united at their base. — they Usually single and Y-shaped — whence • the name be paired in the Plesiosauria and IchthyoThe medial ones of the Sauropoda have two Y-shaped. for the most tail part. in as in some mosasaurs and lizards. replaced by a pair of vertical hypapophysial-like processes (lymphapophyses) Chevrons articulate as a rule intercentrally. III chevron sauria. especially those which the cervical intercentra have migrated forward to articulate or be coossified with the median hypapophysis. have their branches united above in an intercentrum-like bone. At the tip of the they . Chevrons primitively. are vestigial or absent. tigial in the turtles. but sometimes exclusivel)! to the distal part of the preceding centrum with which they may be coossified. In later the two branches articulate separately. More or less ves- they are absent in snakes. reptiles.THE VERTEBRAE most reptiles.

anterior dorsal vertebra. except in the Chelonia. in connection with the segmentation of the vertebrate body.] . middle dorsal vertebra. cal. Cert. at least as far back as the middle of the The first to become fixed or closely united with the vertebrae. Clidastes (Mosasauria). C. and some armoured dinosaurs. which articulates with the end of the diapophysis. Compar. posterior cervical vertebra. or capihilmn.. E. The dorsal ribs are free. The ribs of the Temnospondyli (Fig. from behind. with all vertebrae. 86.-Anat. 86) articulate with intercentrum and arch. Dimetrodori (Theromorpha). the articular surface. Vorlesungen iiber Vergleich Anat. some Ptero- sauria. With the diminution in size of the intercentrum. And this was the original mode among reptiles. usually without differentiation of the articular surfaces. from the side (after Broili). the head. anterior dorsal rib. D. however. and last of the cervi- Vertebrae and ribs: A. from the side. anterior dorsal rib. — Ed. B. like those of the amphibians.CHAPTER III ^ THE RIBS AND STERNUM The late ribs of reptiles. next the lumbar. Ichthyosaurus (Ichthyosauria). after Merriam. primitively articutail. Kingsley. 1921. joins the adjacent ends of two centra across the intervertebral cartilage. Cymbospondylus (Ichthyosauria). Diadectes (Cotylosauria). were the caudal. after all the sacral. continuous to the tubercle. FiG. see Butschli. This continuous articu- lation 1 from the intercentral space to the arch was the almost invari- [For the morphology and variation of the ribs.

centrum. A better for them is holocephalous. the Ichthyosauria. 80 N. the ribs are often dichocephalous (Fig. and Squamata (Figs. 87. the head across the intervertebral cartilage. has continued through those reptiles [see above] and through the mals. and the rib is truly double-headed. the parapophysis. In many reptiles. mam- And this is essentially the mode of rib artic- ulation in the Diaptosauria. Such though usually called single-headed. In the later Ichthyosauria and later Plesiosauria. In the dorsal region there have been many )Z°^ Thaiattosaurus. the head has migrated backward to articulate with a facet or process on the anterior end of the cen- trum. the articular surbecome restricted to the head and tubercle. though connected. and in the dorsal vertebrae of the Squamata and their fig. 86 d). Soon. probably all with a single. however. and the single-headed ribs remained attached more or less wholly to the it is true. there is an emargination of the articular surface between them. ° (After Merriam. and Thalattosauria (Fig. In those reptiles which are here classed under the Parapsida. however. the tuber- cular part of the articulation has been largely or wholly lost. the ribs are described as single-headed in Pleurosaurus. upper temporal opening. So also. Protorosaurus. single-headed and central in the cervical region. in such forms no real a Lower Permian reptile. 73 c-f). both articulations unit- There is. tubercle. imperfectly double- headed in the dorsal region. of articulation of This early mode double-headed ribs. 87). single-headed ribs are those which have name faces lost either the head or the tubercle. that is. 80 o. 86 c). that is.. or dichocephaloiis (Fig. 86 e) and occurs occasion- ally in the ribs. are The ribs of Araeoscelis. Pleurosaurus. and there in the cervical vertebrae of it has remained most reptiles. are not really so since both Theromorpha and even capitulum and tuberculum are present. Strictly speaking. 89) of the .) modifications. The dorsal ribs (Figs. the Proganosauria. the so-called neck. Dorsal verteallies.THE RIBS AND STERNUM able rule 113 among the Cotylosauria (Fig. the tubercle to the diapophysis of the arch. typical. ing with the centrum. in the recent Sphenodon. however. Progano-^ _ sauria. perhaps in part because of the closer articulation of the vertebrae..

The dorsal ribs of the Archosauria. 80 e) and Chelonia 80 m). anterior ones at least. present in all Atlantal ribs. ending in wide. Hist. pz. the Pseudosuchia. in the ancient Metriorhynchus pz.] . 89 r) exclusively with the cen- is. are double-headed. though a vestige often remains. c. Proc. In Cynognathus the thirteenth to the seventeenth ribs shorten rapidly and project widely with a remarkable expansion near the proximal end. articulations . 88). 81). usually also by a single head. Procter. except the Croc- where they are attached exclusively to In the Dino- the intercentrum. expanded and overlapping in ribs in the posterior dorsal series occur some of the Theriodontia. forming more or less of a carapace (Fig. d. anterior zygapophysis. lost in modern ones. that by a single head. and Pterosauria. " Reptiles and Amphibia. Axial ribs are more often present. Dinosauria (Fig. The dorsal ribs of the Eunotosauria ^ Chelonia have expanded to and all meet or fuse with Peculiarly each other. have been odilia. In early crocodiles the axial rib articulated with diapophysis and parapophysis diles Dorsal vertebra of phytosaur: az. but are lost few reptiles. posterior zygapophysis. early reptiles. 1923.114 THE OSTEOLOGY OF THE REPTILES (d) exclusively Sauropterygia articulate with the diapophysis trum. [This leaves out of account the costal plates which enter into the formation of the See Gadow. in not a particularly the Ptero(Fig. In the lumbar series they lose the free portion of the shaft. the Parasuchia (Fig. carapace. 91). in later crocois the diapophysial articulation lost.. — Ed. sauria (Fig. in which the head of the ribs articulates with the centrum throughout. 88. Soc. the cervical ribs (Fig. Zool. 90). but both articulations are with the arch or diapophysis. which overlaps the succeeding rib in a concavity on (Fig. Fig. 92) 1 its anterior border. the first intercentrum bears a small rib (Fig. sauria. some of them at least. Crocodilia (Fig. of rib. and the singleheaded rib has migrated forward on the odontoid. 80 p)." Cambridge Nat. to both arch and intercentrum. And this mode of articulation would seem to exclude their immediate ancestral relationship to the birds.

as Theromorpha. articu- lation for head. are progressively flattened and Fig. remarkably so. The cervical ribs of the Crocodilia (Fig. for tubercle of dorsal rib. />3. in Diadectes and Limnoscelis (Fig. 90. there are three flat dermal plates overlying the following ribs. Fig. in r. and rarely also of the dilated. The four to six ribs of the Cotylosauria. Not only are these ribs so remarkably dilated in Diadectes. anterior zygapophysis. and dorsal vertebrae from in front: «2. d.r. posterior zygapop hysis. Plesiosaur vertebrae: Polycotylus. RIBS ribs AND STERNUM to the greatly 115 Such gave great strength to the lumbar and are perhaps analogous first expanded trans- verse processes of the crocodiles.r. and from /. terior zygapophysis. Vertebrae of gavial from the side (cervical). sometin^es. articulation of dorsal rib. region. for the direct support of the short and broad scapula. but.THE interlocking ends. c. continuing the expansion backward. cervical ribs. front (dorsal): az. r. ancervical rib. 89. posterior zygapophysis. Cervical vertebrae from the side and behind. 95). d. 90) and Dinosauria are . diapophysis. pz.

Fig.ii6 short and THE OSTEOLOGY OF THE REPTILES. 128. i) for instance." either fused or more or ribs of less closely united to arch and centrum. more or less "hatchet-shaped. the tubercular part with the . the capitular part articulating more or less inter- centrally with the preceding vertebra. free ribs continue to the sacrum. 91. four or five vertebrae in front of the sacrum bear no ribs of any kind. 164). Three or four of the single-headed ribs of the Sauropterygia articulating with both centrum and arch are known as pectoral ribs. In certain early cotylosaurs (Figs. True sacral ribs often retain their primitive attach- ments (Fig. Many other reptiles have a variable number of the presacral ribs coossified with the centrum. so-called trans- verse processes. About one sixth natural size. The free cervical Only vestiges of ribs relizards and mosasaurs begin upon the axis. Inner side of carapace of Stegochelys (Chelonia). they are nearly always fused. Seymouria (Fig. main in the pterodactyls and turtles. in others. Sacral ribs. After Jaekel. 93). or centrum and arch.

Fig. 93. from in front. Nothodon lentus Marsh (Cotylosauria). Vertebrae: Cynognathus (Cynodontia). One half natural size. After Seeley. Fig. 92. Sacral vertebra. from above. posterior dorsal vertebrae. 117 . Natural size.

but that term true only of the sacro-lumbars. descended from reptiles with ^ but two sacral vertebrae. as On the other hand. however. while the lumbar or posterior dorsal ribs have often undergone changes. — Ed. and their diapophyses with the ilium.Ii8 THE OSTEOLOGY OF THE REPTILES have been in all cases arch. and Remarkably. since animals their original the caudal ribs have retained more or is less in all attachments.] . and also in the Anomodontia (Fig. though may extend to the ilium. There can be no question. The second added very early in the history of reptiles. but the centra also have retained their primitively amphicoelous structure. The pro- jections from the vertebrae have often been called indiscriminately is transverse processes. that the first true sacral animals^ are identical with the single sacral vertebra Additional sacral ribs have been due to the gradual of Seymouria. It certainly does is not apply to the Amphibia and very doubtful for the Chelonia. In the ilium is these greatly prolonged in front of the acetabulum. as third pairs were growths of the centra (Moodie). 121) primitive in their attachments. 119 c). outilium. It vertebrae of all improbable that there has ever been any is. front. groups that have all been accredited with from seven to ten sacral vertebrae. In the Ceratopsia three lumbar vertefor instance) brae have been fused with the sacrum. In the later pterodactyls there are several such sacro-lumhar vertebrae. additional vertebrae have joined the sacrum in many as three in some reptiles. a primitive type. Not only are the two sacral ribs of the Crocodilia (Fig. Real sacral ribs added from behind. elongation of the basal caudal ribs and their articulation with the shown in the tail of the alligator snapper turtle. ''migration" of the sacral vertebrae. in the Lacertilia evidences of sacral ribs have not been found. Whether tral or not the dinosaurs acquired the third or the fourth sacral vertebrae after their divergence from their immediate ances- stock is perhaps a question. both the Saurischia and the Ornithischia. since the [But few contemporary morphologists would endorse this view. Indeed in some instances {Monoclonius remain on the first. a vestigial free rib may so-called sacral vertebra. But two are accredited to Hallopus. but in such cases the ribs their primitive have not reverted to they attachments if modified. the ilia being supported by transverse processes. but that the allied Crocodilia dinosaurs.

of various forms. Many temnospondylous amphibians have on the hind border of the dorsal ribs an angular uncinate process. they sheath closely the under side of the abdomen. among the Proganosauria. osauria. like that so characteristic of birds. Pterosauria. They are known in Galechirus of the Dromasauria. and Sphenodon of the Rhynchocephalia. they occur more rarely. an ambolomerous type. exterior to these a distinct armor of dermal plates has been observed. of the Stegocephalia. Among the temnospondylous amphibians the parasternum. in greater or lesser ity. though and far from constantly each group. extending on the limbs. in Aigialosaurus and some recent lizards. Homoeosaurus. occur in some members in of every chief group of numbers and complexreptiles. They are arranged in a V-shaped pattern with the apex in front. . if not many. by Gegenbaur and ordinarily known as ventral or abdominal ribs. . They have been observed in only a few of the Theromorpha. Kadaliosaurus. imperfectly ossified procIn esses also occur in the Crocodilia. in the Choristodera. in the Pseudosuchia {Scleromochlus) Phytosauria. They occur in the Jurassic Homoeosaurus and the modern Sphenodon of the Rhynchocephalia. by Baur gastralia. and in many other reptiles they much probability will yet be ossifications. Among the Cotylosauria they are known in three families. and since doubtless the Diapsida began with but two. of the known genera of the order.THE RIBS AND STERNUM II9 and Pseudosuchia have but that number. doubtless remained unossified. and may sheath the whole under side of the body. the Sauravidae. and are certainly absent in some of the families. called body an armature of bony rods. Ossified parasternal ribs. In some cases. found in other reptiles as fused processes or separate Ventral or Abdominal Ribs Many. perhaps most. In Cricotus. Protorosaurus. They are unknown in the Stereospondyli. each composed of an unpaired median piece. Crocodilia. . they are certainly absent in some. Procolophonidae. Captorhinidae. Such ossifications have never been observed among the older reptiles. especially the Branchi- had on the under side of the or plates. Hyperadapedon. Among the Rhachitomi they have been observed in Archegosaurus in the shape of slender rods. Pleurosaurus and Saphaeosaurus of the subclass here called the Parapsida. and numerous lateral ones.

it is seen. meeting in a median. Champsosaurus Three halves natural size.I20 THE OSTEOLOGY OF THE REPTILES sauria. and more than two . bony rods composed of several pieces on each side. (Rhynchocephalia). D. Nyctosaurus (Pterosauria). Ichthyoand Chelonia. reptiles are those of the Cotylosauria and Theromorpha 94 a). B. twelve to fifteen times as numerous as the overlying vertebrae. though frequently absent in forms related to those which possess them. slender. in the Sauropterygia. characteristic of the Reptilia as a whole. They are thus. One half natural size. About one half natural C. of the They sheath closely the whole under side abdomen from the coracoids to the pelvis. Sphenodon The most primitive parasternals known among (Fig. E. Lystrosaurus (Anomodontia). Theropleura (Theromorpha). and Theropoda. V-shaped piece. unpaired. Nearly one half natural size. size. No explanation has yet been given of Fig. (Choristodera). One half natural size. of the Archosauria. Sternum and parasternum: A. 94. their inconstancy.

The have five or six flattened parasternals. consisting of a straight or slightly curved median piece. each composed of two slender rods on each side (not joined in the middle). In the modern Sphenodon (Fig. Clearly endoskeletal in origin. but ble that these are really dermal elements and [not] true parasternals. In the Chelonia they are represented by the posterior it is three pairs of plastral elements. each composed of one broadly V-shaped piece. . In earlier members of the order there was a larger number. nineteen or twenty in number. each composed of a median.THE RIBS AND STERNUM hundred distal I2T number. 121 c) have seven or eight pairs. The Choristodera reptiles. The Crocodiha (Fig. were composed of the usual V-shaped median piece and a lateral splint on each side. Parasternal ribs have long been considered to be of dermal origin. times much more numerous than is These lacertilian ribs are located. (Fig. the posterior ones paired. 94 b) there are about twenty-four such rods. The extinct Saphaeosaurus (Sauranodon) had a full armature of ossified parasternals similar to those of Sphenodon. earlier pterodactyls the splints. it said. as usually accepted. in the rectus abdominis muscles. either connected with the dorsal ribs or cartilaginous. at least. and Ichthyosauria. and usually free. and are united by a dense later pterodactyls sheath of fascia. and three or four lateral splints on each side. Anteriorly they are covered or underlaid by the interclavicle. sometimes paired Only in a few forms have they been ob- served as slender ossifications. In the unpaired median piece has one or two lateral They have also been observed in various genera of theropod possi- dinosaurs. Among the modern lizards abdominal ribs are often present. and some of them. That they are not continuations of the dorsal ribs seems evident from the fact that they are somethe overlying vertebrae. Plesiosauria. the anterior ones broadly V-shaped. aquatic have larger and stouter parasternals. every alternate one of the rib. The parasternals of Sphenodon are in the superficial part of the rectus and external oblique muscles. and have been called distinctively abdominal ribs. they have been supposed to be not true parasternals. The last pair is enclosed in a dense sheath of fascia continuous with the ends of the so-called pubes. in end of the unpaired piece and a lateral splint. 94 c). first eleven attached to the end of a dorsal In the Phytosauria they are similar. espe- cially in the chameleons.

.122 THE OSTEOLOGY OF THE REPTH^ES The abdominal ribs of skin bones which have sunk into the muscles. That they and the sternum other tetrapods are to given origin are really the ends of true ribs which they are supposed to have is improbable. in the Ichthyosauria and Sauropterygia back of the united coracoids and in front of the parasternals. and is thought not to be homologous with the sternum of reptiles. Bulletin. The so-called sternum of the modern amphibians (there was no sternum of any kind in the Stegocephalia) is an ossification of the myocomata. see C. It would seem under side of the body. vol. their an- [For further support of this view. Hist. the lizards are undoubtedly true endoskeletal bones. On either side of the base of the median anterior protuberance it gives articulation to the elongate coracoid. even for a rudimentary one. and that they represent the ends of the dorsal ribs. ^ Sternum The it is earliest recorded occurrence of a sternum or breastbone in rep- tiles is in Anomodontia (Fig. It has been thought that its absence cartilagi- wholly absent. it is more probable that Camp. even as a in these orders 1 is due to. or even more reasonable that the abdominal ribs of all reptiles are of paren- known in chymatous or cartilaginous origin. Amer. 94 e) where. sometimes ossified. xLviii. There was no space. and that the anterior ones fused to form the sternum. sternal Posteriorly in the middle it is contiguous with the parasternal In many reptiles the sternum is nous element. It is figured in Keirognathus as a the distal small. shallow concave bone. Its lateral margins have ribs. ribs. its loss. or outgrowths from them. 389-393] . but without a true keel. since no which the dorsal ribs meet on the approach each other. not derived from the dorsal ribs. Mus. articular facets for four or five. L. and Fiirbringer has suggested that in these animals they are new growths. pp. the best examples of which are the Pterosauria (Fig. supplanting the dermal parasternals which have long since disappeared. it covers the whole under side of the thoracic region with a stout manubrium-like process in front. as a broad. generally present and ossified. 1923. subquadrilateral the coracoids and occur as an ossification bone lying over the posterior extremities of end of the interclavicle. Only rarely does it in other reptiles. according to Broom. Nat. 94 d) where.

also cartilagi- may Rarely. and doubtless the extinct was present in most of members of the order. exceptionally more. The sternum has been found Sauropoda it in not a few dinosaurs. ossified plates. the xiphisternum. In the modern Lacertilia (Fig. probably also the condition in the early Rhynchocephalia. is In the Chelonia there Crocodilia the sternum is no trace of a sternum. cartilaginous plate. It gives articulation on each side to from one to four or nous. 99) and in Sphenodon there is a more or less calcified. even a the Cotylosauria and Theromorpha. thin. Probably this was the condition in all the early reptiles. and the sternum be represented by a pair of small cartilaginous plates or may tions. probably was present as a cartilage. RIBS AND STERNUM 123 small cartilaginous one. Sometimes it has single or paired perfora- Similar cartilaginous sterna have been found in the it DoH- chosauria and Mosasauria. oval. Among the has been recognized in a pair of oval.THE cestors never possessed it. since since in several instances in Rhynchosaurus had parasternal ribs reaching nearly to the coracoids. sternal ribs. contin- ued into a pair ribs. . itself five. be wholly absent. of cartilaginous xiphisternal rods to which dorsal ribs are attached by the intervention of of the cartilaginous sternal Nothing is known it sternum in extinct crocodiles or phytosaurs. rhomboidal plate articulating on each side with the coracoid in front and ending in a single or paired continuation. among There could have been no sternum. both these orders the parasternals have been found sheathing the whole abdomen from the coracoids to the pelvis. no ribs articulate with the sternum. leaving little or no space for a sternum. In the hving six or eight a small.

Pectoral girdle. Diadectes (Cotylosauria). or collar-bone. which early unite with the scapula to form the coracoid process. blade.CHAPTER IV THE PECTORAL AND PELVIC GIRDLES The Pectoral or Shoulder Girdle (Figs. pectoral or shoulder girdle. on each side. or shoulderclavicle. as in that of most mammals. and the there are but two on each side. or possibly two. right side. the scapula. A third bone. In our own skeleton. 95-113) Those bones which form the framework for the support of the an- terior extremities in vertebrate animals are collectively called the Cledkrum'^ lltetacorobCQidj irrf-erclavtcte Fig. In 124 . are represented in most mammals by mere vestiges. 95.

originally is named epicoracoid by Cuvier. Varanops (Theromorpha). from below. C. One half natural size. there are in addition to the clavicles three well-developed bones on each side. The homologies of these. Cacops (Temnospondyli). or rather of . the scapula and two bones articu- Pectoral girdles: A. from above.THE PECTORAL AND PELVIC GIRDLES the lowest living 125 mammals. generally known as the procoracoid. One half natural B. lating with it at its lower end. One half natural size. the anterior of which. D. 96. the posterior one helping to form the articulation for the arm bone. known as the true coracoid. of which Ornithorhynchus and Echidna are the only examples. Diadectes i^). 6'«'7OT0«r»<j (Cotylosauria). from above. One half natural size. size. from below. FiG.

in rep- at least. Cleithrum.126 THE OSTEOLOGY OF THE REPTILES There the epicoracoid. the pectoral girdle is composed of eleven separate and distinct bones. 95. Primitively (Figs. The coracoids. forming more or less of a pectoral buckler — adaptation to their water habits. came subdivided — . Therapsida. or ordinary reptiles. after long existence as a mere vestige. 96 a) may be taken as types. the subdivision Ed. 96 a. though a vestige may possibly be present in our own and anterior one of the two shoulder girdle. a relic from the fishes. are yet doubtful.] occurring only in the Theromorpha. The girdle in the adult land forms. is also a median. the cleithrum is re- duced. and will be discussed later. The posterior two coracoids also disappeared in late Triassic times. of which Eryops (Fig. though much reduced and scapula. the coracoid originally was a single piece which never bein the amphibians.i all of endoskeletal origin. and mammals. Clavicular Girdle The clavicular girdle is variable among the temnospondyl phibians. disappeared in Triassic times. unpaired bone in these mammals. The cleithrum so generally characteristic of the Stegoin reptiles. the interclavicle. In the aquatic types of all Stegocephalia the clavicles and interclavicles are rugose [on the ventral side]. perhaps in a measure analogous to the plastron of the turtles or the extraordinary development of the coracoids in the plesiosaurs. 96). and three bones on each scapula and two coracoids. together composing the the secondary or clavicular girdle. as in reptiles and higher vertebrates. at least in early life: the median interclavicle and a clavicle and side. 108) and Cacops (Fig. amupon the habits of the animals. non-functional in some lizards. except that the cleithrum is larger and the interclavicle less elongate. cotylosaurs. cephaHa 1 (Figs. composing the primary or scapular girdle. unknown in other mammals. clei- thrum on each side. all live of dermal origin. is almost indistinguishable from that of their contemporary cotylosaurs. The cleithrum of the tiles (Fig. In such forms also. the so-called procoracoid. the more terrestrial form. are still present in most reptiles in snakes only are they wholly absent. that is. clavicles. 95). They are smooth throughout in Cacops. heavy a peculiar and broad. in the oldest known reptiles. dependent. 108) was doubtfully ever functional [According to Watson.

It is vestigial in some forms and seems to be quite wanting in others. Clavicles and interclavicle of Ophiacodon (Theromorpha). and either absent or Pterosauria and Dinosauria. 95). and Anomodontia. and in certain was known only members of the Cotylo- Theromorpha. 98) as a rod-like bone at the upper Fig. with a dilated mesial extremity.THE PECTORAL AND PELVIC GIRDLES whatever may have been never large. Mosasauria. where its somewhat spatulate upper extremity partly overlies the front. Among the Theromorpha it has been observed in Edaphosaurus (Fig. It is 127 it its function in the amphibians. articulating on the ventral side of . 97. best developed perhaps of all in Diadectes and its allies of the Cotylosauria (Fig. but none is demonits Its vestigial condition in various cotylosaurs indicates entire disappearance. sent in the Crocodilia. Clavicles are usually present in reptiles. sauria. front border of the scapula. Dinocephalia. clearly a vestige. 96 B-99) they are usually curved bones. articulating below with the stem of the clavicle. and or less vestigial in They are absome Sauria. serpents. (Fig. what has become of it. Clavicles. upper border of the scapula. In crawling reptiles (Figs. more vestigial in the some lizards. There have been several theories as to strable. 107 d) it is In the Anomodontia and Dinocephalia a feeble splint.

fifths Pectoral girdle. firmly united with the scapula and with each other. clavicle. loi) they are normal but very stout. and also with the lower end of the cleithrum when that bone is present. In the Ichthyosauria (Fig. The inner end in some lizards is broad and perforated (Fig. Fig. or its acromion when present. 103). triangular bones. scapula. cl. The clavicles of the Plesiosauria (Fig. The clavicles of the Chelonia are known as the epiplastra of the plastron (Fig. sc. of the interclavicle and sometimes also of an anterior process from the coracoid. Edaphosaurus novomexicanus (Theromorpha). two natural size: c. cleithrum. they are slender.128 THE OSTEOLOGY OF THE REPTILES . 98. 100). 99 c). . 102) are remarkable in some Usually they are a pair of thin. be absent. In the Nothosauria (Fig. 104). they may respects. nor are they expanded mesially in either the Phytosauria or Choristodera (Fig. lying upon the inner or visceral surface of the proscapular process of the scapula (corresponding to an acromion). and all water reptiles. sometimes coossified with each other. 99). In modern lizards the clavicles articulate usually with the front border of the cartilaginous suprascapula (Fig. the end of the interclavicle and a more or less slender stem which articulates with the front border of the scapula. Doubtful vestiges of the clavicles have been reported in the pterodactyls.

Natural size. 100. 99. Zonosaurus (after Siebenrock).Fig. After Jaekel. 129 . Primitive chelonian pectoral girdle: Stegochelys. B. Pectoral girdles (Lacertilia): A. Fig. C. Iguana.

coracoid. . ic. clavicle. Pectoral girdle oi Nothosaurus (Nothosauria). 102. from photograph by E. Pectoral girdle of Trinacrotnerum (Plesiosauria). interclavicle. from above: cl. c. scapula. cl.Fig. ioi. cor. sc. scapula. sc. interclavicle. Fig. Fraas: id. clavicle. coracoid.

Interclavicle. c.THE PECTORAL AND PELVIC GIRDLES 131 Fig. as usual with all later reptiles. Pectoral girdle of Ichthyosaur. 104. an elongate bone with a dilated but not T-shaped anextremity. In the is later cotylosaurs the front end In more is dilated. After Brown. 94 d). the interclavicle is forked in front and somewhat fanlike that of the shaped behind. ^'°^''' Natural . Baptanoden {Ophthalmosaurus). Fig. 103. The stem underlies the approxiinterclavicle in the The mated mesial borders of the coracoids. Pectoral girdle of Champsosaurus (Choristodera). In a specimen referred to Pantylus (Fig. After Gilmore. 107 c) the shape usually like that of the It is Theromorpha and F'°^°^- Cotylosauria. clavicle («v/) ^"^""^ size. very short and fan-shaped in P""^^^ and cor- (Cotylosauna): inter- Broom Lystrosaurus of the Anomodontia (Fig. shaped very much monotremes. 105). usually extending beyond them. 96 B. d) is terior earhest-known reptiles (Fig. a primitive cotylosaur. where attributes its reduction to water habits. the known forms of the Therapsida (Fig.

in of the terrestrial types of the is prac- tically indistinguishable tiles.] Journ. the posterior often found separated (Fig. 96 b. The served. In no other reptile has the metacoracoid been certainly observed. 39-47.132 THE OSTEOLOGY OF THE REPTILES it is In the Chelonia anterior end. — Ed. loi). d) fuse in maturity. the called the and two ventral ones. it has been affirmed in the Rhynchocephalia (Hyperodaerror. 7. like that of the aquatic broad and short. the anterior one commonly procoracoid. (Fig. Among most Therapsida the if three bones (Fig. 107 a. 96 d) tilaginous throughout till remained car- life. Nothosauria When present in the plesio- saurs it is forate. and some plesiosaurs. coracoid. Scapular Girdle The scapular reptiles. in some forms (Fig. has never been observed. all. 102). the Eunotosauria of the Upper Permian. Among is the Theromorpha. c) the union invisible later in life . but that side. and a posterior one. from the girdle contemporary repreptiles. foramen The interclavicle is absent in the Pterosauria. 106). in the earlier forms imperin the later ones with a median interclavicular notch or (Fig. vol. 191 vol. or it united by a loose suture. is Each both the amphibians and early less closely fused: com- posed of three bones more or scapula. 108) are so firmly coossified that their sutural distinctions have rarely been ob- Among became the Cotylosauria (Fig. xxiv. or their sutural divisions was less have occasionally been observed. the metacoracoid. 107 d). loo. is girdle. and in all forms it probably did not ossify of the growth was far advanced. an [For a different view of the fate of the two coracoids see Watson. 103). . not In the Dinocephalia (Fig. the two bones are distinct. 1922. 01. In the Proganosauria the divi- sion between the two bones. b. an oval or triangular bone. firm. three bones of the land Stegocephalia (Figs. or scapulo-coracoid of the aquatic temno- spondyl amphibians of early Permian times. if present. posterior bone The -^ was lost in all reptiles by the close of Triassic times. pp.) it is In the Croco- dilia (Fig. but not in some. 96 a. vestigial in the the entoplastron (Fig. though pedon) 1 . Anat. AnaL. 121 d) and Mosasauria slender and free at the The stem is short in the Ichthyosauria (Fig. Romer. Dinosauria. often called metacoracoid. Record. chameleon lizards. a dorsal one.

and that there was a progressive separation and delayed ossification of the posterior bone in the Fig. formerly placed among the Rhynchocephalia. Paleontological evidence that it is the posterior bone . {sc). Dimetrodon (Theromorpha): scapula and metacoracoid {mcor).THE PECTORAL AND PELVIC GIRDLES It is probable that the three 133 bones early acquired a firm union. This doubtless explains its absence in all known specimens of Paleohatteria. It is known that in Ophiacodon from the Permocarboniferous. 106. ossification of the metacoracoid did not occur till late. line leading toward the modern reptiles at least. both ontogenetically and geologically. and that in Varanops (Fig. coracoid {cor). 96 d) it never ossified.

107. " ib) The true coracoid. and generally are considered to be. Whether this conclusion is the right one so far as the monotreme mammals are concerned is still a de- FiG. though Gregory. a sheet of bone lying immediately above the clavicles. now seems complete. C. lying behind the . the so-called procoracoid. The two coracoids in these mammals seem. batable question. Natural size.134 THE OSTEOLOGY OF THE REPTILES which has functionally disappeared in all modern reptiles. and never reaching the glenoid surface. that lost in the reptiles. was retained. The coracoid of Hzards. B. lizards and monotremes. or so-called procoracoid. D. the metacoracoid. Broom has suggested that in the evolution of the is. however. He thinks that three elements are involved in the problem of their evolution : "(a) The epicoracoid of Sphenodon. Moschops (Dinocephalia). homologous with those of the early reptiles. Ga/^o/)J (Dromasauria). G«/^/)«j (Dromasauria). A. One fifth natural size. Pectoral girdles (Therapsida): Gtf/^fA/r«i (Dromasauria). has offered another solution of the problem that would homologize the anterior or "procoracoid" of the reptiles with the posterior bone of the Monotremata. Natural size. mammals the posterior bone. and who proposed the name metacoracoid for the posterior bone. and not a fusion of the two. and Sphenodon is homologous with the anterior of the two bones. crocodiles. It was Howes and Lydekker who first reached this conclusion. About three fourths natural size.

and its ossification in mammals would be nothing remarkable. and is thence directed forward and upward to terminate the procoracoid in the front border. the Pterosauria. is in the coracoid of later reptiles absent in the Chelonia (except the Triassic Stego- chelys). then modern reptiles have no true coracoid. 96. all it In known reptiles possessing a metacoracoid. ria {Howesia). The supracoracoid foramen. and the bone so called that Gregory's theory ogies are must be known as the procoracoid. the terms procoracoid for the anterior bone. Plesiosauria. 99 b) with that of monotremes will show their identity in relations. always present (Figs. in the The scapula-procoracoid Theromorpha (Fig. and the Thalattosauria — chiefly . It passes directly in- ward to terminate in the free border. 107). A comparison coracoid of lizards (Fig. 96 d). lost in later reptiles (Williston). 113). 106) at least. metacoracoid for the posterior are adopted in this work. Ichthyosauria. 106. until the real homolfully determined. the suture sepa- from the procoracoid enters the glenoid fossa (Fig. in though not in the epicoracoid of the monotremes. 106). 100. originally pierced 135 by the coracoid foramen. The author believes is more probable. all suture. where it jcins the scapular rating suture a little in front of the articular surface. Should it eventually result that Broom's theory is the correct one. primitively forming at least the front part of the glenoid. and usually present (Figs. Cotylosauria and divides nearly equally the glenoid surface in front of the metacoracoid. Rhynchosau- many Phytosauria. 112. originally forming the back part of the glenoid region. except in certain therapsids (Fig. the posterior one of which presumably represents the chief ossification of the coracoid process of higher mammals. but neither have found in numerous of the epi- other bones in the mesenchyme of mammals. 95. and to save confusion for the present. 107). often articulating with the sternum." It is true that such an element as the epicoracoid has not been the early reptiles. But. And doubtless a similar epicoracoid filled in the interval between the coracoids above the clavicles and interclavicles in the early reptiles (Fig.THE PECTORAL AND PELVIC GIRDLES clavicles. "(c) The metacoracoid of Permian reptiles. 99. that both coracoids have re- mained in the Monotremata. and in mammals ossified except when preserved as a vestigial element.

already mentioned. in some.136 water THE OSTEOLOGY OF THE REPTILES reptiles. Pectoral girdle of Eryo/)^ (Temnospondyli). be represented by a notch between the scapula and inal position. 108. in). 108) amphibians has three foramina piercing ramen. doubtless its orig- Fig. 103. coracoid (Figs. The scapular girdle of the terrestrial temnospondylous it: (Fig. It may. it is seen. Two thirds natural size. entering a the supracoracoid fo- little in front of and below the .

entering the glenoid fossa and opening on the inner side in front of the subscapular fossa. however. Its external orifice. 96 d). they were approximated along their mesial borders (Fig. 96 d). it is on the outer face of the scapula in front of the border. 109 a). permitting motion of the ventral and antero-posterior. and in Sphenodon. this emargination is very variable in the mosasaurs. In the pterosaurs (Fig. 95). Doubtless epicoracoid cartilages occupied the interval in front. diameter. oval cavity. It will probably be found in many is other forms when searched in the for." In most other is a simple. convex in the conjugate. 99). The supraglenoid foramen present in the Cotylosauria (Fig. so far as has much. In the modern lizards there are emarginations of the mesial border (Fig. does it enter the supraglenoid fossa back of the border. Chelis onia (Fig. even been observed. according to Dou- articulation reptiles it In the early cotylosaurs and theromorphs (Fig. 112) When the sternum border. are extraordi- narily developed in the Plesiosauria (Fig. In Ophiacodon only. as in many modern reptiles. concave in the dorso ventral. is present the coracoid articulates with anterior lateral The coracoids. as shown by many specimens in which they have been found in place. Turned inward at nearly a right angle from the plane of the scapula. the glenoid foramen. 102). A small artery traverses it.THE PECTORAL AND PELVIC GIRDLES 1 37 glenoid fossa and opening on the inner side at the lower end of the subscapular fossa. dorso- The double coracoids are never elongated transversely. The coracoid of the Pterosauria (Fig. In the single coracoid of later reptiles the glenoid articulation has been completed from behind. The glenoid foramen has not been observed in reptiles. and the supraglenoid foramen entering the supraglenoid fossa near the hind border and opening at the upper end of the subscapular fossa. arm in two planes only. 106) the glenoid is more or less spiral or "screw-shaped. 99). in most modern Lacertilia (Fig. presumably the precoracoids only. varies Theromorpha. more usually. It has also been observed in the procoracoid of the elongate. Theromorpha (Fig. probably the Therapsida. 109 b). its theromorphs. thitt. where they sheathe . and Crocodilia (Fig. the deeper one opposite the supracoracoid foramen. 109) it is saddle-shaped. at a variable distance above the glenoid surface.

apparently a specialization (Fig.138 THE OSTEOLOGY OF THE REPTILES the whole under side of the pectoral region. 109. no). In most reptiles the single coracoid is fused with the scapula in . One fourth After Jaekel. size. Pectoral girdles: A. Stegochelys (Chelonia). B. but in the Fig. natural size. Testudo (Chelonia). D. meeting in a firm median symphysis. Elasmosauridae broadly separated posteriorly by a deep emargination. Nyctosaurus (Pterosauria). One One half natural half natural size. C. in most plesiosaurs throughout their lengths.

and rhynchocephalians. in). 102. 113). phytosaurs (Fig.THE PECTORAL AND PELVIC GIRDLES adult life. and inward. The blade is of a strong proscapular process projecting . and often meeting its mate symphysis. no) is peculiar in the downward. The scapula of the plesiosaurs (Figs. 139 but it is free in the crocodiles. development in a median forward. a character unique among vertebrates. dinosaurs (Fig. and more or less suturally loose in the early pterosaurs.

its directed inward and backward. the shorter and is broader are the scapulae. A distinct angular process on the front margin of the scapula in the Cotylosauria (Fig. the shorter and stouter are the legs. The scapulae is of tail-propelling aquatic reptiles are short and broad. is In Stego- chelys. the proscapular process. In upright-walking reptiles the scapula . proscapular process small (Fig. and interclavicle. 102) extends forward to articulate with the proscapular process or with the clavicles. d). Enclosed within the thoracic cavity has two rather slender branches. mammal-like forms from South Africa. c. Formerly was also supposed to be a separate plesiosaurs. The scapula Chelonia also peculiar (Fig. it 109 b. 106). the . used in the antero-posterior of the paddles. that has been referred to the Chelonia in a wide sense.I40 THE OSTEOLOGY OF THE REPTH^ES little short and small. and on the strength of it a relato tionship was found with the and dilated at is now known is be an less exogenous process of the scapula. one extending toward the roof. It is ossification. corresponding acromion and not to the procoracoid. of service for muscular attachment. corresponds to the acromion. however. and the two coracoids approximating their mates in the median line. fan-shaped (Figs. In general. and downward movement A clavicular process of the coracoids of the later plesiosaurs (Fig. the pectoral girdle is of the primitive type. Probably the great development of the ventral elements of the pectoral and pelvic girdles in the plesiosaurs implies greatest de- velopment of the ventral muscles. A distinctly differentiated among acromion process occurs in reptiles only the Pariasauridae and especially the therapsids. as was once thought always of the to do. In Eunotosaurus a Permian genus of South Africa. flattened its The coracoid It is more or extremity. springing from near the articular this process fossa. a Triassic turtle. as The mode of shown by Andrews. c) and Thero- morpha (Figs. 85. 96 b. the other. unlike the scapulae of tail-propelling aquatic reptiles. and connected with mate by ligaments. having a moderately long scapula. slender clavicles. 100). development proves that to the of the proscapular process. 96 d. to which the clavicle is attached. fused with the scapula. it it is an exogenous process of the scapula. is directed downward and inward to be attached by ligaments to the interclavicle or entoplastron. 112). the pro- coracoid.

Morosaurus 141 . C. B. Allosaurus (Saurischia). After Lambe. Pectoral girdles (Dinosauria): A. Gor^wa«r«j (Saurischia). Fig. 113 Fig. size. 112.Fig. About one twelfth natural Fig. (Saurischia). After McGregor. 113. Scapula and coracoid of Rutiodon carolinensis. D. Triceratops (Ornithischia). After Gilmore. Ill Fig. 112 Fig. After Marsh. size. an American phytosaur. One sixteenth natural After Marsh. One sixteenth natural size. III. One twenty-eighth natural size. Scapula {sc) and coracoid {cor) of gavial (Crocodilia).


in bipedal

more elongated,

forms slender. The scapula of the Cotylois

sauria (Figs. 95, 96, b, c)

relatively short

and broad; that

of the


(Figs. 98, 106)

more elongated, but never narrow; that
of the

of the therapsid reptiles (Fig. 107) relatively narrow, slender in the

The scapula



113 d)



tively long, that of the Predentata (Fig.

13 c) is

much more



it is

most slender and

bird-like of all in the


113 A, b).

The scapula

of the Pterosauria (Fig. 109)

always elon-

gated, very slender and bird-like in


of the earlier forms,


and firmly fused with the coracoid

in the latest.

In the most

specialized of all pterodactyls (Pleranodon, Ornithocheirus) its en-

larged distal extremity articulates with the fused spines of the dorsal
vertebrae, the only

known examples among vertebrates

of the articu-

union of the pectoral girdle with the spinal column.
In the Crocodilia, Pterosauria, and
it is

In the early reptiles the scapula was more nearly erect, or with a slight inclination backward.

as also birds especially,

very obliquely


the upper end turned backward over the ribs.

The Pelvic or Hip Girdle



pelvic girdle or pelvis, in reptiles, as in other air-breathing veris



of three

bones on each



or less firmly

coossified in the adult,
girdle is




as the innominate; the

completed by the sacrum on the dorsal side with which the
never closely united in



not even in the Pterosauria.

The upper or dorsal bone

of the three, that to

which the

sacral ribs or

transverse process of the lumbar vertebra are attached,
the one on the lower or ventral side in front

the ilium;

the pubis; that on the

ventral side behind


the ischium.


the outer side, where the

three bones meet, there

a cup-like depression, sometimes a hole,

the acetabulum, for the articulation of the thigh bone. In only two

groups of

reptiles, the Crocodilia

and Plesiosauria,


the pubis ex-

cluded from union with the ilium.
lizards there are at

In the snakes and snake-like
of the pelvic bones.

most only vestiges


pelvis of the terrestrial temnospondylous amphibians (Fig.

14 a)

almost indistinguishable from that of the contemporary

cotylosaur reptiles in early Permian times.



ilium of the rhachi-

tomous forms


not dilated above, as in the

distinction fails in the

more nearly


reptiles, but even this embolomerous Cricotus, in reptiles.

which the ilium


prolonged backward, quite as in the


pubes and ischia meet in a close symphysis without openings of any kind, except the pubic foramen, a small hole through the pubis below

One half natural size. Fig. II4. Pelvic girdles: A, Cacops (Temnospondyli), from below. B, Seymouria (Cotylosaur), from below. A little more than one half natural size. C, D, Varanops (Theromorpha), below and from the side.

the margin of the acetabulum, in front of the ischiatic suture, for the passage of the obturator nerve. This " plate-like " structure of the pelvis is characteristic of the Cotylosauria (Figs. 114 b, 115),

and more

Therapsida (Fig. 119), Proganosauria, the Choristodera, and early Rhynchocephalia.
or less of the
(Figs. 114 c, 117),



1 1


Limnoscelis paludis.

Diagram of


from the

ilium; pi^, pubis;


Fig. 115. Limnoscelis paludis (Cotylosauria). Pelvis, from below. Two fifths natural size. Cross-section

through pubes at

a\ cross-section

through ischia at


Fig. 117.


A, from the schium.

side; B,

(Theromorpha). Pelvis. One half natural from above, pu, pubis; /'/, ilium; is,





small opening soon appeared where the four bones meet below

in the


(Figs. 114 c, 117),

and increased

in size,



Pelves and sacrum: A, Varanus (Lacertilia), from the right. B, Erythrosuchus (Parasuchia), from the right. After Broom. One tenth natural size. C, Rutiodon (Phytosauria), from below. After McGregor. One eighth natural size. D, Nyctosaurus (PteroFig. 118.



D", prepubis of

sacrum and right innominate bone from within; D', anterior parasternal the same from below.




Pelves (Therapsida): A, Galechirns (Dromasauria). After Broom. Nearly B, Diademodon (Cynodontia). After Broom. About one half natural size. C, GalepHS (Dromasauria). After Broom. Nearly natural size.




reptiles, since Triassic

times at least, this pubo-ischiatic open-

ing extended on each side nearly to the acetabulum, leaving only a

narrow connection between the pubis and ischium

(Fig. 118).




the symphysial ends of the pubis and ischium became connected in

many by Hgaments,

or cartilage (Fig.


later in

some by

Pelvis and sacrum. A, Iguana (Lacertilia), pelvis from below; B, sacrum from below. About natural size. C, Dicynodon (Anomodontia), pelvis, from above and from the side. After Broili. Nearly one half natural size.
Fig. 120.

bone, producing a false obturator or thyroid vacuity on each side.


foramen or vacuity homologous with that in mammals, the so-called

obturator foramen, that
the real


between the pubis and ischium with which obturator or pubic foramen is merged, occurs in the Therioof a thyroid vacuity in the theriodonts

dontia (Fig. 119), Anomodontia, and later pterodactyls (Fig. 118 d).

The formation

may be due


the gradual increase in size of the pubic or true obturator foramen
recession backward, as in the Dromasauria,
till it

finally lies

Fig. 121.

Pelvic girdle and sternum: Alligator (Crocodilia). A, pelvic girdle, from the

right; B, the

same, from above, showing sacrum; C, the same, from below, with


D, sternum and



half natural size.

between the two bones, the pelvis




primitive plate-

hke character with only a small median pubo-ischiatic vacuity. But this will not explain the thyroid vacuity in Pteranodon and Nyctosaurus of the Pterosauria (Fig. 118), since
it is

inconceivable that

these reptiles had an unbroken descent from forms without a median vacuity. In no reptiles is the pelvis more aberrant than in the Crocodilia (Fig. 121). So characteristic is its structure that it at once distin-

or is closely approximated to.148 THE OSTEOLOGY OF THE REPTILES all guishes the order from others. though this has been disputed for some. ibid. are either paired. The so-called pubis It probably homologous with the prepubis of the bipedal reptiles pro- pterodactyls. very The ischium and pubis are closely united into a with a thyroid foramen. They articulated with a tuberosity on the front margin of the pubes and in all probability were continued in front with a ligamentous sheath that enclosed the parasternal ribs. Romer.] . like that of duced anteriorly by the broad plate. If the true pubis of Crocodilia has become vestigial and the prepubis has become the functional pubis. or with a small pubic foramen below the acetabulum. thin. Bulletin. how did the prepubis capture the system of muscle its attachments of predecessor? Ed. though sutures have not been observed. Nat. ilium of the Pterosauria. The pubes and ischia meet in a symphysis below. Its anterior process. With much reason has long been urged that the anIn early terior projection of the ischium represents the real pubis. 1923. p. spatulate extremity The ischium a rather long bone. Amer. as in Rhamphorhynchus. as in Rhamphorhynchus.. its mate only at its inner anterior corner. with a thin but strong plate it of fascia joined to the para- sternal ribs. 1918. with a thin and dilated anterior extremity which touches. In front of the acetabulum it is produced forward to join ligamentously with an anterior process of the ischium. The ilium is a strong bone firmly united with the two pairs of stout sacral is ribs. often called the real pubes. anteriorly. which may be in part the real pubis. has no pubic foramen. with a in a joins its mate median symphysis. 118 d). of which the posterior the larger. or united in a ventral band. Below. to form the acetabulum. much 118 d) and Pteranodon. and is continuous in front with the so-called pubis. ^ life it is largely cartilaginous. proving the normal structure of the pelvis. 606). Mus. articulates This bone is slender. either sides of the vertebrae. enclosing between them a foramen which of considerable size for the passage is of the obturator nerve. the ilium articulates with the ischium only. all is The forms. Pteranodon and Nydosaurus (Fig. As remarkable as the pelvis of the crocodiles is that of the Dino- 1 [The pubis of the Crocodilia gives attachment to a series of muscles which as a whole are homologous with those that are attached to the true pubis in Sphenodon and lizards (Gregory and Camp. Hist. but is becomes fully ossified in the adult. as in so in some more or less Nydosaurus (Fig. as in Pterodactylus.. The prepubes.

or rather of that division called the Predentata. much elongated in the Theropoda. E. or pre- . In the other divisions. After Marsh. B. Ceratosaurus (Saurischia). the Theropoda (Fig. 122 b). pubes have the normal reptilian struc- though unusually stout and strong. One twentieth natural size. Apatosaurus (Saurischia). D. in a firm symphysis. : Each is have been the subcomposed of two pro- jections or processes the anterior one. meeting in the middle below Fig. One sixteenth natural size. One twenty-fourth natural size. One thirty-second natural size. 122. After Marsh. C. 122 a) and (Fig. The sym- physis of the ischia is less The pubes ject of of the Ornithischia (Fig. Pelves (Dinosauria): A. or order Ornithischia. the Sauropoda ture. One After Marsh. the so-called prepuhis. 122 c-e) much dispute and speculation. tenth natural size. Stegosaurus (Ornithischia). Triceratops (Ornithischia). After Marsh. Trachodon (Ornithischia). strong.THE PECTORAL AND PELVIC GIRDLES 149 sauria.

When this peculiarity of by Hulke and Marsh ancestry of birds. however. The is long and slender. of the dinosaurs. Ceratopsia. is less prominent pectineal directed forward. that is. unusually stout in the quadrupedal Stego- saurus. homologous with the pubis of the Saurischia. is observed in many running birds. corresponding to the pubis of birds. Whence it would appear of birds in its that the development of the two processes in the dinosaurs has and cannot be ascribed to a was once thought. According to another view. merely another of the many parallel characters brought about by similar causes. the postpubic process is the real pubis. It it the dinosaurian pelvis was first discovered was hailed as a direct proof of the dinosaurian may be. arising. typically directed forms. however. a more or process. the prepubic process is the real pubis. backward immediately below. which have certainly descended from bipedal postpuhis. An analogous but not homologous structure etc. as clear. from the ilium. however. According to one view. and not the even more bipedal theropods. Why the bipedal predentate common dinosaurs should have acquired such a remarkable structure of the pelvis. slender.. typically flattened and more or less spatulate distally. is directed forward and downward [upward] and does not join its mate tigial in a (Ankylosaurus) median symphysis. The postpubis is vestigial in the heavy quadrupedal Ceratopsia. Geococcyx. There has never any evidence to show that it is derived from a sepa- rate center of ossification.150 THE OSTEOLOGY OF THE REPTH^ES pubic process. At times it may be small or even vesbut is broad and stout in the quadrupedal . the pelvis is more or less open below. where. the prepubic process homologous with the prepubis of pterodactyls or crocodiles. It has been ascribed to differences in the posture of the tail in running. posteriorly directed pubis similar to the post- pubic process of the dinosaurs. heredity. the slender ischium and not meeting its mate in a symphysis. where apparently it again functions as the normal pubis. is not yet entirely arisen in response to similar causes. like that The pubis its turns backward from embryonic development normal position. however. been. rather to have been . the ostriches. in addition to the normal. or postpubic process. to the author at least. It is. but would seem. the postpubic process an outgrowth from it. as in birds. possibly as a reinforcement to the ischia in the support of the heavily armored body.

In the earlier ichthyosaurs the broad ischia and pubes were separated by the broad pubo-ischiatic opening. 124) the pelvis. of the usual type. Pelvis of A^oMoJ««r«j (Nothosauria). as usual. but firmly connected. as in the birds. as in The little.THE PECTORAL AND PELVIC GIRDLES due to differences in procreational methods. 124. from below. the 151 of the open pelvis predentates permitting larger eggs to be extruded. Fig. ichthyosaurs. 123. Fig. and the pubes and were united without a pubo-ischiatic opening. in other like that of the water animals. The narrow ischia and pubes meet sym- physis. After Andrews. the pelvis was reduced. The slender ilium of the mosasaurs. and the pelvis was con- nected with a sacrum. and there is a pubic foramen. shows only a moderate aquatic adaptation in the broad pubes and ischia. that the acetabulum of all dinosaurs is perforate. In the later forms. 'PtW\s oi Platecarpus (Mosasauria). In the Nothosauria (Fig. It may be added the birds. the rod-like ilium lay loosely in the ischia flesh. . lay loosely in the flesh with its upper end in apposition in a or ligamentously connected with the end of a transverse process or rib of a single vertebra. pelvis of amphibious or aquatic reptiles It lost all connection is also modified not a with the spinal column in the Mosasauria is and later Ichthyosauria. however.

152 THE OSTEOLOGY OF THE REPTILES ilium is The firmly connected with the sacrum. is and there is a pubic foramen. articulates at its distal extremity with the 125. 125. Upper Cretaceous //. plesiosaur. Fig. Pelvic girdle of Trinacromerum osborni. an from above: />. the pubo-ischiatic notch small. In the Plesiosauria (Figs. ilium. ischium. ihum. connected ligamentously with a sacrum of three or four vertebrae. pubis. 126). the slender . is.

A.Fig. 153 . Fig. ilium. Pelvis: Testudo (Chelonia). Pelvic girdle oi Elasmosaurus (Plesiosauria): p. from the side. 127. pubis. from below. il. ischium. is. 126. One half natural size. B.

where supported by the united or contiguous diapophyses of the false sacral vertebrae. from the primitive closed and plate-like type. often divided in the middle by a cartilaginous septum. Ceratopsia (Fig. is no separate pubic foramen or notch. as in mammals. secondarily plate-like. 122 such forms have short toes. like that of the plesiosaurs. — prob- ably separated by a ligament in In some genera. has its been modified by is peculiar relations to the carapace and plastron. and. velopment of a vacuity tion of the anterior process of the ilium. are and flat. a. The pubes and ischia. in the Pleurodira. is coossified with it.154 THE OSTEOLOGY OF THE REPTILES is ischium only. is no. like the coracoids. producing a false thyroid foramen with which the obturator foramen is confluent. like that of the Chelonia. by the elongaand by its closer union with additional true sacral or lumbar vertebrae. or. the two bones are secondarily broadly united at their symphyses. rarely absent The ilium. it is unusually long in the Anomodontia (Figs. as in the plesiosaurian Sthenarosaurus. there is in reptiles. directed upward and very broad backward. like the pectoral girdle. b) as also in more or less some Coty- Therapsida (Cynognathus). The ischia are triangular or "hatchet-shaped. 122 and Pterosauria (Fig. As in the plesiosaurs. sometimes at the entire expense of the postaceIt is tabular process. possibly of the gluteal muscles. and usually the large pubo- ischiatic vacuity broadly connected across the median line life. . is elongate and directed upward and backward to the firm sacrum. or only a but always a postacetabuis al- In upright.walking animals the preacetabular process ways 120 well developed. e). The ilium is helmet-shaped in the Saurischia (Fig. however. lar one. short in the long-necked. meeting in a more or less horizontal symphysis. The There pelvis of the Chelonia (Fig. c. Sthenar osaur us or Thaumatosaurus. Usually in crawling reptiles (Figs. by the progressive debetween the ischia and pubes. 114-118 a) there small. c. as we have seen. losauria. 118 d). The pubis is larger than the ischium and has a stout tuberosity which rests upon the plastron. 119). preacetabular process to the ilium. There is is no pubic foramen." elongated in the short-necked forms. but broadly ossified in the land tortoises. a large pubo-ischiatic vacuity. and Theromorpha (Casea) it is — all due to the greater expansion The evolution of the reptilian pelvis has been. lumbar vertebrae. 127). for instance.

145. and the big toe or hallux^ on the anterior or preaxial side. or metapodials. 128). The digits of fleet. elongated and slender (Fig. conveniently called propodials.g. composed^ mammals. the or pollex.. in volant reptiles [pterosaurs] they are longer than the hind pair. with the big toe on the inner side.. as though directed backward in the the epipodials parallel. the palms of the composed of metacarpals hands and soles of the feet turned downward or to the ventral side. and a variable number of finger and toe bones. axis of the body. or very are The articular surfaces of the limb joints of aquatic reptiles (Figs.CHAPTER V THE LIMBS Two as in pairs of limbs are almost always present in reptiles. 141 i). the wrist peSj and ankle bones. the hind pair the longer. In climbing and cursorial reptiles the limbs more or less. the little finger and little toe on the posterior or postaxial side. the thumb thumb on the outer side. or epipodials. known as phalanges. As the hallux is analogous with the pollex. unextensive. they are smaller much smaller. The fore and hind Hmbs of terrestrial reptiles equal length. sometimes very much. The limbs are best understood and described as though directed outward from the long axis of the body (Fig.g. 155). mosasaurs] the front pair are often the larger. 149. and more or less carti- laginous. and should not be used for any vertebrates. In bipedal reptiles [e. much later Theropoda] or those usually assuming this posture in locomotion. crawHng reptiles are long. this nomenclature places them on the opposite sides are of approximately [e. are short. in which the digits of the two sides are brought more nearly parallel to each other. or mesopodials. the forearm and leg bones. those of the more upright-walking kinds (Figs. 155 . In aquatic reptiles ichthyosaurs. and usually the longer. of four analogous segments: the arm and thigh bones. The hind extremities are sometimes described as though parallel with the long axis of the body. 158) are poorly developed. The terms outer and inner are often applied to the anterior extremity. the manus and and metatarsals.

on the other hand. with 3 for the front. 128) reptiles were pentadactylate. The limbs of some Lacertilia and most Ophidia are wholly absent. . Primitively {e. these are more specialized. Among aquatic and volant reptiles. and most reptiles still retain these characters. in springing. 3. All other known have four functional limbs. mosahave short propodials. and only the first digit wholly In the more upright-walking kinds. As a rule the hind limbs of terrestrial reptiles. and tends to confirm Huene's contestation that the ancestors of these reptiles were originally more upright in locomotion than are their descendants.. g. some snakes have vestiges of the hind pair. and the ones remaining are more developed than those of the front feet. g. unlike other crawl- ing reptiles. reptiles. In certain lizards (Phelsuma) the first digit has become vestigial. 3. i.g. 141. hand. Figs. in which the posture in locomotion is more like that of mammals. plesio- Swimming saurs] The epipodials of ordinary terrestrial reptiles are always somewhat shorter than the propodials. and some lizards only reptiles vestiges of either pair or the front pair only. as of terrestrial mammals. 2. 4 for the the hind pair. 4. 140 a. 154). saurs. ichthyosaurs. possibly also of burrowing. 157). or volant they On the other may be consider- ably longer than the propodials (Fig.156 THE OSTEOLOGY OF THE REPTILES reptiles with propelling tails [e. the others are well developed. This weakening of the postaxial digits is especially noticeable in the dinosaurs (Figs. 5. there are fewer bones. are more specialized than the anterior ones. may be obsolete or and both the fourth and fifth and very rarely the third also. the propodials of limb-propelling water reptiles [e.. that is. Greater shortening of these bones is indicative of swimming habits. where locomotion is chiefly effected by the fore limbs. in a few dinosaurs is The first digit to be lost lost.. 5. on the other hand. 155). those in which the feet of the two sides are brought more nearly parallel in walking. 4. 156) and turtles (Fig. lost. and in strictly aquatic reptiles they are always very short. leaping. the greater strength of the foot passes digits more to the preaxial side. the phalangeal formulae 2. indeed the degree of water adaptation may be gauged by the proportional lengths of the epipodials. proganosaurs] are elongated. the fourth digit the longest and strongest. The same character is also observed in the CrocodiHa (Figs. sometimes very short. is fifth.

Captorhinus (Cotylosauria): Skeleton. One half natural size. 128. 157 . from below.Fig.

134) the articular sur- more or less spiral-like. 131. 1 29-131). or first bone of the anterior ex- tremity. stouter-limbed face is Theromorpha (Figs. In most of the Cotylosauria (Figs. directed forward obliquely. corjd. dorsal side. is brought backward in walking. permitting movement in an antero-posterior direction with a concomitant partial rotation as the hand. The bone was not depressible below out dislocation. free. femur. 129. 128. One half natural 'size. flbuU Theromorph limbs: Naosanrus. ventral side. humerus. 132) and Cesser troca.. extending around the head from the ventral postaxial to the dorsal preaxial side. usu- by a more or less complete. The articular surface of a horizontal plane withthe pterodactyl humerus .iS8 THE OSTEOLOGY OF THE REPTH^ES Propodials The humerus ally (Figs. articulates in the glenoid fossa of the scapular girdle. 130. ball-and-socket joint. mecioa/t'pvt- "rO/cLi/auc ulnar Fig. permit- ting rotation. 129.

Seymouria (Cotylosauria). D. E. A.THE LIMBS (Fig. radius. C. That on the preaxial ventral side (Fig. <? from side.ventral. b from front. usually situ- ated above the middle third. near its articular part. left femur. are two more or less prominent processes for the attachment Fig. At the upper or proximal end of the bone. 130. from behind. right femur. muscles. undetermined. from Cacops bone-bed. natural size. but often descending nearly to the middle or even below the middle in stout-limbed reptiles. femur. is the . B. 141) is 159 saddle-shaped. permitting motion in two planes only — of antero-posterior and dorso. right humerus. ventral side. left tibia. F. tibia. Humerus. 131).

is lateral tuberosity the shaft usually round or oval in cross-section. as . 131. Doubtless in these animals. the bicipital fossa in such cases looking more dorsad than ventrad. three fourths natural size. 131. C. distal is The always great in the Cotylosauria some Theromorpha (Figs. or some of them at least. radius and carpus. 131). A. or expansion also in may nearly equal 130. distal end. is Among all reptiles the lateral process It is most developed in the pterodactyls (Fig. 129. the head. left ulna. Ophiacodon mirus Marsh (Theromorpha). at other times approximating or even exceeding a right angle. 134). left humerus. one half natural size. one half natural size. on the ventral Immediately below the the bicipi- fossa (Fig. (Figs. also largely developed in the Cotylosauria (Figs. is Between the two. tal is the ulnar or medial tuberosity side. nearer radial or lateral tuberosity or process. one half natural size. B. 130). Fig. 131). 129. the peculiar humerus is to be correlated with the screw-like motion in the glenoid and Anomodontia. 133). Theromorpha (Figs. left carpus. ventral side. ventral side. 133). 130. descending below the middle of the bone. 128. in stout-limbed reptiles like the Cotylosauria (Figs. The expanded extremities of the humerus are the angle sometimes slight. tremity The width of the more expanded distal ex- may be it less than an eighth of the length of the bone. 141). left humerus. D. 129. and often not well marked. sometimes in divergent planes.i6o THE OSTEOLOGY OF THE REPTILES On the opposite side. and Anomodontia. dorsal side.

131. The humeri but Kttle from of many known temnospondylous amphibians differ those of the Cotylosauria. mode of progression. in the short-limbed cotylosaurs and theromorphs as also the is temnoas the spondyl amphibians. the radial condyle. In some of these it is replaced by a groove. there is a stout process on the radial side above the epicondyle. Dinosauria. That on the ulnar side. for articulation of the ulna. 129 A. 133 sc. 130.). Proganosauria.) and theromorphs (Fig. ectocondyle. g. and the latter is present in the Mosasauria and young Plesiosauria. as also the stout-legged temnospondyls (Fig. d) side. for the articulation of the radius. 131). A similar foramen on the radial side. ectep. there is a more or less convex surface. the Nothosauria. known radial epicondyle. The distal expansion of the humerus on the ulnar or postaxial side is commonly known as the entocondyle or entepicondyle. Theromorpha.THE LIMBS fossa. In the very stout-limbed cotylosaurs (Figs. but more distal and the ulnar condyle or trochlea. Crocodilia. 130.). the horizontal position of the l6l in humerus locomotion. occurs in all Cotylosauria. or capit-ellum. and Phytosauria. save in the absence of the .J. The on the radial or preaxial side. the ectepicondylar foramen (Fig. {e. or pre-epicondyle. ectepicondyle. Both the ectepicondylar and entepicondylar foramina occur in some Theromorpha and Anomodontia. misleading terms (Figs. for the passage of the radial nerve.f.ior the passage of the median nerve. distal extremity of the humerus (Figs..h. sometimes turned more dorsad. cially correlated It is espe- with short digits and doubtless a more turtle-like mode of progression. and the more turtle-like are never long. dorsal. the entepicondylar foramen {entep. Pleurosaurtis etc. above the radial condyle (Figs. The digits in such animals and the ungual phalanges are short and stout. 158 c. 133). on the At the preaxial and more or less ventral side. and most therapsids. 131. Araeoscelis. The Pterosauria. It may be known as the supracondylar process. 129 a) are very characteristic foramina in most reptiles. 136). Ichthyosauria. 129 a.a. 133 ewi. and * Plesiosauria have no epicondylar foramina. . Chelonia. Contiguous with is it on the postaxial Fig.p. Choristodera. Piercing the condylar expansions more or less obliquely (Fig. is characteristic of most Lacertilia. In aquatic reptiles both of these are simple projection or process facets at the extremity of the humerus. 131).s.. and in not a few mammals. Rhynchoceph.).

i62 THE OSTEOLOGY OF THE REPTILES is entepicondylar foramen. 132) and pterodactyls (Fig. accounting. but sometimes. Its articulation in the acetabulum with the is by a more or less convex head. vol. This foramen reported for Cochleosaurus. The dinosaurs (Fig. no and mammals. consequently. Mus. the convexity in the Chelonia.. have the proximal preaxial border of the femur tion more or less curved. . (Fig. as well as outward. like the variable in shape. perhaps. Broom. 132). the the turtles. then. 135). but also apparent in the quadrupedal. motion. The thigh-bone. while Amalitzky. but is known in no other amphibian. Hist. so characteristic of birds from the shaft of the bone by a more or less well-marked neck. 155) only. Amer. XLVi. Many of the Therapsida though without a diiferentiated neck.^ and like femur is them probably never was brought below a horizontal position in walking.. Figs. or directed more or less up- ward. Nat. — Ed. cannot be dis- brought parallel with each other in the same direction without location from the socket. So propodials of Lystrosaurus and doubtless of other Anomodontia were directed horizontally in loco- On 1 the preaxial ventral side. Femur. An ectepicondylar foramen is quite unknown in the class. and is known in Diplocaulus of the LepospondyU. of a neck to the femur is set off indicative of crawling or aquatic habits. for the reduction of the phalangeal formula in that genus. as in Diadectes.] . There is. have the head real neck. is {e. g. descend[ButRomer {Bulletin. The femur body the bone is is of most reptiles is turned artic- outward from the long axis of the if in locomotion. as in the short-limbed Cotylosauria. for instance. mammal-Hke mode Pariasaurus of the Cotylosauria has also been restored in a more upright posture. in such reptiles. 129 b. usually on the upper third of the bone. plate xlvi) shows that the pariasaur femur (Propappus) differs in significant features from the femora of cotylosaurs. 1922. though. more on the dorsal side. and Romer are agreed that the femur of paria- saurs was directed obliquely downward. as The two femora of a lizard. the mode of locomotion was probably more like that of also. more its it upright. with the articula- more on the preaxial side. ulation at the extremity. giving evidence of a of progression. a rhachitomous temnospondyl. most noticeable in the bipedal types of dinosaurs. because of the more or less vertical or antero-posterior position of the femora. The absence. or femur humerus. but quite like that of the earlier cotylosaurs.

5607 B Therapsid femora: A. 132. lower row. proximal end. Upper row. ventral. After Gregory and Camp. 163 . Aelurosaurus. Moschops. Cynognathus. Fig. B. Scales various. dorsal. C. middle row.

long and pendent. 129 b). ing below the middle. to the fibula. The preaxial condyle. are separated by a groove in front and another behind (Fig. the fourth trochanter. aspera of mammals and may be called the adductor ridge or On the opposite side. have near the middle on the ventral preaxial side a rugosity or prominence. The femora dicated in of the dinosaurs (Fig. is is 1 54) [and in certain Therapsida. 154). or nearly to. 132]. a rugosity more or less pronounced ridge or roughening descends toward.^ from which usually a postaxial condyle (Figs. is [Recent evidence (Romer. 132). half ""*""' ^"- the length of the bone. and nearer the head. The condyles. but usuarticular more strongly developed. sometimes markedly sigmoidally in curved (Figs. at the upper extremity in but broadly ventral in most other forms. the lesser trochanter. It corresponds to the linea crest. and in part to the is tibia behind. It is more slender Fig. sometimes. Fig.] A better name for it is . of the The femur ally the temnospondylous amphibians 151 a) is sometimes indistinguishable from that adductor ridge is of the cotylosaurs. gives articulation to the tibia. kinds. chanter. but also in- the Sauropoda. the great trochanter. right femur. a depression or fossa [intertrochanteric]." — Ed. the or eminence. at the distal extremity of the femur.164 THE OSTEOLOGY OF THE REPTILES is. the postaxial condyle. obsolete or even absent in ordinary crawling reptiles but well devel- oped in the Chelonia (Fig. always longer and its niore sleudcr • than the humerus. 157). distal After Giimore. One sixth Width scMom i i i t if ever equal to more than (Fig. as in Camptosaurus. Dinosaur femur: Camptosaurus. 129 b. Between the two there turtles (Fig. and the ends are 1 less well ossified. 1924) indicates that this process not homologous "internal tro- with the true "lesser trochanter" of mammals. there especially in crawling forms. The the shaft of the femur 155. 132 bis. 132 bis)^ especially the bipedal Predentata. usually the smaller.

flexibly in the aquatic types. ventral One fourth natural size. One half natural size. foreleg.THE LIMBS Epipodials 165 Radius and Ulna. Fig. to the by which the dorsal sur- and hand is turned forward at right angles of the radius (Fig. . withthe out rotation of the radius. chelonians (Fig. noscelis. they may be more or less fixed in Fig. 145 a). The two bones of the forearm or antihrachium are always complete in reptiles and movable upon each other. humerus without pronation or rotation 145 a). as mounted. Cotylosaur limb: left Limside. that is. wrist. freely in most terrestrial reptiles. Ophiacodon: Anterior extremity. The forearm in this order has a peculiar twist on the humerus face of the forearm. though not crossed. 133. 134.

articulating with each In some of the later plesioill other throughout their adjacent sides. normally with the intermedium and ulnare of the carpus. 134). are shortened. or elbow. They retain some of their land characters in the early plesio- saurs and ichthyosaurs. which more often the with both condyles of the femur. permitting a wide range of lateral movement in the foot. in later reptiles it is closer and firmer. on the postaxial side. by a hinge. like the posterior epipodials. This joint in the early reptiles was extensive and loose. It articulates of the always the larger in is terrestrial reptiles (Fig. hind-leg is The tibia on the preaxial or big. in reptiles. on the postaxial or little-toe side. less Its proximal extremity is expanded into a more or for the is prominent cnemial crest on the dorsal side immediate attachment of the extensor muscles. though chiefly with the preaxial. D. since there rarely sesamoid bones of no patella. distally. the olecranon is but feebly or not at all produced. 133. on the thumb. saurs a third and even a fourth bone. unlike the radius. In terres- the ulna produced more or less into an olecranon. as in the pentedactylate humerus by a more or mammals. 134). under- may articulate with the distal end of the in humerus on the postaxial side. The fibula. 158 c. or the astragalar part of the fused bone. 133. A third bone is also known some ichthyosaurs — an accessory epipodial (Fig. especially in bipedal forms. as in mammals.toe side smaller. more specialized of both groups they are wider than long. 136) present no characters by which they can be distinguished from those of the Cotylosauria. The less articulates with the preaxial condyle of the concave and rotating joint. but somewhat spiral joint. The ulna (Figs. distally. 159). Tibia and Fibula. its distal is usually also at trial reptiles postaxial angle with the pisiform. 135. In the aquatic reptiles the two bones. 151. normally with the radiale of the carpus. It articulates proximally exclusively with . whose homologies are stood. 158 c). 135). and any kind. 153) articulates exclusively with the astragalus. articulates with the trochlear condyle of the humerus.1 66 THE OSTEOLOGY OF THE REPTILES radius (Figs. The radius and ulna of the temnospondylous amphibians (Fig. is more slender than the tibia in land reptiles. but in the (Figs. The distal extremity (Figs. radial or preaxial side. sometimes losing all resemblances to the terrestrial forms.

in the later pterodactyls (Fig. . in is which the astragalus closely united or fused with the tibia. B. Varanops. It 167 has more of a sliding or It arthrodial joint in land reptiles rotating the foot in extension. articulates distally — primitively with the calcaneum and astrag- FiG. Its lower end in those is reptiles. 135. alus. 155 c) the whole bone has disappeared. Theromorph limbs: A. Casea. One half natural size.THE LIMBS the postaxial condyle (Figs. especially cursorial reptiles. 135. 151). as a separate ossification at least. obso- lete or lost.

Fig. A.Temnospondyl limb: Eryo/)^. B. reconstruction. Miner. and Noble. in . dorsum. One third natural size. under After Gregory. side. C. or upper side. American Museum of Natural History. Cope's original specimen. 136. left foreleg.

articulates with three bones. side. corresponding quite to the scaphoid. except that they are somewhat. pus and tarsus of The carpus (Fig. and Trematops. intermedium. and unciform human wrist. The elongation of the tibia and fibula. Watson has recognized a small third centrale in the curious genus (Fig. 158 a. in the 151 b) from the middle Pennsylvanian. utnare. or much. pyramidal. the radiate. and ulnare. the carpaha. as also in the Proganosauria (Fig. known reptiles is composed none small: four in the first row. 146-148. 155 c). two in the second row. Eosauravus (Fig. (Fig. 137 e) from South Africa. from the preaxial to the postaxial intermedium. 136) same in in but two temnospondylous amphibians. the first four corresponding to the magnum. Mesopodials Numerous modifications have occurred in the structure of the carreptiles in adaptation to diverse habits of life. As in the front leg there may be accessory epipodials in both the plesiosaurs and ichthyosaurs. In both. the radial or trapezium. the supis posed radiale. trapezoid. first. and Thalattosuchia 150). so characteristic of aquatic animals. is known The radius of Eryops. and pisiform of the human wrist. however. the preserved bones are number as in the early reptiles and some modern ones. 153 a) and Choristodera. Aigialosauria. reached the maximum The in the Pterosauria (Fig. of the centrale. and five in the third row.THE LIMBS The 169 posterior epipodials in aquatic reptiles (Figs. and pisiform. unknown as an ment in other reptiles. 158) are almost indistinguishable from the anterior ones. is seen to a moderate extent in the earhest known is reptile. so characteristic of the cursorial or leaping forms. though perhaps represented by a the young of the modern Sphenodon. It much more pronounced Mosasauria (Figs. lunar. b). 159. while the pisiform large. tibia and fibula of some temnospondylous amphibians are quite indistinguishable from those of many Cotylosauria. smaller. and the ulnar or second. This short- ening of the epipodials. Broomia ossified ele- cartilage in Carpus The carpus Eryops the (Fig. Thalattosauria. and an articular surface on the postaxial distal margin of the ulna . called respectively. 134) or wrist of the earhest of eleven freely articulated bones.

Primitively both centralia were large (Figs. 134). it seems to the author. but not the fourth carpale. 133. Moreover. Two and two only. but not always.. 133). Unfortunately. as correlated with the longest and strongest digit. 141 i. Every other bone may be absent in different reptiles. even this early. while the specimen of Eryops. seems certain. had. brought about a modified structure in the carpus and tarsus of Limnoscelis (Fig. is at least since early Eocene times.170 THE OSTEOLOGY OF THE REPTILES seems to indicate the original presence of another bone that would correspond better in position with the real pisiform. centrale. 141 f) and Stegosauria (Fig. That there has been an actual loss of the first to centrale and fifth distale. the single known specimen of the carpus of Trematops has but two centralia. it is most first part. — Ed. Nat. here figured of Dr. their actual loss in hving reptiles has long since been affirmed. where unoccupied spaces for cartilaginous elements have been preserved. more after the mode of the tortoises. retarding or pre- The first centrale and fifth carpale are always absent in mammals. and short epipodials. 1923. short. and Lystrosaurus been doubtless true for the It is the general belief that the loss of mesopodial bones has due to their fusion with adjacent ones. Gregory. intermedium. and ulnare that was very persistent. There are five well-developed carpalia in both forms. manus of Eryops see Gregory. Noble Amer. Miner. supposed water all habits have delayed the ossification of some of the mesopodial bones — supposedly. There was a perforating foramen between the second venting the fusion of these three bones (Fig. 137 e). but it is a curious fact that have similar feet. proving conclusively the presence hand. that similar walking or digging habits. Hist. Diadectes. Mus. broad toes and ungual phalanges. unless it be in certain quadrupedal dinosaurs like the Sauropoda (Fig. It is not impossible. the be lost in both carpus and tarsus. very broad humeri. vol. as it also is the first to of the carpus be ossified in the human wrist.^ of five digits in the In some of the early cotylosaurs and anomodonts. but the second centrale 1 often and [For a different interpretation of the in Bulletin. as shown in specimens of various early reptiles. has a small space which may represent a small third centrale. by the kindness were present in Trematops. bones preserved in the proximal row. j).] . And this carpale is the most persistent bone in the carpus. 134. xlviii. as was also the fourth carpale.

Natural size. After Watson. left tarsus. Galesphyrus. About twice natural size. Natural size. 137. D. front and hind feet. left tarsus. E. left carpus. Ga/echirus. Therapsid limbs: A. Broomia. B.Fig. Natural size. 171 . Broomia. C. After Broom. After Watson. After Broom.

the car- (Fig. or a radial sesamoid. Among . Scymnognathus (Therocephalia). G. an error. hind leg of same. One D. as figured by the same writer (Fig. THE OSTEOLOGY OF THE REPTILES In Procolophon only. as morpha. E. F. After Broom. 138 b) the Dromasauria has a small intermedium. Nine eighths natural size. Throughout the Theronumerous forms. One Among pus is ill the reptiles collectively known as the Therapsida. front foot. dorsal side (rearranged from half natural size. Seeley). part of tarsus. One third natural size. as it also is in Dicynodon and its allies of the Anomodontia. Araeoscelis (Protorosauria). A small element found near the pale was referred to a possible prepollex. 137 a) as known. 138. In Galechirus of figured by Broom. in nous in one known genus. Enlarged. The carpus of Scymnognathus of the Theriodontia. Theriodesmus (Therocephalia). of the Cotylosauria. C. Sceloporus (Lacertilia). as known plete. Protorosaurus (Protorosauria). front leg. ^^lJb Limbs: A. B. is is this centrale absent. After von Meyer. front leg. Varanops. the carpus is primitively comsave that the first centrale and fifth carpale remained cartilagialso affirmed of the radiale.172 present. and the fifth carpale is represented as fused first car- with the fourth. the primitive structure is retained. F probably immature. half natural size. Fig.

Sphenodon. Rhynchocephalian limbs: A. After Lortet. traces of the missing bones have been shown in dotted lines. also lacks the fifth carpale. The carpus 140 a). of the carpal bones. — Ed. Pleurosaurus.] . Acta Zobl. B. In Rhynchosaurus.^ 138 a) of the same group. indicating a primitive carpus save for the pisiform which less present. in a specimen figured by Newton.THE LIMBS reptiles there is 173 Theriodesmus (Fig. as restored by the author from Seeley's figures. D.. 307-360. The complete carpus is unknown members the only of the Therapsida. 1922. C. Extinct members of the order and its allies of the Diaptosauria are not sufficiently well known to determine it whether been this primitive structure is general. After Howes and Swinnerton. was doubt(Fig. Sauranodon. Nine eighths natural size. size. 139 b) of the RhynchocephaUa is modern which has retained the primitive structure and arrangement Fig. 139. It is of the Crocodilia has of four been strangely modified all composed bones only in forms so far as known 1 [But see Steiner. Sphenodon reptile (Fig. no evidence of a lost prepoUex. though the intermedium in other is not small. pp. Nine eighths After Lortet. though doubtless has for the most part. About seven eighths natural natural size.

Limbs: A. dilia). articulates with the postaxial border of the ulna and ulnare. distally with the fourth carpale only. . pisiform. but smaller. lating also with the pisiform and radiale. are represented by cartilage. and fills perhaps also the centrale. Alligator (Crocodilia). articu- FiG. 140 b) and Crocodeleimus from the two carpals are yet more elongate. as they are usually called. The first three carpalia. Natural size.174 THE OSTEOLOGY OF THE REPTILES The very large and elongate. B. is a very ancient one as shown in the carpus of AlliJurassic. where. 1 40. It is supposed to be the fused radiale and intermedium but possibly is the intermedium only. Alligatorellus (CrocoTwice natural size. radiale is its and preaxial border of the ulna. D. The pisiform. dilated at ends and the radiale. The ulnare. of considerable size. the out the interval between the end of the radiale and the metacarpals. One half natural size. gatorellus (Fig. Amblyrhynchus (Lacertilia). ulnare. and fourth carpale. is aparticulating with the radius proximated to the middle part of the distal border of the ulna. of similar shape. which This structure indeed. C.

After Gilmore. About one sixth natural size. After Riggs. AlloAfter Gilmore. . J. C. After Gilmore. One fourth natural size. one eighth. One fifth natural size.After von Huene.] H. eighth natural size. Thescelosaurus (Ornithischia). After Gilmore. One half natural siye. L. M. Trachodon (Ornithischia). saurus (Saurischia). One twelfth natural size. One ninth natural size. Leptoceratops (Ornithischia). and one fourth natural size. Plateosaurus (Saurischia). F. Stegosaurus (Ornithischia). I4I. One eighth. One Fig. After Lambe. D. I. One eighth natural size. Gryponyx (Saurischia). Morosaurus (Saurischia). G. K. Dinosaur pedes: A. After Marsh. After Brown. Allosaurus (Saurischia). After Brown. After Broom. E. One fourth natural size. Morosaurus (Saurischia). [Should be Brontosaurus. B. Gorgosaurtts (SauOne twelfth natural size. rischia).

j) the postaxial one of the two has been found in young specimens in three parts. (Fig. 141) has suffered greater reduc- The carpus tion than in any other order of terrestrial reptiles. 141 i. After Plieninger. 141 m) of the Ceratopsia. is. though identified as the first and second in Ornitholestes and its immediate alHes of the Theropoda. In Stegosaurus (Fig. the intermedium. in Leptoceratops (Fig. as would be expected from the constantly reduced fifth finger. In no form has a centrale been reported. In the quadrupedal forms there are but two proximal bones. medium radiale. except the Trachodontidae in most cases the and fourth carpale. Fig. that bipedal forms. Rhamphorhynchus. Pterodactylus.176 THE OSTEOLOGY OF THE REPTILES in the Dinosauria (Fig. In the Theropoda. and ulnare seem distinct in though not The second row of carpals has disappeared in the (Fig. and pisiform perhaps that was also the case in the Sauropoda A small bone may possibly represent a vestigial inter. American Museum of Natural History. 141 r). ulnare. 141 i). Nearly three times natural size. One half natural size. The carpus other third in the Trachodontidae (Fig. doubtless because of the upright posture. the all. large. Sauropoda (Fig. both large and massive. and iguanodont orthopods. 141 h) is more reduced than in any other . 1 42. 141 f) and Stegosauria Two have been found in all . intermedium. Pterosaur limbs: A. and the fifth carpale is doubtfully present in any (Camptosaurus). B. known forms.

dorsal. The most remarkable modifications of the carpus are those of the volant Pterosauria (Fig. D. dactylus (Fig. right hind foot. on the radial side. dorsal. articulating chiefly with the [distal] carpale. articu- lating with both radius and ulna. some forms there are and the usual lateral one in. the second with the fourth or wing metacarpal. supporting the pteroid. unless it 177 re- be some aquatic mosasaurs. etc. the carpus its allies of the Upper Crehomologized reduced to three bones: a proximal one. a third carpale. This great consolidation of the carpus . all the bones of the proximal row except the pisiform. but two small bones maining. of Chameleon. probably the former. 142 a). with one. fibula. the first articulating with the first three metacarpals. The earliest stages we do not know. Rhiptoglossa. except that in two in the proximal. taceous. 143. the centrale. much enlarged. A. though certain progressive modifications are observable from the In Pteranodon and is earher to the later. In the earher Rhamphorhynckus (Fig. 142 b).. with tibia. 142). a distal composed either of the greatly enlarged fourth carpale. and perhaps to be Fig. or possibly neither. left innominate. right hand. there are two distal carpals. Limbs. and tarsus. may be either the first carpale. or a fusion of two or three. Squamata. C. two in the distal row. B. This is also the structure in Pterofive bones.THE LIMBS reptiles. probably the radiale and fourth carpale. right scapulocoracoid.

right front and hind legs. 144. Pleurodira: Thalassochelys. Chelonia. which ulnare. The first examined. and pisiform. intermedium is visible in the various forms though sometimes small. Marked modification in the structure of the carpus is also charin acteristic of the Lacertilia (Fig. 140 c).lyS THE OSTEOLOGY OF THE REPTILES maximum of firmness with but little mobilis pterodactyls resulted in a ity. is No may also be interpreted as the radiale. It reported to be present only in the family Lacertidae. Fig. The carpus of the Pseudosuchia and Phytosauria practically unknown. . cen- A centrale is usually present. which was not needed in the volant hand. There are but three bones the proximal row.

Indeed. and posteriorly placed pisiform in the first row. third. In the marine Chelonia (Fig. 143 a) the carpus is reduced to four functional bones. are usually large. and the carpale radiale was doubtless cartilaginous. among the terrestrial tortoises (Fig. a large amount of cartilage remaining in the joints. first centrale. and The radiale and intermeis least reduced. not only of the mesopodial bones. The as usual. except that the radiale. unless be the element sometimes called the centrale. reached almost to fifth chelys. which may represent the first and second carpalia. fused with the fourth. The pisiform is greatly enlarged and has lost its primitive location between the ulna and ulnare. At the opposite extreme. dium are more or less elongate. some lizards. The may be ab- sent. The earliest ob- served effect of water habits delayed ossification. probably the latter. though much modified. all and fifth carpalia and the pisiform may be coossified. ulnare. the ulnare is small. 144) the carpus is broad and flat. or the second and the centrale. the proximal bones were less elongate. around which the metacarpals revolve. or separate and distinct. but of the bones of the skeleton in general. the centrale large. In the curious hands of the highly speciahzed perching Rhiptoglossa (Fig. fourth. This was the structure of the marine turtles as far back as the Cretaceous in Protostega. the second. who found Emydura the two centralia in their normal positions. though enlarged. this is the explanation given by Baur. The fused centralia in such early forms as Idiofrom the Jurassic. first is is large. is left of it but a which has usurped 145 a) the radiale has disappeared until nothing nodule of cartilage united with the its place. The changes of the wrist and hand is in adaptation to aquatic life are more profound than those of terrestrial reptiles. and a large. it and fifth The first absent. hemispherical. in some old animals the third.THE LIMBS trale is also identified in 179 fourth centrale [carpale]. Between the first metacarpal and radiale there are in the more specialized types two minute bones. fused third and fourth carpale in the distal row. becoming attached to the ulnare and fifth metacarpal or the latter alone. The two centralia are often present. in At least. the radius. often fused into the large single bone. Partial chondrification of the wrist and ankle occurred as early as the cotylosaurian Limnos- .

the same. 148) the most highly specialized of all mosasaurs. . since space them in many specimens as they have been found in the rocks. C. vanced aquatic type. 146) of the Mosasauria. first and fifth carpalia were not. In Tylosaurus (Fig. Chelonia. etc. intermedium. the ulnare. front leg. B.) dorsal side. third and fourth carpalia. A. hind foot (tarsus. radial side. essentially a surface-swimming the four proximal bones of the wrist are ossified. there are but one or two bony nodules is left left. In Platecarpus (Fig. THE OSTEOLOGY OF THE REPTILES 133) and Diadectes. Pleurodira: Tw/Wo. but the centralia. dorsal side. 147) a more ad- FiG. leaving only the radiale.i8o cells (Fig. one of which is certainly the fourth carpale. In Clidastes lizard. (Fig. I45. and second carpale have also disappeared. pisiform. probably marsh animals. All the others disappeared as for bones but remained as cartilage.

Platecarptis (Mosasaur). <^^? - Fig..::-C:::. ulna. Fig.. (^=^ £>-2 %. I47. radius. coracoid.^^ . humerus.. h.^' C. right front paddle. 181 . r.. left front paddle. sc. scapula. I46.-. Tylosaurus (Mosasaur). left front paddle: c. I48. Clidastes (Mosasaur). «.s^JC:r. Fig.

. the radiale apparently the most con- its absence. Nothosaurian limbs: Lariosaurus. Fig.l82 THE OSTEOLOGY OF THE REPTH^ES (Fig. 149) o^ the In the swimming feet of Lariosaurus the carpus spicuous for is Nothosauria also reduced. 149. About four times natural size.

visible. ber. and paddle-like left front After Fraas. presumably a cotylosaur though the It has but skull is reptile. eight. The carpus and hand strictly aquatic or the marine reptiles are so like the ankle and foot that they may be discussed to- gether (p. from the Permian of South Africa. two bones in the prox- imal row. one corresponding to each metatarsal. Geosaurus (Thalattosuchia). 183 and hand in the marine Crocodilia of them.THE LIMBS Very interesting are the modifications of the wrist (Fig. not known. from the middle Pennsylvanian. of which are undoubted tarsalia. five and only six. 150. and probably also two the cotylosaurian genus Labidosaurus. 152). these. of very broad and flat. the first supposed radiale. 193). left two centralia in the middle row on the tibial side. corresponding quite to the astragalus and calcaneum of mammals and Beyond bones are the typical reptiles. six. The whole numwas the most known any reptile until recently. 150). Elon~ hind leg. Tarsus The earliest known tarsus is that of Eosauravus (Fig. the astragalus and calfrom the Fig. Nine bones are present in the in tarsus of Ophiacodon (Fig. have in also Since this discovery two been found by Watson in the genus Broomia 137 d). one may be a centrale. The second . uppermost Pennsylvanian or basal Permian of New Mexico: two in the proximal row. gate leg. and caneum. centralia (Fig. correspond- ing to the elongated ossified bones of the terrestrial forms. the But two is carpals remain. 151 b). five tarsaha in the distal row.

a. A little . but the fifth tarsale all is absent in is reptiles since Triassic times. 152. 151. About twice natural size. than one third natural size. calcaneum. Fig. less One half natural size. B. Fig. though present in most mammals. (Cotylosauria). Limbs: A. Trematops (Temnospondyli).i84 centrale is THE OSTEOLOGY OF THE REPTH^ES usually retained in later reptiles. astragalus. from mounted skeleton. Eosauravus Ophiacodon (Theroniorpha): right hind leg. and a free centrale absent in all living reptiles. c.

and the ichthyosaurs and plesiosaurs. the navicular corres- ponding to the second centrale. sponding to the greater length and strength of the fourth The tarsus is known in but two temnospondylous amphibians. a from the specialized to the generalized condition of the Amphibia seeming impossibility in evolution. then. altogether improbable that both intermedium and tibiale have ever been present as separate bones in reptiles since early Pennsylvanian times. 149).^ Since Gegenbaur. the largest of the series. and these reptiles are generally additional bones of the amphibian tarsus. there are three and In the former. and according to Baur in the latter bones in the proximal row. Zool. corretoe. This is denied by Baur. But. and crocodiles. Howesia of the Rhynchocephalia. it is generally believed that the intermedium is fused with the tibiale to form the astragalus. four centralia in the tarsalia in the distal — twelve in middle row. who says there is no evidence of such union. or unless such forms have enjoyed an uninterrupted and independent it is descent of which we have no knowledge from the Amphibia. in this respect. A separate intermedium has been accredited to certain reptiles. There have been various theories as to what has become of the (Fig. the inter- and the third and fourth centralia. primitively. Trematops Archegosaurus.THE LIMBS The mammalian foot. 151 a). also. is 185 even more primitive than that of the Hzards. 1 [For an excellent review London. medium. mamSoc. Otherwise we must assume that there has been a reversion in these animals. 192 1. . (Fig.] of this subject see in Proc. 143-155. tibiale. Nine bones. the tibiale represented by the tibial sesa- moid. inter- medium. Moreover. and five all. we may assume was the primitive number of tarsal bones in the reptiles. The fourth distale. which occurs in certain lower animals. the tibiale. there are but two bones in the proximal row of the" tarsus of the Nothosauria to supposed have a real genetic relationship with the plesiosaurs. both from later rocks than Eosauravus. and Three fibulare. the cuneiforms and cuboid to the four tarsalia. mammals but it is is unknown as such in In this uncertainty better to use the two Broom. Others have thought that the intermedium alone forms the astragalus. Oudenodon {Dicynodon) of the Anomodontia. of these have been lost in all known reptiles. as in the carpus and is as a general rule in all reptiles. turtles. pp.— Ed.

The second known to fuse with the astragalus in the . Limbs: A. 153. Three fourths natural size. Sauranodon (Protorosauria). fibulare. Of the Fig.1 86 THE OSTEOLOGY OF THE REPTILES malian names astragalus and calcaneum and abandon the names tibiale. the third with the fourth tarsale. Natural size. centralia the most probable theory is is that the fourth of the amphib- ian tarsus has united with the astragalus. Modified from Lortet. and intermedium for the reptilian tarsus. B. Modified from McGregor. Mesosaurus (Proganosauria).

is first and second b). the mammals). Four tarsalia are usually present. And very probable that the first centrale of the amphibian and rep- tilian tarsus ceased very soon to be ossified. 145 154 d. 54 c) is In the Chelonia the small calcaneum sometimes free (Fig. 155 d) i). Zool. and most c. Sclerosaurus. 156 reptiles. is has been shown by Baur and others that the fifth tarsale not fused with the fourth. known reptiles. tarsalia are either vestigial The tarsus of the chameleons (Rhiptoglossa). and Telerpeton of the later Cotylosauria. have not more than seven tarsal bones. 187 (Fig. There is no centrale or fifth tarsale. and the cal- caneum is either fused or some Dinosauria (Fig. In the kionocrane Lacertilia (Fig. It even in a fused condition. true tibiale has disappeared in later reptiles. but has disappeared. until their discovery four tile. The fourth tarsale always large. the fifth tarsale being invariably absent. Baur based the order Proganosauria chiefly upon this All other character. 143 ^ But two bones remain in the mal rov/ of the tarsus of the very primitive Seynwuria. 154 c) perhaps at other times it is lost. like the wrist. 139 a) of the Rhynchocephalia. Lacertilia (Fig. the astragalus and calcaneum are fused into a single bone. [Watson (Proc. 1919) reports the presence of three bones in the proxiand adopts the view that the only. 153 a). (Fig. The been recognized in the Proganosauria tarsalia. Sphenodon (Fig. but there are five were the most known in any repexcept certain Therapsida (like Indeed. 152).THE LIMBS modern Chelonia it is . 140 d) and Chelonia (Figs. Soc. in any later reptilian tarsus. very curiously modified (Fig. In Pariasaurus. the small calcaneum either indistinguishably fused with the astragalus. the Cotylosauria there are usually eight tarsal bones. the astragalus representing the inter- medium — Ed. ^ fifth tarsale In Pariasaurus the centrale and are not In the Theromorpha eight are present in known with cerall known forms centrale has not except Ophiacodon (Fig. The is centrale is never free.] . and the or lost. and A free centrale is absent in all modern though sometimes suturally fused with the astragalus in 1 the Chelonia (Fig. or suturally attached in the adult. lost in the Pterosauria (Fig. g). and is not represented. the third sometimes suturally united with the fourth. which has nine. 143 b) there is a similar condition. 154 c). Among tainty.

hind leg from postC. B. D. F. around which all the short metatarsals closely articulate in two groups of two and three. front foot. Cistudo. dorsal side. Limbs and feet of Chelonia. postaxial side. left hind foot. G. Chrysemys. Podocnemys. G Fig. articulating together enarthrodially.1 88 THE OSTEOLOGY OF THE REPTILES highly speciahzed species. Trtonyx. the astragalo-calcaneum and the large. hind foot. left hind leg. axial side. Chrysemys. dorsal side. Natural size. dorsal side. A. 154. hind foot. Podocnemys. dorsal side. right front forearm and foot. Chelydra. hemispherical fourth tarsale. dorsal side. . E.

second.THE LIMBS The hind foot is 189 In Galechirus poorly known in the Therapsida. like the carpus. c. as in the young of birds a parallel character which has no genetic value. 156). the Dromasauria the and a sent. 137 c). 155 d). the calcaneum fused with the astragalus.. apfourth is always single when present. but a small (Fig. There is a tendency in all for the two proximal bones. is highly specialized. e) fits more or less closely in a depression or groove on the under and anterior side of the tibia. in the later forms (e. 137 b) of one has been recognized in the related genus Galesphyrus (Fig. — always absent. pulley-like articulation. The astragalus of the Theropoda (Fig. the astragalus and calcaneum. like the carpus. like Pteranodon or Nyctosaurus. In the early pterodactyls there were at least three other tarsals. the fifth tarsale is ab- The is tarsus is The carpus. in the later ones. tlie The tarsus of Pterosauria (Fig. Ornithomimus. The Anomodontia have the astragalus and calcaneum. 156 g) even the fourth is said to be wanting possibly a vestige yet remains. parently. there are but two free tarsalia. 156 e) developing a high ascending process in front. 139 and fifth tarsale a). In the Sauropoda (Fig. probably the fourth and the fused second and third. small. The tarsalia. or fused tralia are first. tarsus of unknown in other groups. 156 b. g. to articulate closely with the leg bones. four tarsalia. frequently unossified centrale. In addition to the fused calcaneum and the astragalus. like the carpalia. . 156 i) there is a less close union. unlike the highly specialized. In the early forms the astragalus suturally united with the tibia. the calcaneum fused or forming a large. really absent it is the only known example among reptiles of the absence of all the bones of the distal row. Cen- unknown in all. end of the tibia. is In the later forms the astragalus indistinguishably united with the lost as in birds. and third. If — . Fig. the The centrale and first and fifth tarsalia are The second and third tarsalia are often fused. are absent in the Trachodontidae (Fig. perhaps due to the larger amount of cartilage in the joints of these animals. fifth tarsale is lost. the modern Sphenodon (Fig. the centrale first have disappeared and is three tarsalia are fused in the adult. The tarsus of the dinosaurs (Fig. has been much modified in adaptation to upright-walking habits.

Ha/lopus (DinoAfter Marsh. /iraeosceiis iProtorosauna). IQO . C. 155. sauria). Three fourths natural size. About five sixths natural size. Pteranodon. B. Fig. Pteranodon (Pterosauria). About one thirdl natural size. D.Limbs: A. One half natural size.

After Marsh. A little more than one sixth natural size. One sixth natural size. Plateosaurus (Saurischia). About one eighth natural size. Struthiomimus (Saurischia). Anchisaurus (Saurischia). After Hatcher. C. B. One twelfth natural size. E. One seventeenth natural size. One eighth natural size. Allosaurus (Saurischia). Trachodon (Ornithischia). After Brown. One sixteenth natural size. D. H. Dinosaur pedes: A. One I. Thescelosaurus (Ornithischia). After Brown. F. After Marsh. Ornithomimus (Saurischia). G. 156. After Huene.] 191 . Morosaurus (Saurischia). After Osborn. After Gilmore. Monoclonius (Ornithischia). About one nineteenth natural size.Fig. After Osborn. fifth natural size. [Brontosaurus.

Hallopus (Fig. B. the heel-like process. the first and fifth tarsalia produced into a and the centrale are abre- second tarsale is small. as is the case in Scleromochlus. Araeoscelis (Fig. 157. f erred Alligatorellus. Crocodilian limbs: A. 157 a. Broomia (Fig. 137 d). dorsal and postaxial. 155 b). left After Lortet. left femur dorsal. and other genera of the Pseudosuchia. About three fourths natural size. usually FiG. .192 THE OSTEOLOGY OF THE REPTILES of the Crocodilia (Fig. C. to the Dinosauria. 155 a). D. and other leaping or springing reptiles. hind limb of Alligator. also has a heel-like calcaneum. b) is The calcaneum sent.

or at most two. left front paddle. with the divaricated in land lizards. c). Ophthalmosaurus (Ichthyosauria). right hind leg. has sometimes been called the pisiform in both front and hind limbs. and metatarsal. Clidastes (Mosasauria). though represented by cartilage in the adult. In Platecarpus and Clidastes (Fig. Ichthyosaurus platydactylus (Ichthyosauria). reptiles: A. fifth In Tylosaurus. and fibulare. one sixth natural Not more than six bones of the plesiosaurs can be called tarsals. One eighth natural size. left front paddle. Platecarpus (Mosasauria). Fig. One sixth natural size. like the carpus (Figs. but as there never was in any terrestrial reptiles a pisiform in the tarsus. but one. calcaneum. About B. 158 b) the astragalus. (Fig. 158 a) 193 In the web-footed Mosasauria the tarsus. C. One third natural size. Other tarsal bones remained unossified. Limbs of aquatic size. The three in the first row are usually called the tibiale. . small bones remain. on the postaxial side. that name is of course incorrect. as fourth tarsale alone remain. They have the same shapes and relations as the carpal bones and cannot be distinguished from them except by their smaller size. intermedium. 158. D. right hind leg and tarsus.THE LIMBS 146-148). a fourth. and their homologies are doubtful (Fig. 159 b. the most speciahzed of mosasaurs. progressively became more cartilaginous.

159. may be first. /. ulna. /. fibula. right hind paddle of Thaumatosaurus. There of is the same objection to the recognition an intermedium tarsi in this order as in the plesiosaurs. Paddles of Plesiosaurs: A. r.ktT Fraas. right front paddle of same individual.194 THE OSTEOLOGY OF THE REPTH^ES for There are valid reasons doubting the reappearance of the inter- medium It after its loss in the terrestrial ancestors of the plesiosaurs. . «. humerus. B. fb. and the fourth tarsalia. The homolexists. a. femur. h. whatever may be the corresponding bone in the carpus. tibia. the enlarged centrale. where a like similarity between the front and hind limbs Fig. The bones in the distal row may be the fused second and third. even more doubtful. d). C. radius. 158 c. right hind paddle of Trinacromerum. are ogies of the mesopodial bones of the Ichthyosauria (Fig.

140 a. reptihan hands and feet of of the hands or feet and it is purely an assumption that the were evolved from forms like the later ones Permocarboniferous times. Pseudosuchia. see Gregory. must be remem- bered. Therocephalia. the modern Sphenodon having the bones arranged in the same ways. We can hardly number of digits or number of digits. and but three in the fifth. that we know nothing whatever of the earliest amphibians. in the but not or markedly divaricated. three in the There are two phalanges digit. the rhachitomous temnospondyls. 3. Mus. 133). had known to be the case in some of the ancient either It 2. Of the Temnospondylous amphibians no complete hand is known. That there were five functional digits is certain. Nothosauria. in The phalangeal formula was 3 or 2. xlviii. Rhynchocephalia. Amer. however. vol. 1923. five in the fourth. Nat. 139 d). b) as well as of has not (Fig. — Ed. 3. 1 the early Araeoscelis (Fig. 136) often assumed that all and Trematops. 4. and Noble in Bulletin. feet like those of the ancestors of the reptiles In all probabiUty the embolomerous had the phalangeal formulae of both front and hind conceive of an increase either of the known earhest reptiles. with fewer [For a different interpretation. and the group called by the author the Acrosauria. The first and fifth metacarpals are more movable on the wrist than are the other three.^ since there are five freely functional carpalia in both Eryops (Fig.THE LIMBS Metapodials and Phalanges 195 The most sauria. or at least some members of the group. as is amphibians of the past. 4. Stegocephalia. 157 a. it is seen. 128. Hist. is The five metais carpals increase in length to the fourth. 155 a). that Sauranodon phalanges.] . primitive hand or manus known is that of the Cotylo- from the Permocarboniferous (Figs. however. the fifth shorter. In crawling reptiles the structure of the 139 A. is. 3. as of the present. b). changed much to the present time. four in thumb second the third. Thelizards (Fig. the postaxial fingers are in In the Crocodilia (Figs. Pleurosaurus (Fig. 3. phalanges in the earliest reptiles. d) most modern same number romorpha. 139 c).. 4. 1380). It is but four fingers. 140 c. all cases shorter and weaker. pollex. and (Fig. Phytosauria. Miner. This primitive phalangeal formula is that of the Cotylosauria. Protorosaurus (Fig.

142) in the later ones they were mere splints lodged loosely in the flesh at the distal end of the fourth meta. This wing finger has generally been supposed to be the fifth. leaving the normal numdigit. A feet general reduction of the postaxial digits of both front and hind is characteristic of the Dinosauria (Figs. so great indeed that there is dispute as to the homologies of the ones that remained. It known is seems more probable that the wing finger the fourth. with four phalanges for the sup- port of the patagium. as in the third digit of the foot. the hand func- . 156). hand recognized. as in the bats. We can conceive of no cause for such hypo. the fourth usually. but an additional phalange has been added. especially Nycto- saurus and Pteranodon. two phalanges of the fifth The fifth finger is absent in all Theropoda since the In Gorgois early Jurassic. joint. One of the phalanges of the third finger is short. and even in these. fourth finger is On the postaxial side the very long and strong.196 THE OSTEOLOGY OF THE REPTH^ES In no other reptiles has there been as great modification of the fingers as in the Pterosauria (Fig. The fourth metacarpal. is Only in the primitive Anchisaurus and Plateosaurus (Fig. 141 a) plete a nearly com- finger are gone. carpal. where there are three very short and weak fingers on the preaxial side. 142). the terminal ones in the shape of strong claws. 141. Its distal articulation is a very perfect pulley-like permitting flexion of the first phalange through almost one hundred and eighty degrees. and four phalanges. only the first of them retaining a very slender connection with the wrist.and hyper-phalangy in the hand in these volant reptiles. on the other hand. saurus (Fig. the first finger or pollex represented by a slender bone as turned backward from the wrist toward the humerus and the pteroid. the fifth being absent. 141 c). The first three metacarpals of the early pterodactyls articulated normally with the carpus (Fig. three. while each of the preceding three digits has lost one phalange.from the uppermost Cretaceous. In the development of the patagium the claw of the wing finger would in all ber for the fourth probabihty disappear. The maxi- mum of changes was reached in the latest forms. not only has the claw been converted into a long membrane-supporting phalange. with two. progressively till it increased in length much exceeded the length of the forearm. If it is really the fifth. as originally so called by Cuvier. the third sometimes.

156). 141 a-d). (g). so far as the reduction of phais concerned. (d). 5. 4. 2. indeed. and in Trachodon Among the . 157 a. 141 h-m). 4. the formula. i. curved and sharply pointed in the Theropoda (Fig. a related genus. o being the usual one. is The foot or pes of reptiles similar in structure to the hand. 4. 54 c.3. lizards Chelonia (Fig. usually one. d) . 141 is less l.THE LIMBS tionally didactyl. sometimes two. 3. 5. It is often reduced among the 1 tarsal left. being o. i Among the Predentata (f-h) the phalanges of the fifth toe are invariably absent in known forms. according to Lambe. A very few also have lost the fifth toe. the claws lacking in the two postaxial digits. claw the largest. though seldom entirely absent. h. of the normal phalanges are in the The greater strength of the foot in this order as dinosaurs is more to the preaxial side. though primitive in Propappus. In Pariasaurus. 3. The langes foot of dinosaurs (Fig. In all herbivorous forms the outer fingers are reduced.ngers being the larger. f. for the most part hoof- Predentata (Fig. more obtuse in the Sauropoda (Fig. 3. 3.4. except in the fifth toe. The known Sauropoda (i) have 2. the extreme of specialization 197 among reptiles. In Trachodon a greater reduction has occurred. SindStru. 141 a-e) the the hand thumb is the stoutest digit. the fifth metatarsal a vestige. 3. 2. sauria. In the Trachodontidae (Fig. o. 2. the phalangeal formula never exceeding 2. o. the first finger is absent. unhke most other lost. 5. 3. 1 Allosaurus (c). thiomimus phalanges. the Theropoda a-e) retaining the original formula. the formula. from the lower Cretaceous. 3. b. almost the maximum the fifth is among reptiles. the phalangeal formula slightly reduced. the reduction of the toes being usually anticipatory of the fingers in the terrestrial forms. There was one more phalange in the fifth toe than in the fifth finger primitively. 141 like in the f. 4. 156 is less specialized than the hand. m) preaxial. Plateosaurus (a) and Anchisaurus (b). 4. 3. 2. The loss of the fifth toe is rare among reptiles. In the herbivorous dinosaurs (Fig. o. 5. reptiles. In its the Theropoda (Fig. only. g). have the formula 2. (Fig. of the Cotylois sauria. 3. d) have only the fifth metaand the fourth toe has but four phalanges. 2. 141 h). from the uppermost Cretaceous. aside from the DinoThe crocodiles (Fig. from the Trias. The ungual phalanges of both front and hind feet are characteristic. the first and second fi.

b). 145 c. 3 in both front and hind 145 a) in feet. sauria (Fig. Similar reduced phalanges are seen in the Struthiomimus and the tree sloths hand of the theropod among mammals. this arguing perhaps a more sauropod-like mode of progression. Hypophalangy. with a further reduction to 2. singularly identical ing phalanges of the . 4. The greatly elongated feet were adapted more for perchearliest pterodactyls The ing or clinging than for locomotion. 2. Cynodontia. In the Chelonia (Figs. 2. a character which has been adduced in proof of their 139 A.198 THE OSTEOLOGY OF THE REPTILES (i) quadrupedal Sauropoda less so in the axis of the foot is more to the preaxial side. 2 (i) (Fig. as in the mam2. many tortoises. either ossified or cartilaginous. The river turtles (Triony- choidea. 3. An increase of the phalanges . 143). 144. 154). the metatarsal straight. as has digits occur among the Cotylosauria {Pariasaurus). 154) proximally. Crocodilia. In those reptiles which have a is fifth tarsale. Dromab). the primitive phalangeal formula suffered a reduction to 3. 145. A peculiarity of the fifth metatarsal among the Diapsida (Figs. The chameleon lizards have the phalangeal forphalangy. Fig. or many of them. 3. which may rather be ascribed to a secondary hyper- More than three phalanges have also been observed in some Pleurodira. 137 a. and the absent. though the oldest theropods (a. (Fig. 154) have normally four phalanges in the fourth and the fifth digits. and especially the Dino- been mentioned above. had two or three phalanges in the fifth toe. Anomodontia. 153 b). and various examples of partial reduction of the postaxial sauria. 144 b. A striking peculiarity is seen in with the correspond- the greatly reduced second phalange of the third toe and the second and third 138) of the fourth toes. the phylogenetic relationships. 140 d) and more or less hook-like shape. mals. in other dinosaurs it is the third toe that is the stoutest. the later ones (Fig. Hyper phalangy and Hyperdactyly. 4. 3. 155 d) have only the hook-shaped metatarsal left. in all cases doubtless to be ascribed to the grasping or clinging habits. 2. 3 for both fore and hind feet (Fig. lates In all such cases the metatarsal articufifth tarsale is with the fourth tarsale. mula 2. hand of the therocephalian Scymnognathus (Fig. and the Sauria is Chelonia (Figs. and perhaps also in those reptiles in which its the foot had become more or less erect or digitigrade before entire loss.

and from two additional cartilaginous phalanges in the to ten ossified ones in the Cetacea. and except in the Chelonia. known in the ichthyosaurs only among reptiles (Fig. 153 a). theories have been proposed to account for their That they cannot be due to the ossification and separation of the normal epiphyses in reptiles epiphyses. . 146). of the fifth toe. 140). is of the far num- ber of phalanges in both front and hind the normal number. one or two sirenians. though without Mesosaurus (Fig. 153 a) the fifth toe has two extra phalanges. 151 b) from the middle Pennsylvanian has the same number and arrangement of the phalanges as in the Cotylosauria (Figs. and the only examples of hyperphalangy in the order Chelonia. 128. and and Tylosaurus (Fig. observed in 151 b). Fig. 6. chondrification of the mesopodials. and the Cretaceous bird Hesperornis. Platecarpus (Fig.THE LIMBS above the normal number (hyperphalangy) and dactyly) is 199 of the digits (hyper- known only in swimming animals. It may perhaps more as sculling organs after the manner of seals. In web-footed swimming animals there is sometimes especially a tendency toward the elongation Eosauravus (Fig. there was an increase feet. 159) the digits are elon- gated. as the earliest foot known {Eosauravus. certainly an acquired character. 147). 3. for these reptiles at least had no Like the additional epipodials of the plesiosaurs and ichthyosaurs. In some if not all Proganosauria (Fig. 2. in the genera Clidastes (Fig. In the Mosasauria there was a progressive increase in hyper- phalangy as observed (Fig. and as many as six in the fifth. a river turtle. 158. as (Fig. they are accessory. that is. sea otters. 158 c. as twenty-two phalanges are many known ichthyosaurs have even more. possibly but very improbably a primitive character. concomitant with the progressive In certain plesiosaurs as in the third digit. In all strictly aquatic reptiles (Figs. Lariosaurus 149). 4. 148). In Trionyx. new ossifications in the mesenchyme and not reversions to a primitive fish-like fin. is quite evident. 133) and modern lizards (Fig. 5. Hyperdactyly. five phalanges have been observed in the fourth toe. Various origin. sometimes beyond A like hyperphalangy observed in the marine mammals. 148) and Tylosaurus from one or two to as many as six or eight additional phalanges. d). indicate the use of the hind legs additional phalanges. in all of which the fifth toe is very long and strong.due to is and certain the same causes.

sides of the paddles. which the . shows an extensive lateral movement of the foot upon the leg in locomotion. Some of these accessory digits seem to have arisen at the It is a question . In such reptiles the toes always are and must be long. the direction of the foot in the is turtles. On the other hand. more like the feet of ducks and In crawhng reptiles (Figs. and the motion of the foot upon the epipodials is largely lateral. Anomodontia. ac- may count for the coossification of the heel bones. In the modern lizards (Fig. In such locomotion long toes would be a hindrance. 128) the feet are directed more outwardly. The structure of the feet in the early forms. both by a reduction in the numbers and by a shortening of the segments. frogs. and likewise resulted in the reduction of the phalanges and shortening of the toes. Doubtless this same more mammal-like or turtle-like mode of progression was characteristic of the Dromasauria. were oar-hke and both the plesiosaurs and ichthyosaurs the feet of the mosasaurs were webbed. i. and Theriodontia. 139 a) the angle between the leg and foot in locomotion probably much more so than in the early forms. and especially the tortoises (Fig. The paddles flexible. and that number are accessory more probably the hypothat all above dactyly occurred after the ichthyosaur paddle was essentially evolved. More nine. The same acute angle between the foot and parallel in leg and the elevation of the heel bones have also resulted in the firmer ossification of the tarsal bones and their fusion. have but three while other later ones may have as many as whether three was the primitive number. but certain early forms digits. like Pariasaurus and Telerpeton have the astragalus and calcaneum fused. and this is acute. even as late as Sauranodon (Fig. and the results began to be seen in the reduced phalangeal formula of Pariain saurus. with the main axis of the foot more postaxial.200 THE OSTEOLOGY OF THE REPTILES usually the feet are pentedactylate. 140 d) and Sphenodon (Fig. Except among the Sauropoda and Stegosauria. as shown in Figure 158 c. more forward than lateral the digits of the feet on the two sides are brought more nearly locomotion. others by a of splitting of the digits. . One can imagine the difficulties of locomotion if our toes were six inches long! The Cotylosauria have short and broad feet. and they have been shortened. 153 b). with a large astragalus and calcaneum. and many of the later ones. Possibly the mode of progression was more turtle-like than lizard-like. 145).

In the still dactyls (Fig. the foot is more mesaxial is or preaxial. alus. the dinosaurs have rather long digits. to injury with the heel elevated far above In the bipedal Theropoda (Fig. as in the toe. digitigrade. c). the astragwhile perhaps never fully fused with the tibia. as formerly. Short toes and reduction of phalanges. was functionally between the leg and tarsus as in mam- mals. and the joint. c. In reptiles it is that is. where the fourth very seldom the strongest The chief joint the end of the tibia and the intratarsal. then.THE LIMBS toes 20I have become shortened and the phalanges on the postaxial side reduced. shortening the portion resting upon the ground. In those reptiles which walked more or less upon the there was a progressively closer articulation between the tibia and the astragalus. In the turtles has been produced by the exigency of the immovable shell. while actually. In all such reptiles with the more mammal-like mode of locomotion. like the reduction of the digital formula in the plantigrade reptiles. mam- mals. as in birds. giving a firmer and closer ankle which otherwise would have been subject the ground. or lesser twisting of the epipodials upon the pro- . between the foot and legs in mammals is between first row of the tarsals. acquired a long ascending process which fitted closely into a groove in front of the distal end of the tibia. 156 a. between the first and second rows of the tarsus. 155 d) the astragalus more elongated feet of the pterobecame indistinguishably fused with the tibia. intratarsal. toes. and of locomotion mode by the greater podials. but they had become distinctively digitigrade. mal-like this mean a more mam- and posture of the feet.




it and frogs when seems impossible that all should have been evolved from the same ancestor with procoelous vertebrae. for instance. and the problem of all classification is to distinguish between them and those characters due to heredity. called zygosphenal. the lesser branches genera and famihes. Did they occur rarely in organisms they would not trouble us much. In the adaptation of its progeny to various provinces and modes of hfe they have divided into innutimes. pterodactyls. others have continued to modern none has ever reunited with another branch. It is easy enough it is relatively to distinguish the twigs and smaller branches. . and especially the early or primary ones.CHAPTER VI THE PROBLEM OF CLASSIFICATION One who has studied attentively the skeleton of reptiles cannot fail to be impressed with the fact that similar or even apparently identi- cal structures have arisen in different orders. lizards. and each division again into two. but often very difficult to determine where the limbs united with the boughs and where the boughs joined the would be dichotomous. Until we have learned to distinguish common them our classification must remain more or genealogy. and branch dividing into two. and we must often treat groups of organisms as though perfect classification first trunk. differing very slightly from its ancestors. The twigs we call species. The true end of all classification is Some time in the Carboniferous period there was but a single kind of reptile. and from this reptile has descended all the kinds that have ever lived. occur in crocodiles. Procoelous vertebrae. the limbs orders. less artificial. and the main boughs subclasses. For such resemblances the convenient term homoplasy has been proposed. Snakes. A limb. each bough. some lizards. but merable branches. but an approximation even to such a classification cannot be attained. even though small. but their evolution from a common ancestor is impossible. classification is Our object in to determine these branches. but they are everywhere in nature. Many of these branches were feeble and of short duration. and certain Stegocephalia have a peculiar mode of articulation of the vertebrae.

As the only air-breathers of paleozoic times were amphibians and reptiles. since man has exerted a powerful influence upon them. that in classifying them. in classification to trace the history of each species. or have been the result of common environmental influences. that organs or functions . possibilities of great developments. a few hundreds. And our chief difiiculty in doing this is to determine whether the resemblances that they show to each other have been due to descent and common heritage. and order to its separation from allied forms.2o6 THE OSTEOLOGY OF THE REPTILES common center. family. and rapidly or slowly in different kinds of organisms evolution and extinction have replaced the faunas and floras many times. for instance. Many of the first groups of species that branched as twigs from the common stem were the ancestors of orders. and paleontological history is and always will be imperfect. and doubtless there has been no time since the first general invasion of land by air-breathers that the number has been less it may have been greater. The problems are hard and always will be hard because actual proofs of heredity must ultimately rest on the facts of paleontology. we are aided by general laws which have obtained recognition among students of extinct animals. the minor ones greatly is. succeeding species became more and more is restricted in their potentialities. There are to-day living upon the earth about twenty thousand species of air-breathing vertebrate animals. radiating from a And it is also evident that such divisions occurred rapidly. there must have been. to the Jurassic. probably every bone of the human skull to the primitive rep- . then. . would be much easier. genus. Nevertheless. we can back trace. Our chief object. and to give to each minor and major group a name and place. First of all. all of them. for they held. In all probability the earth since remote ages has always been as densely populated with living organisms as it is at the present time. — — scores of thousands of their kinds. but never the original ones. by the law that evolution is irreversible. By the general law that there has been a continuous loss of parts. we know but increased. in tracing the genealogies of organisms. the major problems Had we records of all that have lived. from the Mississippian during the time that they reigned supreme. millions of years. once lost can never be regained by descendants similar organs or similar functions often.

Chameleon lizards have only about sixty vertebrae. may vary extraordinarily within an order. we have seen. that surrounding the and in consequence it furnishes the reliable characters for the discrimination of the larger groups. vis chief factors of evolution is have been environment and There more or less impulse due to heredity. And the teeth of reptiles. and exits crescences occur only in the later history of any race. horns. The chief problem. the subclasses or super- orders. Furthermore. that is. to unite in the same group a lizard and a turtle because each happens to have eight cervical vertebrae. in any classification nation of the more conservative hereditary characters of the skeleton. have little value as criteria of relationships. then. The two heredity. other lizards may have a hundred and ninety-four. of articulation of the ribs. and sometimes perhaps not more than specific. More conservative is the arrangement and of all has mode brain. but there were twice or thrice as many bones in the older forms as there are in recent ones. Some forms. The pectoral and pelvic girdles have been influenced less by environmental conditions. never at beginning. the structure of the feet stfll less in adap- tation to life conditions. spines. It would be absurd. The number is seldom of more than generic value. small animals has never been evolved from a race of large animals. though their location may be more conservative. And there has been in general an increase largest animals and a race of in bodily size in every phylum. while snakes of the same order may have as many as four hundred and fifty. those which have been influenced less by environmental conditions. The have always lived at or near the end of their race. .THE PROBLEM OF CLASSIFICATION tilian or 207 amphibian skull. protuberances. the determi- how much to similar environmental influences. so adaptable are they in shape and number to food habits. unlike those of mammals. Most conservative most been the structure of the cranial region of the skull. a sort of a iergo that related forms. though seems to influence evolution along parallel lines in we are never sure how much is due to it and is the relative importance of structural characters in the absence of the actual connecting links insensibly uniting different forms. parts of the skeleton are very variable even in nearly related The number of vertebrae in the spinal column. for instance.

It is only in the gradual crystallization of opinions that stability finally results. largely a matter of the personal opinion. the turtles. the have twenty-two or twenty-three pairs and five or six unpaired bones. merely represents the present state of our of the opinions of those best qualified knowledge and the consensus to decide as to their value. and more and more subordinate details. The crocodiles have but twenty-four pairs and six unpaired bones in the adult. then. however. one pair that is fused in the crocodiles. Fortunately. they could by possibility have regained them when once lost. But not all are the same. is and this crystallization never complete. and all have lost some bones of the feet. No two persons see them from the same viewpoint and consequently no two persons whose opinions deserve consideration ever wholly agree as to the value of characters in classification. the lizards. Any system of classification. The persistence or loss of bones furnishes many certain evidences of relationships and descent. however. for per- fection postulates complete knowledge. the lizards have at the most twenty-nine pairs and five single bones. and so on. The crocodiles have turtles three or four pairs that have been lost in the turtles. . So long as science endures. more or less influenced by the classifier's individual opinions. the increase of knowledge affects less and less the major principles. The relative importance of all such characters in classification classifier's is. No classification will ever be perfect.2o8 THE OSTEOLOGY OF THE REPTILES primitive reptiles that The most we know had no less than thirty- seven pairs and four single bones in the skull. Each order must have de- no scended from ancestors that had the persistent bones. several bones lost or fused in the turtles. All reptiles since Triassic times have lost four bones in the pectoral girdle. new facts will be discovered to influence our opinions.

209 .

in two or more rows posteriorly.] Pariasauria. Cotylosauria. Triassic to Cretaceous. Labidosauria. D. Ed. G. Most primitive. Skeleton primitive. carapace with peripheral plates. not perforated.] [But see footnote on page 126. 1 Jurassic to Recent. Eunotosauria. a single coracoid. Insectivorous. plates. Jurassic to Recent. Triassic. A. 3. Lower Permian. Pleurodira. Lower Permian. Lower versely molariform. B. crushing. A. at least eighteen dorsal vertebrae. in upper jaw and dentary. Skeleton primitive. no peripheral River turtles. Cervical vertebrae amphicoelous or concavo-convex. the posterior transMalacophagous. posterior teeth transverse. Permian. Middle and Upper Permian. an acromion. Neck retracted vertically. . Permian. two coracoids. Temporal 1. interparietal on dorsal surface. Teeth blunt or pebble-like. Skeleton not primitive. their ribs expanded to meet on the dorsum. Cretaceous to Recent. Seymoiiria. Trionychoidea. Upper B. pelvis united with plastron. Tail long. Cryptodira. Teeth anisodont. interparietal obsolete or absent. crushing. their ribs expanded to meet on the dorsum or a dermal layer of bony plates. in single rows [E. no supratemporal. two coracoids". conical. Procolophonia. Lower intertemporal and otic notch. ten dorsal vertebrae. ten dorsal vertebrae. LiMNOSCELis. the posterior flattened F. no supratemporal. in single row. Amphichelydia. Teeth anisodont. Middle Permian. neck not retractile. C. numerous on palate and coronoid. and a dermal layer of bony plates. region of skull roofed over. in single row. teeth conical. interparietals large on dorsal surface. No mesoplastra. or secondarily emarginated. Incisors conical. Lower Permian. C. [compressed] and crenulate. Neck retracted laterally. DiADECTOSAURiA. their ribs not expanded. Neck retracted vertically. Pantylosauria. Testudinata (Chelonia). Teeth heterodont. interparietal on posterior surface. D.CHAPTER VII A SYNOPTIC CLASSIFICATION OF THE REPTILIA ANAPSIDA. Teeth anisodont. 2.

3. teeth small. phalangeal formula primitive. no teeth on palate. teeth small. dorsal spines short. teeth strongly anisodont with diastema. C. B. Vertebrae platycoelous no dorsal intercentra. Dentition subisodont or absent. NoTHOSAURiA. 211 SYNAPSIDA. Placodontia. III. vertebrae notochordal with persistent dorsal intercentra. coracoids and feet unknown. vertebrae deeply amphicoelous. articulating with diapophysis. Plesiosauria. phalangeal formula 2. long. dorsal spines more or parietal less elongated. no secondary palate. A. Malacophagous. Dorsal spines elongate. Skeleton rarely notochordal. less downward rate in locomotion. no secondary palate. Anomodontia. less primitive. interparietal and tabulars present. interparietal or tabulars. propodials turned more or Therapsida. 3. Triassic. later forms. and tabulars present. Edentulous or with long canine. A. Theromorpha. Temporal opening extending to parietal in large. 3. propodials horizontal in locomotion. an acromion. or canine and molars. 3. phalangeal formula primitive [?]. Carnivorous. C. Lower Permian. Sauropterygia. quadtemporal opening surrounded by postorbital and squamosal. Lower Permian. dorsal spines not elongate. Neck more or less elongated. subisodont. 7. Triassic to . Edaphosaiiria. 3. Malacophagous. feet webbed. A. Insectivorous. limbs paddle-like. A single temporal opening. .A SYNOPTIC CLASSIFICATION OF THE REPTILIA II. vertebrae amphicoelous. Dromasauria. Skeleton primitive. Permian. D. with bars. two coracoids. 3. Upper Triassic. Upper Permian. 6. body with dermal bones. dorsal ribs double-headed. teeth on palate bones. no supratemporal. Lower Permian. . no interpterygoidal opening in palate. PoLiosAURiA. Middle and Upper Triassic. dorsal intercentra absent or unknown. at least one pair of caniniform teeth. isodont. Carnivorous. A single coracoid (? Placodontia). Pelycosauria. conical. Theriodontia. Marine. Triassic. DiNOCEPHALiA. the propodials palate with openings. hyperphalangic close of Cretaceous. a rudimentary secondary palate. SYNAPTOSAURIA. 3. dorsal ribs single-headed. dentition more or less heterodont. Jaws and closed palate with heavy pavement teeth. isodont. teeth small. Skull massive. D. primitively below the poslorbito-squamosal arch. 5. inter- B. 4. 3. phalangeal formula 2. phalanges and teeth variable. Amphibious. Caseasauria. Upper Permian. A single temporal opening bounded below by postorbito-squamosal arch. B. phalangeal formula primitive. phalangeal formula often reduced. Lower Permian.

212 IV. {d) Amphisbaenia. 9. the brain-case more or . An epipterygoid. L. but small. Mus. procoelous. the neck Phalangeal formula 2.] — Ed. — [Sometimes present. Phalangeal formula primitive. Proganosauria. Protorosauria. V. Not more than seven cervical vertebrae. 3. Quadrate fixed. hmbs paddle-like. procoelous. five cervical vertebrae. Middle Triassic to Upper Cretaceous. Lower Permian to Jurassic. Ophidia (Serpentes). A [For a more comprehensive classification of the Lacertilia. see C. 289-481. 3. Vertebrae Limbless. An epipterygoid present-.. Ribs articulating more or less exclusively with centrum. ijb) Nine or more cervical vertebrae. Ichthyosauria. Quadrate freely articulated proximally (strep- tostylic) or secondarily fixed. Lower Permian. (c) hyperphalangic seven cervical vertebrae. separated by postorbito(? Youngina). K. limbed or limbless. mem- branous anteriorly. vertebrae am(a) Kionocrania phicoelous with persistent dorsal intercentra. vertebrae procoelous. Marine reptiles with short neck and all aquatic adaptations. the quadrate fixed.1 . vertebrae procoelous. Hmbless or with vestigial . No epipterygoid or clavicles^. A. Rhiptoglossa. no supratemporals or tabulars ^ openings. mandibles united by ligament. Vertebrae amphicoelous (PSaphaeosauridae). {e) front legs. Squamata. 4. between parietal and postorbito-squamosal arch. no epipterygoids no temporal arch. 4. supratemporal (tabular) persistent. the quadrate secondarily fixed. 3. 4 (6). phalangeal for- mula B. 2 ^ Amer. DIAPSIDA. Hist. 8. . Marine reptiles. 10. G. Brain-case enclosed by descending plates from parietals and frontals. XLViii. Nat. Platynota. THE OSTEOLOGY OF THE REPTILES PARAPSIDA. Vertebrae amphicoelous. Cretaceous to Recent. Upper Cretaceous. Aquatic or terrestrial. [Absent in Dibamidae. no chevrons. No epipterygoid or temporal arch. Vertebrae procoelous. Bulletin. Two temporal squamosal arch. with zygosphenes. Lacertilia {Sanria)} Parietals never united to basispheless noid by descending plates. and tail elongate. 11. no dorsal intercentra. 2. Lower Cretaceous to Recent. A single coracoid. Quadrate fixed. or procoelous and no dorsal intercentra. 1923. Oligocene to Recent. N. Pythonomorpha. Skeleton largely primitive. — Ed. Aquatic. eight cervical vertebrae.] Camp. an epipterygoid present. Skull imperfectly known. Cretaceous to Recent. Phalangeal formula primitive. Phalangeal formula primitive. A single temporal opening. 4. 5. Oligocene to Recent.

. Littoral reptiles. Rhynchocephalia. Skull elongate. External nares terminal. or tabulars. no supratemporal. I). A. vertebrae amphicoelous or procoelous. usually an antorbital opening. A. Triassic. no false palate. pubo-ischiatic opening small. Face elongate. dichocephalous. a well-preserved antorbital vacuity present. Uppermost Cretaceous and Lowermost Eocene. 15. A. No epicondylar foramina. phalanges not reduced. 213 no clei thrum. holocephalous. External and internal nares terminal. not meeting in ventral symphysis. body with dermal scutes. Skull only known. EusuCHiA. Cursorial reptiles. Sphenodontia. is ^ [According to Broom. no supra temporal. Terrestrial or marsh reptiles. the epipodials long. articulating with arch only. Imperfectly known. Premaxillae beak-Uke. Phytosauria. 12. Pelvis more or less plate-like with small pubo-ischiatic vacuity. Phalangeal formula primitive. Parasuchia. Legs short and stout. with palatal teeth. Often reduced postaxially. AA. vertebrae amphicoelous. no sclerotic plates. Upper Triassic. with decurved premaxillae and crushing teeth on palate. Jurassic to Recent. With dermal plates. Pubes excluded from acetabulum. PsEUDOSUCHiA. dorsal intercentra not persistent. ? Proterosuchia. External and internal nares remote from extremity of slender face. Triassic. Rhynchosauria. No antorbital foramen in skull. Pelycosimia. an entepicondylar foramen. External and internal nares near extremity of face. Only skull known. Vertebrae amphicoelous. Teeth protacrodont or acrodont. Upper Jurassic to Recent. no parietal foramen. C. no teeth on palate. postaxial digits reduced. tabulars or interparietals. with external nares ? Thalattosauria. Triassic. A. Limbs paddle-Hke. ?Eosuchia. interparietal and tabulars present. palate with teeth. Subaquatic. posterior.] . Skull broad. Upper Permian. — Ed. No antorbital vacuity^. an antorbital vacuity. interparietal. DIAPTOSAURIA. B. Skull elongate. dorsal intercentra persistent. Littoral reptiles. Dorsal ribs articulating with intercentral space or centrum and arch. vertebrae variable. legs elongate. Amphibious. C. Ribs attached by single head to centrum. Typically with large pubo-ischiatic vacuity. A secondary palate.A SYNOPTIC CLASSIFICATION OF THE REPTILIA single coracoid. Triassic. Triassic. Aquatic. 13. labyrinthine. 14. 16. a parietal foramen. ARCHOSA URIA Dorsal ribs. the nares terminal. Choristodera. pelvis with large pubo-ischiatic vacuity. no teeth pubo-ischiatic vacuity. Crocodilia. B. anteriorly at least.

Ceratopsia. 19. Triassic to close of Cretaceous. paddle-like. 18. Quadrupedal. Without dermal bones. DINOSAURIA. pubes composed of a spatula te anterior process not meeting in symphysis. quadrupedal. presacral vertebrae amphicoelous. Volant reptiles. tail long. Jurassic. Pelvis normal. limb bones not hollow. Cretaceous. the pubes meeting in a ventral symphysis. A. Front legs reduced. Fourth finger greatly elongated to support patagium. and a more or less elongate posterior process. Saurischia. No predentary or rostral bones in skull. Theropoda. C. Lower Jurassic to close of Cretaceous. Pterosauria. No dermal ossicles. Postaxial digits reduced. the front legs more or less reduced. Pterodactyloidea. More 17. with dermal armor of plates and spines. THE OSTEOLOGY OF THE REPTILES Thalattosuchia. Stegosauria. A. B. Uppermost Cretaceous. tail short. Skull very large with bony horns and a posterior expansion fringed with scutes or spines. the bones pneumatic. Wing metacarpal longer than forearm. Herbivorous.' Neck shorter. Skull with or without teeth. the front legs but Httle or not at all reduced. a predentary bone in mandible. Vertebrae procoelous. Postaxial digits reduced. Pterodermata (Rhamphorhynchoidea). Upper Jurassic to Upper Cretaceous. Sauropoda (Cetiosauria. Upright-walking. bipedal. Ornithopoda. . . or some or all opisthocoelous. Ornithischia iOrthopoda) Herbivorous. B. or less upright-walking reptiles. B. Marine. Vertebrae amphicoelous a terminal caudal fin.214 B. Jurassic to close of Cretaceous. Jurassic. A. . Skull with teeth. Wing metacarpal not longer than forearm. Carnivorous or secondarily herbivorous. bipedal in gait. Jurassic. no dermal plates. skull small. sclerotic plates in orbits. Opisthocoelia). neck and tail elongate. bones solid. Quadrupedal.

and possibly also in the tail Free ribs are present to the sacrum and in in Seymouria only. Pantylus. the neck was always short. some Captorhinidae. The earliest known member of the order dates from the Middle Pennsylvanian. the tail of . None has all the primitive characters given in the Hst. Diadectes. The ectopterygoid has not yet been certainly demonstrated in any genus. not emarginated: skeleton primitive. The body was never slender. invertebrate feeders less for the marsh reptiles. Dermal ossifications of any kind are known in but a few genera. nor the legs long. lowland. but the any form are few. or secondarily emargisides. The parietal foramen is absent in Pantylus. not only the oldest geologically. structure. They may be discussed under three groups. probably the ancestral stock of all later Amniota. varying in size from than one to about ten feet in length. or chmbing. Two pairs of sacral ribs are present in all genera except Seymouria and Diadectoides There are from twenty- two or twenty-three to twenty-six presacral vertebrae. the latest from the Middle Triassic.CHAPTER VIII THE SUBCLASS ANAPSIDA Temporal region of skull wholly roofed over. never cursorial or most part. 1. I. nated from the not perforated. Parasternal ribs are known only . the Seymouriidae the supratemporals are absent in the . the body was probably covered with horny plates or scales. They were very variable in size. order of reptiles. though probably present. Captorhinidae. subaquatic. which include their chief evolutional modificalosses or modifications in tions. absent wholly in the lumbar region of the others. Carboniferous and Lower Permian The intertemporal bone and the otic notch are known in but one or two genera. and the Pariasauridae. and habits: as known. in the last-named family and Sauravus. the tabulars also in Labidosaurus. ORDER COTYLOSAURIA An Temporal roof complete. but more primitive in structure than any other.

Fig. Enlarged. Eosauravus: Part of skeleton. i6i. Speci- 2l6 . men in National Museum. from above.

One on jaws and dentaries. Small in reptiles six to eight or ten inches length. Dichocephalous ribs are known is only in Seymouria and Pantylus. Vertebral spines vestigial..THE SUBCLASS ANAPSIDA moderate or considerable length. United States. the vertebrae with broad arches. insectivorous rep- than three feet in length. About two feet long. reptiles Eosauravus (Middle Pennsylvanian). Thevenin. Family Seymouriidae. Primitive. fully 217 Two centralia pedis are doubt- known only in Labidosaurus. Family Gymnarthridae. Teeth conical in a single row Fig. B. Skull Family Sauravidae. France. twentieth natural size. (? Conodectes Cope). about one foot in length. 162. Skeleton and life restoration of Cotylosaurs: A. Diasparactus. Teeth obtusely . Slender and intercentra unknown. The cleithrum vestigial in some. Seymouria. United States. insectivorous or invertebrate feeders. tiles less terrestrial. After Case. Sauravus. Seymouria Broili Cleithrum vestigial or absent.

Desmatodon Case. Family Captorhinidae (Labidosauria). Small pointed. Skull short. Pariotichus Cope. vestigial. insectivorous or invertebrate feeders. transversely molariform posteriorly. Family Pariotichidae. one or more on mandible. Phanerosaurus Meyer. recurved in single rows. United States. Elongate. No supratemporals and. Family Diadectidae (Diadectosauria). Cleithrum rather large. Family Stephanospondylidae. Diadectoides Case. Teeth obtusely more rows. known. Chilonyx Cope. short. Cardiocephalus Broili. Bathyglyptus Case. Maxillae with two rows of transCleithrum large. Teeth subcorneal. THE OSTEOLOGY OF THE REPTILES Parasphenoid large and broad. Bolhodon Cope. conical. United States. . United States. Imperfectly known. Teeth conical. Ectocynodon Cope. Skull short and high. Saxony^ Germany. Pleuristion Case. in one or terrestrial reptiles about one foot in length. From one in length. Cleithrum Spines of vertebrae Parasternal ribs present.Archeria Case. Family Bolosauridae. Teeth and part of skull only. United States. Helodedes Cope. Genera Incertae Sedis: ChamasaurusV^i\\iston. Lahidosaurus Cope. United States. Animasaurus Case and Williston. no tabulars. cuspidate. Captorhinus Cope. imperfectly known. Diadectes Cope. Isodectes Cope. From about five to about eight feet in length. Dermosupraoccipitals confined to occiput. in Teeth obtusely flattened. two or more rows on maxillae. United States. subaquatic or marsh reptiles seven or eight feet in length. verse teeth. Appendicular skeleton unknown. Teeth incisiform in front. Diasparactus Case. Family LiMNOSCELiDAE. Small reptiles. in Lahidosaurus. and the skull imperfectly known. to three feet invertebrate feeders. Bolosaurus Cope. Gymnarthrus Case. as also the skeleton. Stephanospondylus Stappenbeck. Puercosaurus Williston. their arches broad.2i8 pointed. Vertebrae with hyposphene and hypantrum.

m^ 219 .

saurusW2it's. Astragalus and calcaneum sometimes fused. relatively smooth. Embrithosaurus 3. tail Skull elongate and compressed in front. intimate structure not well known. Scapula with acromion and screw-shaped fifth tarsale glenoid fossa. No parietal foramen. DermosupraoccipiPost-temporal Spines of Tabulars large. Two or three sacrals. scutes. and calcaneum fused centrale and unknown. six or Teeth in a convex on the outer side with seven cusps arranged Astrag- around alus their borders. Family Pariasauridae broad and short. 4. Middle and Upper Permian (Pariasauria) . South Africa. in a single row. with palatal and coronoid jaws and mandible. elonParietal foramen large. Family Pantylidae (Pantylosauria) two feet in length. primitive in others. including between them and the squamosal a large otic notch. fiat. Ectopterygoids distinct. pos3 in sibly absent. size. Limnoscelis Williston. 3. so far as known. Pantylus Cope (? Ostodolepis Williston). Russia. Propappus Seeley. Coracoids free in maturity. 3. Pariasaurus or an allied genus. teeth. 2. cleithrum. Terrestrial reptile less than Skull low. its Large lowland cotylosaurs Skull with protuberances. Body covered with small bony scutes. Small reptiles a foot more in length. Lacrimals small.on. reaching nine or more feet in length. No supratemporals. Pariasuchus Haughton and Broom. . a single row of short teeth in [upper] Skeleton imperfectly known. . Phalangeal formula of front feet unknown. . Skull triangular. one genus. gate anteroposteriorly. Body with several rows of bony dermal Pariasaurus Owen. single row. Lower and Middle Triassic or Family Procolophonidae (Procolophonia). United States. openings of considerable vertebrae small. Radiale and fifth carpale unossified. United States. Anthodon Owen. Orbits very large. Brady- Watson. short. elongate. Phalangeal formula believed to be 2. conical. sometimes excluded from nares.2 20 THE OSTEOLOGY OF THE REPTILES Cleithrum small. also centrale and fifth tarsale. Parasternals No sometimes present. . Teeth in front behind transverse. tals small or vestigial.

the palatine primitive. covered with dermal ossifications. 2. repre- sented by incomplete specimens which have recently been described by Watson. with numerous teeth. This group at present is known by a single species. dal. Eunotosaurus africanus Seeley. feet unknown. Tail probably small. . Family Elginiidae. External nares terminal. apparently in median and lateral rows. capitular attachment of ribs on anterior Pectoral and slender. with long horns in tabular region and numerous conical protuberances. The known characters ally more closely with the Coty- losauria than with the Chelonia. as either order will require Watson has urged. from the Middle Permian of South Africa. limbs. from whom genus is the above characters are taken. Femur moidally curved. contiguous at their borders.THE SUBCLASS ANAPSIDA South Africa. and an incomplete carapace dermal bones. ORDER EUNOTOSAURIA of Primitive terrestrial reptiles with reduced dorsal vertebrae. South Africa. Intimate structure and skeleton unknown. Koiloskiosaurus Huene. Crowns of teeth denticulate. Elginia Newton. Saurosternon Huxley. Europe. Dorsal region strongly convex. Sclerosaurus Meyer. For that reason it may be left in an independent position until further discoveries furnish more details regarding the skull. broad behind. Maxillae and premaxillae with a single row of small teeth. the second to ninth ribs with a vestigial tubercle. Orbits not elongate. notochor- with vestigial spines. Thelegnathus Broom. progressively broadened. Middle Permian. Scotland. Leptopleuron 221 Procolophon Owen. Skull triangular. Vertebrae slender. pelvic girdles primitive. is The skull is doubtless wholly roofed over. Eunotosaurus Seeley. ex- panded ribs. That the it intermediate between the true Cotylosauria and the Che- Ionia seems very probable. To locate with many modifications in their definitions. sig- part of centrum. Owen {Teler- peton Mantell). it and cara- pace.

3 or 2. The prefrontals meet in the median line. Only rarely. covered by the large postorbitals. Ten dorsal vertebrae enclosed a more or less complete carapace. and the phalangeal formula 2. 2. supracoracoid foramen. have the outer fingers of the hand more so twisted than three phalanges. pelvis with large openings. 2. In addition to the bones mentioned above.22 2 THE OSTEOLOGY OF THE REPTn. Because of the inflex- ible fied. broad body. in certain aquatic types. roofed over.ES 3. jugals. A single coracoid. 2. the temporal region. dependent upon the character of the carapace. species. the former with but a single living Regarding the general dispute. the latter with more than two hundred. the turtles [beaked]. a rather short skull. dermosupraoccipitals. the stapes is slender. The prevomers are united. As in the dinosaurs there is a greater or less reduction of the postaxial fingers and a strengthening of those on the other side of the feet. there is no interpterygoidal opening. the supratemporals. carapace the structure and posture of the limbs are is much modi- upon the humerus that the foot is The forearm brought more or less directly forward in walking. always reduced. The more generally (in- accepted classification recognizes four suborders. 3. 3. and are always toptery golds and usually the lacrimals are absent. exposed by the emargination of the roof bones. Dollo. a flexible neck of eight vertebrae. probably because of an incipient hyperphalangy. with the fifth toe usually more so. squamoand quadratojugals. 3. the Athecae and Thecophora. the nasals are often absent. the Cryptodira . usually without a when sals. a short. No order of reptiles is so unequivocally distinguished from all Jaws and except in Stegochelys of the Trias they are wholly toothless. Eight cervicals. an ossified plastron. The pectoral girdle is composed of a furcate scapula and a single coracoid. The carpus and is tarsus are much modified. Phalangeal formula always reduced. to still 2. and an osseous carapace and plastron. and there may be an incipient secondary roof to the palate. and Hay there are two chief divisions or suborders. ORDER TESTUDINATA OR CHELONIA of skull usually Temporal region tabulars absent. classification of the Chelonia there is still According to Cope. tortoises. and in. the postfrontals and ecothers as the Chelonia.

Carapace united to Upper Jura. since the Eunotosauria have both the expanded dermal ribs corresponding with the ordinary carapace of the turtles. Vomer and parasphenoid with small teeth.THE SUBCLASS ANAPSIDA 223 eluding the Athecae). Europe. Skull wholly Toofed over. plastron by narrow buttresses. ^ [ = Triassochelys Jaekel. Turtles in which the primitive dermal armor lete or abolished. (Jaekel. much strengthened. the Pleurodira. Helochelys Meyer. Cretaceous. Amphichelydia. is Thecophora. One sacral vertebra. Dermochelys coriacea. the interpretation of the is characters of Eutiotosaurus. Scapular girdle with short proscapular process or acromion. amphicoelous.^ Proterochersis Fraas. of various Cretaceous Thecophora. perhaps. the carapace formed obso- by expansions of the ribs.] . Cervical vertebrae Family Pleurosternidae. the cervical vertebrae Mesoplastra present. Platychelys Wagler. as given on a preceding page. Pleurosternum Owen. ? Stegochelys Jaekel. — Ed. Skull elongated. this theory is If however. Chelyzoum Meyer. Neck am- phicoelous or concavo-convex. Family Proganochelydidae. coracoid distally expanded. North America. Nasals and lacrimals distinct. not retractile. but they have been ac- counted for in other ways. a supracoracoid foramen. and usually peripheral plates. A single living species. neural plates. Germany. and an overlying carapace composed of rows of plates like those of Dermochelys. correct. Progano- chelys Baur. Turtles which retain the primitive dermal armor. The definitions of the Athecae and Thecophora given by Hay are as follows: Athecae. A. with at least traces of the subdermal expansions connected with the ribs.) Upper Trias. Evidences of this primitive external series are found as vestiges. and Trionychoidea. with ribs. Neck with free dorsal plates. coracoid short. Germany. Nine dorsal ribs (costals) and seven cervical vertebrae. Suborder Amphichelydia short. Glyptops Marsh. more or less fused with scapula.

Chelodina Fitzinger. Skull roof complete with horn-like protuVery large turtles. Africa. ? Polythorax . Eocene. Pelomedusa Wagler. distinct. Miolania. South America. Temporal roof of skull complete or much emarginated. Africa. Suborder Pleurodira Neck withdrawn laterally. Family Miolanidae. India. Family Pelomedusidae. Eocene. Eubaena Hay.Siy Naomichelys'H. Head withdrawn in a vertical flexure. Neurankylus Lambe. Vo- Upper Cretaceous. Eocene. Charitemys Hay. Australia. North America. B. Pubes and ischia suturally united with plastron. K. Epiplastra in contact with hyoplastra. — G. Taphrosphys Cope. Carapace united tresses. Peripheral bones of carapace present. Ba'ena Leidy. Cope.224 THE OSTEOLOGY OF THE REPTILES to plastron Family Baenidae. Vomer frontals separated. Boremys Lambe. United States. C. Recent South America. present. Mesoplastron mers present or absent. Thescelus Hay. Probaena'H. Cervical vertebrae for the most part with but one end concave. Mesoplastra absent or present. Hydraspis. United States. Naiadochelys Hay. N. Sternothaerus Bell. Upper Cretaceous. by strong but- Skull short. Africa. Europe. Pelvis not united with plastron. berances. Pleistocene and Recent. ? Lower Cretaceous. North America. Amblypeza Hay. Suborder Cryptodirai Carapace with marginal plates. Ba'ena Leidy. Pleistocene. No mesoplastron. Stereogenys Andrews. Miolania Owen.] . No nasals. Pterygoids not separating quadrates from basisphenoid. ' [Many genera omitted. the pre- Family Chelydidae. Pterygoids separating quadrates and basisphenoid.aiy. Bothremys Leidy. Uppermost (?) Cretaceous. AustraHa. Podocnemis Wagler. Pliocene.

at least two claws on front {? Toxochelys Cope Cynocercus Cope). Upper Jura. ? Allopleuron Baur. Catapleura Cope. Miocene. Argillochelys Lydekker.THE SUBCLASS ANAPSIDA over-roofed. Eochelone Dollo. Chelyopsis Beedin. loosely Small palatine foramen. connected with carapace. Skull mostly roofed over. Upper Cretaceous. Internal nares far forward. 225 Family Thalassemyidae. Desmatochelys Williston. Family Toxochelyidae. usually with one or more fontanelles. Limbs paddle-Hke. America. Nasals present. loosely attached. North America. Europe. Temporal region of skull more or less Neck short. Eosphargis Lydekker. Internal nares far forward. Eurysternum Wagler. Protostega Cope. North Family Cheloniidae. land. Europe. the plastron Skull large. Archelon Wieland. Europe. Allopleuron Baur. Eocene ?Lytoloma Cope. feet. Upper Cretaceous. Upper Cretaceous. Africa. Temporal region largely over-roofed. Palatines entering into internal nares. Porthochelys North America. Europe. . Lytoloma Dollo. temporal region wholly roofed over. Plastron loosely articulated. Procolpochelys Hay. Eocene. Scyllomus Cope. Lemhonax Cope. Skull almost wholly roofed over. Atlantochelys Agassiz. OHgocene. Plastron aquatic. Williston. Seeley. Europe. Osteopygis Cope. North America. Europe. paddle-like without claws. North America. Thalassemys Riitimeyer. turtles of large size. Upper Jura and Wealden. Marine Neptunichelys Wie- Family Protostegidae. turtles with paddle-like Hmbs and retractile. Family Desmatochelyidae. shields. (Rhetechelys. Tropidemys Riitimeyer. Ambulatory turtles. North America. not under-roofed. Erquelinnesia) Upper Cretaceous. Subaquatic. Pelohatochelys Chitracephalus Dollo. Head not Plastron loosely united to carapace. North America. the limbs Carapace greatly reduced. Europe. Marine elongate fingers. Plastron loosely connected with carapace. Erquelinnesia (Pachyrhynchus) Dollo. North America. Europe. Peritresius Cope. Carapace with Cretaceous.

Marsh turtles. Feet elongate. ? Anosteira Leidy. Basilemys Hay. Africa. Psephophorus. Baptemys Leidy. polygonal. webbed. Graptemys Agassiz. bony plates. in the dermis of the Eocene. Caudals procoelous. North America. Anosteira Leidy. North America. ? Clemmys Gray. Gafsachelys Stefano. the skull incompletely roofed. former grade into the — Ed. webbed. Europe. ? Acherontemys Hay. with mosaic of small. Marsh and river turtles. ? Gyremys Hay. Alamosemys Hay. Basilemys Kallistira Hay.226 THE OSTEOLOGY OF THE REPTILES turtles. North America. [Family Dermochelyidae. Agomphus Cope. as the . the carapace well Upper Cretaceous. Echmatemys Hay. Miocene. North America. Homorophus Cope. Family Emydidae. Tretosternum Owen {Peltochelys DoUo). Chelydra. Europe. North America. Eocene. the feet tortoises. Macrochelys Gray. Entoplastron T-shaped. Eocene.] unite the Emydidae with the Testudinidae. Egypt. Middle digit rarely reduced.^ Temporal region not tile. North America. Subaquatic. Recent. Emys.] Family Chelydridae. Upper Jurassic. Trachyaspis Meyer. Family Dermatemyidae. Anosteira Leidy. Marine turtles. Hay. 1 [Many modern taxonomists latter. Xenochelys Hay. roofed. North America. Caudals mostly opisthocoelous. Europe. Europe. Chelydra Schweiger. Leathery back. North America. Pleistocene and Recent. Pseudotrionyx Dollo. Neck retrac- Carapace low-arched. Paleotheca Cope. Marsh ? Upper Cretaceous. Notomorpha Cope. ossified. with reduced plas- tron loosely joined to carapace. Miocene. Europe. Oligocene. Plastron No parietosquamosal arch. Chrysemys. Dermochelys. Hoplochelys Hay. Temporal region not suturally united to carapace. Compsemys Leidy. Zygoramma Cope. Europe. Adocus Cope. Ohgocene. Eocene. not ankylosed to the ribs. Eocene. roofed. Europe.

North America. Testudo Linne. Eocene. Clemmys Ritgen. Africa. Terrepene Merrem. hypoplastra. Tes- Oligocene. and xiphi- Legs unknown. Helopanoplia Hay. gen. Testudo Linne. North America. D. Plastomenus Cope. Terrepene Merrem. Clemmys Ritgen. Skull SUBORDER TRIONYCHOIDEA in the Cryptodira. Asia. Europe. Axestemys Hay. Pleistocene. Openings between hyoplastra. North America. which lacks the peripheral bones and is covered with a leathery skin only and is flattened.THE SUBCLASS ANAPSIDA Miocene. the neck long. Temporal langes not roof reduced. Family Testudinidae. tudo. Uppermost Cretaceous. Chrysemys Gray. Europe. Pliocene. North America. Pseudemys. Hyoplastra. terrestrial. More than three phalanges in third digit. North America. Trachemys Agassiz. Stylemys Leidy. Miocene. Family Plastomenidae. feet elongate. plastra closely united. Hadrianus Cope. Ptychogaster Pomel. Uppermost Cretaceous and Lower Eocene. Africa. Plastron suturally united with the highly arched carapace. ? ? Clemmys RitPseudemys Gray. plastra. North America. Pha- more than two in number. Neck retractile. Three claws only. North America. Cotichochelys Family Trionychidae. the basiphenoid separating and neck as the pterygoids. 227 Trachemys Agassiz. Hay. North America. Aspideretes Hay. Oligocene. Miocene to Recent. herbivorous. Deirochelys Agassiz. Achilemys Hay. Plastron ligamentously united with carapace. Small to very large tortoises. Eocene. . hypoand xiphiplastra. River and lake turtles. Amyda Oken (= PlatypeUis Fitzinger). North America.

interclavicle and clavicles always present. The above definition will distinguish fairly well the Lower Permian forms from the Middle and Upper ones. the intercentra persistent throughout. name was proposed by Cope to include not only the all of Cotylosauria but the African genera of the order Therapsida as . amphicoelous.CHAPTER IX THE SUBCLASS SYNAPSIDA A SINGLE. Feet always pentadactylate. articulating intercen- more or less and with the arch. Fig. 164 his. Pelvis more or less plate-like. lateral. motion horizontal. jugal. Dorsal ribs double-headed. trally temporal opening. sometimes with vestigial cleithrum. Nor can we hope to reach a very satisfactory solution of the numerous problems much more is known of them and especially of the later Permian and Triassic forms included here in the same subclass. and postorbital only. Pectoral girdle with two coracoids on each side. There has been much discussion as to the rank and limits of the Paleozoic genera included under the above definition. ORDER THEROMORPHA Limbs and palate primitive. About seven cervical vertebrae. and the order Theromorpha till may Originally the be therefore accepted for the present with these limitations. Sphenacodon (Theromorpha). 4. Propodials in loco- Vertebrae notochordal or deeply biconcave. Restored skeleton. bounded primarily by the squa- mosal.

Skeleton and life restoration oi Dimetrodon (Theromorpha).Fig. 229 . 165. About eight feet long.

No dermal bones have been discovered in any member of the order. But the primitive characters were less constant. Doubtless they were more active and agile animals. and the two former are always confined to the occipital surface when . smaller. tabulars. The quadratojugal is The teeth are is often wanting on the prevomers the postsplenial never present in the mandible though there the posterior one of which is is a possibility of an additional coronoid. some may be wanting. sacral vertebrae. present. The intertemporal bone is never Fig. the longer neck. and their adaptive radiation was greater. the entepicon- dylar foramen is always present. THE OSTEOLOGY OF THE REPTILES and is still used often in that original sense or with the exclusion of the Cotylosauria. the only constant differ- ences from which are the perforated temporal roof. present. always present. and parasternal ribs are known only in the PoKosauridae and Ophiacodon- . There are two or three The fifth tarsale is rarely unossified. The humerus has an ectepicondylar foramen only in the Edaphosauridae. the lacrimal seldom extends to the nares. and supratemporals are always smaller. i66. Skeleton oi Edaphosaurus (Theromorpha). the interparietals. There is greater diversity among the Theromorpha as thus dis- tinguished than among the Cotylosauria. and usually longer legs. The plate-like pelvis never has a large pubo-ischiatic or a true thyroid foramen.230 well.

b n 231 .

232 .



Permocarboniferous (Uppermost Carboniferous and Lower-

most Permian).

Suborder Pelycosauria

Family Sphenacodontidae

Carnivorous reptiles

from four to eight

feet in length,

with long, often very long, dor-

sal spines; three sacral vertebrae.




Dimetrodon Cope, Texas.

Clepsydrops Cope,




Matthew, Texas.

Bathygnathus Leidy, Prince Edward Island.

Suborder Edaphosauria
Subaquatic or

Family Edaphosauridae (Edaphosauria).

invertebrate feeding reptiles, from six to eight feet in length.

Spines of dorsal vertebrae very long, each with transverse processes.
Skull small, short, high, with

numerous palatal and coronoid conical

Edaphosaurus Cope, Texas, New Mexico. Naosaurus Cope, Texas, New Mexico, Ohio, Germany, Russia.

Suborder Poliosauria
Lizard-like, insectivorous, four or five
of vertebrae short;

Family Poliosauridae.
feet in length.

Teeth conical; spines

two sacral


Texas and







Poecilospondylus Case, Arrihasaurus Williston, Scoliomus Williston

and Case.

Family Ophiacodontidae. About

six feet in length, carnivorous.

Skull narrow; teeth slender and conical or flattened; temporal open-

ing small, an upper one also in Ophiacodon;

ribs holocephalous;

limbs short and stout; two sacral vertebrae. Texas and
saurus WiUiston.

New Mexico. Ophiacodon Marsh, Theropleura Cope, Diopeus Cope, Secodonto-


Suborder Caseasauria

Family Caseidae (Caseasauria)

Thickset, crawling and probably

burrowing, invertebrate-feeding reptiles about four feet long.


Fig. 169.

Skeleton of Theropleura (Theromorpha), from below.


sixth natural size.












broad, short, with large pineal opening, and palate and coronoids

covered with conical teeth. Three sacral vertebrae. Texas.

Casea Williston,


Trichasaurus Williston.

SUBORDER Uncertain
slender reptiles.

Family Paleohatteriidae.


seven presacral vertebrae; three sacrals.

Parasternal ribs present.

Intimate structure of skull unknown. Skeleton feebly
ably young animals, the metacoracoids not
notochordal, ribs holocephalous.




Lower Permian. Germany. Haptodus Gaudry,

? Callibrachion Boule, France.

Incertae Sedis. MycterosaurusW\\\\s,ton,GlaucosaurusW\\\i'&tOTi,
Tomicosaurus Case, Metamosaurus Cope, Emholophorus Cope, Texas.
Archaeoholis Cope, Illinois.

Aphelosaurus Gervais, Autun, France.

Gaudry (? Sphenacodontidae), Autun, France. Doubtfully members of the order: Ammosaurus Huene (Triassic).



Less primitive, more upright-walking
or less inclined in locomotion.
chordal, dorsal intercentra

the propodials more

Vertebrae amphicoelous, rarely noto-

unknown. Palate and limbs less primitive

pelvis with larger pubo-ischiatic vacuity or thyroid opening.


stated on a previous page, sharp distinctions between the


bers of this order and the preceding one cannot be made.
tive characters




both orders are largely included in the
differences presented


But the very great

by the

later, dif-

Triassic, forms, especially those included

under the Cynodontia,

ferences as great as those between

any other two orders

of reptiles,

render a division or divisions imperative, even though



so often the case in other groups of animal

in the structural differences

between members

of the

and vegetable Hfe, same group

being greater than those limiting the groups themselves.
division, it

seems to the writer,


be best made at the present

time between the Lower and Middle Permian types, that


upon the stages

of evolution chiefly.

Perhaps when more



of the various


and diverse forms included


both orders, a better

and more scientific division may be made on genealogical characters. But such are not available at present.

less so

characters, as a whole, of the Therapsida are primitive, but

than those of the Theromorpha, and they are increasingly inconstant. The vertebrae are known to be notochordal only in the

Dromasauria and Dinocephalia, and the intercentra are seldom
ever persistent throughout the column; there



be as



seven sacral vertebrae; the boundaries of the temporal opening are

constant; in a few words, no characters seem to be more primi-

tive than in the

Theromorpha. The



present, are

fused into a single bone, which

rarely the case in the


The supratemporals

are always, the postf rentals often, the quadrato-

jugals usually, absent.^


palate and teeth undergo

many changes;

the pterygoids are less free, palatal teeth are less constant.



seldom present and always small,



to divide the various

groups into orders seems not to solve

but rather to add to the


For that reason, perhaps

it is

better at present to consider the whole group as one order, as


has suggested, clearly differentiated from


others save the Theroits

morpha by

the skull and pectoral girdle, and to treat


under the chief

many, of the genera is any order of reptiles or other organisms until everything about them is fully known, a result greatly to be wished, but never within the limits of human endeavor. The classification adopted is that of Broom and Watson in numerous publications and in Uteris, with but
few modifications.

Of course the distribution of some, perhaps more or less provisional, as must be the case in

Suborder Dinocephalia
size of a

Powerful reptiles from the

boar to that of a rhinoceros.

Skull very massive, especially in the cranial region. Temporal opening bounded by the postorbital and squamosal, the jugal sometimes

intervening below.

Lacrimals and quadratojugals small, the interlarge.

and tabulars


dermal bones fused in midline.

Parietal opening large, opening in a protuberance or boss.


[The quadratojugal has recently been identified in anomodonts, gorgonopsians,

and cynodonts, by Watson.

— Ed.]


less flattened

more or

and denticulated along


border/ not more

than eighteen in either jaw, subisodont or with a large caniniform tooth; no teeth on palate. Pre vomers, palatines, and pterygoids united in midline, concealing the parasphenoid. Quadrate large.
Vertebrae deeply concave or notochordal. Atlanto-axis as in Dimetrodon (Theromorpha) four sacral vertebrae. Ribs dichocephalous,

probably no parasternals.

Shoulder girdle massive; procoracoid

barely entering glenoid fossa; a feeble cleithrum sometimes,


Fig. 170.

Skeleton of MoJfAo/)j (Dinocephalia). After Gregory. Skeleton in American Museum. size.

One twenty-second


always, present.







Pelvis with small pubo-ischiatic vacuity.


entepicondylar fora-


Legs stout; epipodials and

digits short; phalangeal


unknown, probably

Middle and Upper Permian. Delphinognathus Seeley, Lamiasaurus'^ Watson, Moschognathus Broom, Mormosaurus Watson, Moschops Broom, Moschosaurus Haughton,

Family Tapinocephalidae.

Phocosaurus Seeley, PnigaUon Watson, Struthiocephalus Haughton,

[This statement refers only to the cheek teeth; the premaxillary teeth and the first

three or four in the dentary have a long conical crown, greatly expanded posteriorly
at the base,

and long


[Cranium, Fig. 170.

— Ed.] — Ed.]

Galeops Broom. parasternals present. Rhopalodon. Edentulous. Suborder Anomodontia mouse to that of a tapir. Zool. Upper Permian. Galesphyrus Broom. tarsus with 2. pubic large. quadratojugals obsolete or absent. Eichwald. About the size of Skull short. Family Galeopidae. London). quadratojugals small or obsolete. squamosal. the face short. or absent. and probably the terparietal.subisodont. lacrimals small. Middle Permian. Titanosiichus Owen Suborder dromasauria a rat. 3. 3. Middle Permian. parietal foramen large. South Africa. Family Rhopalodontidae. in front of. in 1923 (Proc. teeth isodont. two or three foramen presacrals. probably twenty-eight no acromion and no cleithrum. Eccasaurus Broom. South Africa. Large temporal opening bounded by postorbital. "Lamiasaurus" [snout]. Ural Mts. Deuterosaurus. phalangeal formula Family Galechiridae. pelvis plate-like. — Ed. present. South Africa. vertebrae notochordal. possibly the preparietal. or without a fifth tarsale. South Africa. vegetable or inver- From the size of a tebrate feeders. below. 3. and jugal. 3. quadrates and squamosals large. Taurops Broom. Galepus Broom. and jugal.i Upper Permian. Family Deuterosauridae. in- squamosal. . 2 [See page 243. orbits large. Family Macroscelesauridae. Upper Permian. C. Scapanodon Broom. A single row of subisodont teeth. intercentra unknown.THE SUBCLASS SYNAPSIDA 239 Tapinocephalus Owen. sacrals. Eichwald.] — Ed.^ Preparietal usually present. Macroscelesaurus Haughton. Ural Mts. or sur- [A number of new genera of South African Titanosuchidae were described by Broom ^oc. Archaeosuchus Broom. Galechirus Broom.] . carpus primitive. Family TiTANOSUCHiDAE. Skull typically short and wide. B. lacrimals con- tinuous to septomaxilla temporal opening bounded by postorbital.

rarely primitive. South Africa and Russia. A No parasternals. and generally with a in number series. and in canine. Stapes large. Diaelurodon Broom. and small tabulars. . An sternum. so far as known. Gordonia Newton. Cistecephalus Owen. Upper Permian. no teeth on palatal bones. There is a small cleithrum in Cistecephalus. Taognathus Broom. Upper Triassic. but there is never any canine present. Lystrosaurus Cope^ Prolystrosaurus Haughton.. Myosaurus Haughton. Legs short and stout. four to A mandibular foramen. Carnivorous Therapsida with more or coronoid. and Sclerotic surangular large. angular. of small molars often irregularly arranged Molars are always present on the mandible if in the maxilla. Emydops Broom. Scotland. Family Dicynodontidae. South Africa. no intercentra back of seven sacrals. covered with horny beak. few or no teeth on palate cleithrum. The shoulder girdle has the coracoid and precoracoid well developed. more than one Dentary. A rudimentary false palate. Suborder Theriodontia less differentiated dentition. Vertebrae amphicoelous. South Africa. Maxilla usually with an enlarged. Prodicynodon Broom. Dicynodon Owen. ilium projecting in ossified front of acetabulum. including at least one pair of upper caniniform teeth.3. Cryptocynodon Seeley.3. Wyoming. Dicynodon Owen. Prevomers fused. bones. Endothiodon Owen. Brachyhrachium Williston. mula 2. Dicynodon Owen.3. Chelyrhynchus Haughton.. permanently growing is absent in the females of some genera. parietal foramen. Pristerodon Huxley. Placerias Lucas.3. no coronoid. and possibly present known in Dicynodon and most other genera. Lower and Middle Triassic. Middle Permian. plates in orbits. and a distinct but short acromion. which. Emyduranus Broom. Kannemeyeria Seeley. Emydorhynchus Broom. hands and feet short. Occipital condyle tripartite. Diictodon Broom. atlas. Phalangeal forthyroid foramen in pelvis. an entepicondylar foramen. Tropidostoma Seeley. Eocy clops Broom. No Manus and pes. South Africa. a prominent Vertebrae never notochordal.240 THE OSTEOLOGY OF THE REPTILES An interparietal rounding. Geikia Newton. however. life Premaxillae fused and always toothless. D.

241 .

5. Aelurosuchus Broom. single occipital condyle. Tribe Gorgonopsia Prefrontals and large postf rentals contiguous over orbit. Parietal region wide. 1 [Made "the type of a new suborder. Cynodraco Owen. coracoids relaing tively small. Upper Triassic. Scylacognathus Broom. A single vomer (? fused prevomers). South Africa. Burnetiamorpha. 1923. A dis- tinct preparietal in front of small parietal foramen. a median. Scylacops Broom. 3. Temporal open- bounded above by united postorbital and squamosal. Phalangeal formula primitive 2. no clei thrum. 2. Galesuchus Haughton. Sesamodon Broom. Middle and Upper Permian. Family Gorgonopsidae. large posterior palatine vacuities. rhinus Broom. Inostrancevia Amalitsky. Melinodon Broom. No acromion on scapula. Lower Africa. Theriodesmus Seeley. Ictido- Family Burnetidae. Gorgonops Owen. 4.242 THE OSTEOLOGY OF THE REPTILES 1. Asthenognathus Broom. an ectopterygoid. strong incisors and grinding molars. A group intermediate. according to Broom. Tigrisuchus Owen. Arctognathus Broom. Aloposaurus Broom. 3- a large proatlas. No acromion on scapula. \ Ed. frontals. Arctops Watson. quadrate foramen present or absent. parietal squamosal small. Arctosuchus Broom. below by squamosal and jugal or squamosal only.^ South Tribe Bauriasauria A well-formed secondary palate. Russia. Family Ictidorhinidae. Bauria Broom. Burnetia Broom. South Africa.1 . South Africa. no postlarge. Middle and Upper Permian. Microgomphodon Seeley. Scymnognathus Broom. Scymnosaurus Broom. Cyniscodon Broom. Cerdognathus Broom. Aelurosaurus Owen. by Broom. unpaired vomer. Triassic. Ididomorphus Broom. Gorgonognathus Haughton. between the Thero- cephalia and Anomodontia. No secondary palate. the pterygoids extend to quadrates.

Middle and Upper Permian.^ No quadratojugals-. Alopecopsis Broom. Family Alopecopsidae. the postorbital and squamosal . Lycosaurus Owen. Scylacoides Broom. Middle and Upper Permian. Alopecorhinus Broom. Simorhinella Broom. one or two large upper caniniform teeth. 243 Tribe Therocephalia Temporal opening large. Cerdodon Broom. Middle and Upper Permian. Middle and Upper Permian. Scylacosaurus Broom. Scaloposaurus Owen. interpterygoidal opening. long dentary. Family Ictidosuchidae. Icticephalus • Broom. 1 [In typical Therocephalia. Alopecodon Broom. Troclwsuchus Broom. Scymnopsis Broom. South Africa. Family doubtful. Middle and Upper Permian. palate usually narrow. and Postcranial skeleton large coronoid. posterior elements not reduced. Eriphosto7na Broom. no preparietal. Family Lycosuchidae. bounded below by squamosal and jugal. parietal region Mandible with loose symphysis. Alopecognathus Broom. South Africa. a parietal foramen. Family Scylacosauridae.] by Broom. South Africa. above by the parietal or the connected postorbital and squamosal. largely unknown. prevomers separated or fused (Scaloposaurus) an . or a rudimentary one (? Scaloposauriis) . Pristerognathus Seeley. Akidnognathus Haughton. Lycosuchus Broom. Scymnorhinus Broom. no secondary palate. South Africa. as described do not connect with each ^ other. South Africa.] — Ed. and five to nine smaller ones posteriorly. Family Scaloposauridae. Scylacorhinus Broom. South Africa. PardosucJms Broom. four or five in pre- maxilla. with few or no teeth. Ictidosaiirus Broom. quadrates small. — Ed. Ididosuchus Broom. Teeth conical. [See page 239.THE SUBCLASS SYNAPSIDA 3. Family Whaitsidae. GlanosiicJms Broom. Whaitsia Haughton. large posterior palatine vacuities. above. postfrontals small or absent. Arnognathus Broom. Lycorhinus Broom. Middle and Upper Permian. South Africa. Ididognathus Broom. Hyaenasuchus Broom. squa- mosals large.

the pterygoids do not reach the quadrate. probably a small ectopterygoid vomer .244 THE OSTEOLOGY OF THE REPTILES 4. Jj] posterior mandibular bones less re- duced. 3. saurus Owen. pha2. A clei- small acromion on scapula. GaleMiddle Triassic. tabular large . Family Nythosauridae. Dentition composed of from three to five incisors. Ictidopsis Broom. reduced posterior mandibular bones. Platycraniellus v. quadrate small. Tribe Cynodontia Especially characterized by a heter- odont dentition. South Africa. Nythosaurus Owen. dorsal intercentra. no langeal formula thrum. Twenty-eight pre- sacrals. A thyroid foramen in pelvis. large. 3. 3. Temporal opening bounded by parietal and postorbital above. a secondary palate. rarely thirteen. scapula with reflected anterior border. . . four sacrals. but no preparietal bone . stapes long. a small parietal foramen. so far as known. unpaired. stout or slender. a canine. Hoepen. Fifth carpale unossified. frontals small. parietals narrow. molars. usually by squamosal and postorbital only below. molars less cuspidate. secodont or gomphognath or cuspidate. ex- cluded from orbital margin by the union of the prefrontal and postorbital postfrontals absent. Feet imperfectly known. Septomaxillae on face. and two occipital condyles. Coronoid large. the digits Vertebrae amphicoelous no short. 3. and seven to nine.

Theriodontia (?) Incertae Sedis bolodon Seeley. [posterior] mandibular bones more reduced. ? Dromotherium Emmons. ? Upper Triassic Cynochampsa Owen. Triassic. South Africa. 5. Probably each a distinct family. South Africa. Upper Triassic. South Upper Triassic. Diademodon Seeley. Pachygenelus Watson. South Family Diademodontidae. Africa. Middle Triassic. Lycochampsa Broom. — too is imper- the type of — R. TriKaroomys Broom. South Africa. Upper Africa. [May be either primitive mammals or cynodonts fectly known to enable one to decide.] . South Africa. Septomaxillae within nares. Broom. molars cuspidate. Trirachodon Seeley. Gomphognathus Seeley.THE SUBCLASS SYNAPSIDA Family CYNOSUcmDAE. Cynognathus Seeley. North Carolina. Family Cynognathidae. Tritheledon Broom. Lower Jurassic. ? 245 Cynosuchus Owen.

A single. would trace from the Cotylosauria. Pelvis with large pubo-ischiatic opening. general characters of the skeleton are of more or less modified by aquatic adaptations. The boundaries the temporal region seem to be those of the upper opening of the diapsid reptiles. and there are many who is believe that it really is the upper one. however. Parasternals stout. even in the Nothosauria. by the loss of the lower arch. dermosupraoccipitals. large temporal opening. chief groups. especially the Cynodontia. . or quadratojugals. A SINGLE. No sternum. Quadrate A parietal foramen. and these reptiles are confidently believed to have descended from theromor- phous reptiles is with a typical lower opening. large coracoid on each side. or secondarily a thyroid foramen. we must await the discovery of more terrestrial ancestors in the early Trias. the The order is clearly divisible into two Nothosauria and the Plesiosauria. There is still much doubt The as to the derivation and genealogical relationships of this order of reptiles.CHAPTER X THE SUBCLASS SYNAPTOSAURIA 6. Cer- vical ribs attached exclusively to the centrum. below by the postorbital and squamosal. but in the present uncertainty they may be left in an independent their Whatever has been their origin. ORDER SAUROPTERYGIA No fixed. bounded above by the parietal. the dorsal ribs ex- clusively to the arch by a single head. The author group. The modificalife tions of structure in adaptation to aquatic are very pronounced. is bounded quite like that of some members of the Therapsida. it is seen. that the Sauropterygia are related to the anomodont-like their descent directly Some. Girdles stout. Vertebrae platycoelous. the tail never long. others from the Diapsida. The more general opinion reptiles. nearest related to the Progano- The opening. chiefly because of the structure of the temporal region. believes that the first of these views is the correct one. and that the order sauria. Neck elongated. tabulars.

Microleptosaurus Scuphos. like the plesiosaurs. Lamprosaurus Meyer. are very stout. or with the loss of one phalanx in the fourth finger. of exclusively Triassic age. Dactylosaurus Giirich. upon minor characters For the present they may all be placed in a single family. the Nothosauridae. and subsisted chiefly brates. though less broad than that of the and it is not probable that they were rapid swimmers. Doliovertehra Huene. Lariosaurus Curioni. and also in the unusually stout clavicular girdle of both. as do the crocodiles. sacral. the orbits situated far forward. Anarosaurus Dames. vertebrae. Several famihes have been proposed. From twenty to twenty-five cervical. based of the skull chiefly. Neusticosaurus Seeley. Proneusticosaurus Volz. Skull depressed.THE SUBCLASS SYNAPTOSAURIA A. Nares about mid- way between the orbits and extremity. Family Nothosauridae. Clavicles stout. The parasternals. looking upward. like those of the Plesiosauria. two to five and a moderately long tail. twenty-five to thirty dorsal. Digits probably webbed in life. Nothosaurus Miinster. Phalangeal formula primitive. Upper and Middle Trias. for the capture of upon fishes and inverte- were well pecuHar parallel adaptation to that of the contemporary aquatic Labyrinthodontia is seen in the forward position of the eyes fitted. Cymatosauriis Fritsch. except the large internal nares. Lacrimals possibly absent. but not exclusively so with but minor aquatic adaptations. curved teeth A in the flat skull. They doubtless lived in the shallow waters. line. . slender. which their slender. com- ing frequently to land. more or less elongate. Pistosaurus Meyer. Palate without openings. apparently also an aquatic adaptation. the vomers and pterygoids meeting in the middle line throughout. the feet still retaining terrestrial characters. Parthano- saurus Scuphos. The elongated coracoids meet in the middle Epipodials much shorter than propodials. 247 Suborder Nothosauria reptiles Crawling or swimming from three to seven feet in length. Simosaurus Meyer. The body was never plesiosaurs. The Nothosauria were all aquatic in habit. the interclavicle vestigial.

five pectoral and about twenty dorsal vertebrae. other openings variable on the palate. it is seen. clavicles An with extensive and long-lived group of purely marine reptiles. Their phylogenetic relationships with the Nothocould have been actually ancestral to sauria are incontestable. Jurassic. though not nearly so broad as represented in most modern restorations. in the Stuttgart museum. of which a figure copied from a photograph is reproduced here. contiguous in midline.ES B. sometimes vestigial. but many minor modifications. reptiles from eight to about with paddle-like. The neck was extremely variable in length. no interclavicular foramen. long. much longer than broad. Europe. but not flat.248 THE OSTEOLOGY OF THE REPTn. Coracoids very large. Thaumatosaurus Meyer. and interclavicle small. hyperphalangic limbs. epipodials dials. with from thirteen to seventy-six cervical vertebrae. Premaxillae continuous to parietals in middle. Family Pliosauridae. situated in front of the external. Family Plesiosauridae. composed of about Cervical ribs double-headed. Skull moderately five to long. widely distributed over the earth. SUBORDER PLESIOSAURIA fifty feet in length. The body. Skull moderately broad to very Nares small. terpterygoidal openings divided A pair of posterior inin the by the parasphenoid always present. Orbits with sclerotic plates. situated remote from the extremity and near the orbits. above with a well-developed sacrum of three or four vertebrae. Scapulae closely approximate in midline. Skull nineteen vertebrae. neck short. No distinct nasals. no accessory epipo- Coracoids contiguous throughout. The squamosals meet middle line posteriorly. though the closed palate of the latter indicates that no known form them. Marine slender. is broadly oval. articulating below with ischium only. The body was broad. Scapulae not contiguous in the middle. the cervical ribs double- headed. The most perfect specimen known and the author has seen most of them in the collections of the world is that of Thaumatosaurus victor. protected below by the extraordinary — — developments of the pectoral and pelvic girdles and intervening parasternal ribs. Internal nares small. Ilium rod-like. coracoids con- . From thirty- [seventy-six] cervical vertebrae. Plesiosaurus Conybeare. as a whole clearly defined.

Fig. 173. 249 . One twentieth natural size. After Fraas. Skeleton of Thaumatosaurus (Plesiosauria).

174. lating with parietals. Cry ptocleidus Seeley. with from more than fifty to seventy-six vertebrae. Ischia short.250 THE OSTEOLOGY OF THE REPTILES Two or three epipodials. Fig. Elasmosaurus Cope. and the coracoids are contiguous throughout. no interclavicular foramen. is shorter. Trinacromerum Cragin. four epipodials. Three or Upper Cretaceous. broader. process separates the scapulae in the midline an interclavicular fora- men. all short. Family Cryptocleididae. Europe. Peloneustes Lydekker. a Cretaceous plesiosaur. Ischia long. Picrocleidus Andrews. Polycotylus Cope. North America. Europe. Very much Hke the but the neck epipodials. North America. with from thirty-two to forty-four vertebrae. Pliosaiirus Owen. Skull short. Jurassic. Large or very Jurassic. neck very long. as broad as long or large. with from twentythree to twenty-six vertebrae. Premaxillae articuNeck not longer than head. Family Polycotylidae. as mounted in the University of Kansas Museum. Family Elasmosauridae. Sthenarosaurus Watson. Coracoids broadly separated on their posterior half. Microcleidus Watson. The precoracoidal . Leurospondylus Brown. all From two to four short. Two epi- Upper Cretaceous. short. Muraenosaurus Seeley. The scapulae meet in midline. Ogmodeirus Williston and Moodie. . Ischia elongate. tiguous throughout. Skeleton of Trinacromerum osborni. ? Piratosaurus Leidy. ribs single-headed. Skull very slender. Tricleidus Andrews. coracoids contiguous throughout. Cervical ribs single-headed. ribs single-headed. podials only. Head short. following family.

because of our ignorance of the skeleton. with isolated ones upon the skull. Jaws and palatines with few. and squamosal. neck very short. But their presence has been denied. Orophosaurus Cope. Lower Cretaceous. shell-eating reptiles has long been a problem. Spondylosaiirus Fischer. Lioplenrodon Sauvage. Paddles imperfectly known. Europe. Europe. Upper Cretaceous. . Mauisaurus Hector. " Plesiosaurus^' Conybeare. as examples. 7. with but Cervical ribs singleheaded. Jurassic. Pterygoids not reaching to vomers. as believed by Huene. . North America. Uronautes Cope. sedis ^'Plesiosaurus" Conybeare. Embaphias Cope. On the other hand. postorbital. very large. North America.OligosimusL. hypantrum. Genera incertae Triassic. Taphrosaurus Cope. This singular group of littoral. however. North America. ^^ Plesiosaurus^' Conybeare^ North America. shorter than skull. Remainder of skeleton unknown. the supratemporal and interparietal are really present. Pantosauriis Marsh. " Muraenosaurus^' Seeley. among the Therapsida would be im- Placochelys has a carapace of bony plates. Their presence or absence. or no more importance than Squamata. include Some would If others would give to them among them the same rank among the Therapsida. Simolestes Andrews. both above and to of below. New Zealand. all of which seem is be wanting in Placodus. 251 Skull long. Polyptychodon Owen. Megalneusaurus Knight. their location proper. ORDER PLACODONTIA Temporal opening bounded by parietal. Colymhosaurus Seeley. thirteen vertebrae.eidy Brimosaurus Leidy. Upper Cretaceous. Eretmosaurus Seeley. postfrontal. A parietal opening. Piptomerus Cope.THE SUBCLASS SYNAPTOSAURIA Family Brachaucheniidae. Brachaiicheniiis Williston. Cimoliosaurus Leidy . in the Dinosauria. with hyposphene. Vertebrae amphicoelous. if there should prove to be but a single coracoid on each side in the pectoral girdle. Europe. Ribs double-headed. flat crushing teeth. they would certainly find no place among the Sauropterygia. Ischyrodon Meyer. the Sauropterygia as a separate suborder.

The temporal vacuity is bounded as in the plesiosaurs. Family Placodontidae. Upper Triassic. and with a heavy have all is shell-eating reptiles. Europe. . The placodonts were ten feet in length. perhaps an adaptation for grubbing in the mud after in- vertebrates. perhaps eight or undoubtedly slow in movement. Possibly there was a moderate adaptation in Placodus for life in shallow water. skull. the nares situated rather far back. Cyamodus Meyer. though there skull that Uttle in the structure of the would them to an ordinal rank. the is Until more known entitle group may remain in an independent position. the As might be suspected brae is number of presacral verte- reduced. as reptiles of considerable size. and also in some theriodonts. Placochelys Jaekel. of the skeleton. The maxillae are large.252 THE OSTEOLOGY OF THE REPTILES in such forms. Placodus Agassiz {Ano- mosaurus Huene). shell-eating animals with crushing teeth occur in various orders.

Until the skull the Sauropterygia. though highly speciahzed coid. 3. probably side. the earhest retain many primitive characters. with a single. 6. by others among tail and elongated. articulating with centra. phalangeal formula pes (in Mesosaurus and Noteosaurus at reptiles. eleven or twelve cervicals. 1 [Much further evidence for this view Ichthyosaurier des Lias und ihre is given by von Huene in his memoir Die Ziisammenhdnge. Primitive. a single corapelvis with small pubo-ischiatic coid. long and slender. 1922. Numerous parasternal ribs. Free ribs on all presacrals except atlas. ORDER PROGANOSAURIA tail. upper temporal opening. Because of the articulation of the single-headed and the probable possession of but a single.CHAPTER XI THE SUBCLASS PARAPSIDA 8. carpus and tarsus primitive. however. doubt remains as to their relationships with other reptiles. vacuity. the nostril near orbits. upper temporal opening on each Face long and Teeth numerous. unknown. in the scapular girdle with its single cora- known. Propodials long. probably also on other palatal bones. Ed. the first least) 2. Aside from the Dolichosauria and certain dinosaurs they are the only known aquatic reptiles with both neck is better known. single-headed. By some they have been placed with the double-arched reptiles. the premaxillae elongated. intercentra slender. Scapula fan-shaped. 4to. Vertebrae deeply amphicoelous. dorsal ribs stout. 4.^ ribs especially. Humerus with entepicondylar foramen. body. their natural association seems to be near the ichthyosaurs and lizards. clavicular girdle primitive. 5. of epipodials short. Berlin. eighteen to twenty-two dorsals. two or Structure of skull imperfectly known.] — . These small known in geological history with marked aquatic adaptations. aquatic reptiles with long neck. the fifth toe elongate. two sacrals and sixty or more caudals. small teeth on vomers. and three feet in length.

254 .

posterior nares. The posture of the hind leg is slightly modified. short neck. 176. 9. short pro- . Mesosaurus Ger- Noteosaurus Broom. 255 Lower Permian. Mesosaurus Gervais. After McGregor. sclerotic plates. South Africa. vais {Ditrichosaiiriis Gurich). no sacrum. ? Slereosternum Cope {Notosaurus Marsh).THE SUBCLASS PARAPSIDA Family Mesosauridae. Fig. ORDER ICHTHYOSAURIA all Marine reptiles with aquatic adaptations of the tail-propelling type: elongated face. flattened or dilated tail. Brazil. Restoration of Mesosaurus. elongated body. long.

tetra. Middle and Upper Switzerland. postf rental. Mixosaurus Baur. deeply amphicoelous. Delphinosaurus Merriam. free paroccip- large stapes. Nevada. and often hyperdactyly. The large upper temporal vacuity is bounded by parietal. Spitzbergen. No lateral open- Vertebrae short. labyrinthine in structure. Teeth more or less anisodont. Both the Mixosauridae and Shastosauridae. California. The ichthyosaurs were adapted to aquatic plesiosaurs. epipodials shorter. saurus Wiman. Phalaradon Merriam. pelvis more reduced. dichocephalous. Middle or Upper Triassic. Chevrons Y-shaped. are more primitive. No sternum. hyperphalangy. Feet dactylate. Family Shastosaueidae. Triassic. without persistent dorsal Scapulae small. a single coracoid. no ectopterygoids or dermosupraoccipitals. feet pentedactylate. slightly decurved. A parietal foramen. but numerous parasternals. Pelvis more lar or less plate-like with small pubo-ischiatic vacuity. Body more elongate. Toretocnemus Pesso- Merriam. Cymhospondylus Leidy.256 THE OSTEOLOGY OF THE REPTILES podial and epipodial bones. none on palatal bones. Germany. clavicles and inter- intercentra. and especially the Ophthalmosauridae. Cervical ribs Tail distinctly expanded and decurved distally. life exclusively marine reptiles. which Merriam gives only sub-family values under the Mixosauridae. Premaxillae long. inserted in sockets. Face less elongate. Spitzbergen. maxillae short. hind limbs . Epipodials relatively long. Epipodials relatively long. Shaslosaurus Merriam. more perfectly it than any other known ones unless be the They varied from about two to about thirty feet in length. chevrons separate or fused. and tabular (supratemporal) . ing. cephalous. Merriamia Boulenger. Teeth inserted in sockets or grooves. Family Ichthyosauridae.or tri- Chevrons Y-shaped. with less perfect aquatic adaptations than the later forms of the Ichthyosauridae. Family Mixosauridae. of Prearticu- bone mandible distinct. itals. Fewer presacral vertebrae. Cervical ribs for the most part holo- Tail with a preterminal dilatation. tail with a broad terminal fin. clavicle present. dorsal ribs dichocephalous.

257 .

? ORDER 0MPHAL0SAURIA2 Marine reptiles Family Omphalos auridae.] the evolution of which he traces from the Triassic to the Upper Cretaceous. their broad. Upper Upper Cretaceous. Africa. Europe. and in the more discoidal form of the phalanges. shell- Mandibles short. . Upper Jurassic. the presence of three epipo- bones in the front paddles. the ordinal rank and relationships provisional or conjectural. however. it is A widely distributed genus as ordinarily accepted. Asia. Europe and North America. the largest about fifteen milUmeters in diameter.] .258 THE OSTEOLOGY OF THE REPTILES more reduced. The incompletely known remains of these reptiles. the fusion of the ischium and ilium. are very suggestive of a new type of shell-eating aquatic reptiles. Ophthalmosaurus Seeley {? Baptanodon Marsh). crushing skull. It pre- South and ? North America. Cretaceous (Upper Greensand) ? Ophthalmosaurus Seeley. the more reduced hind paddle. the dentaries united in a strong symphysis. — Ed. Nopcsa) class the Omphalosauria with the Ichthyo- — sauria.[Recent authors (von Huene. tially From the same horizon in Spitzbergen similar teeth have been to pertain to the described by Wiman. division.^ numerous minor modifications that might justify its Family Ophthalmosauridae. Triassic to in usually grooves. . dial Differs from the more typical Ich- thyosauria in the more reduced teeth. Teeth inserted Face longer. described by Merriam. In them- but their associations fully and their age make it not at all improbable that when known essen- they will justify the rank provisionally given to them. superior surface beset with several rows of low-crowned. but until more is known they are merely suggestive. convex. in the apparent entire absence of chevrons. which seem same kind of 1 [Von Huene (1922) divides the old genus Ichthyosaurus into several phyletic lines. with a short. frequently hyperdactylate. Ichthyosaurus Koenig {Proteosaurus Howe)." Skeleton otherwise unknown. sents. button-like crushing teeth. selves the characters are not of ordinal rank. Vertebrae amphicoelous. Ed. New Zealand. Australia. "Palate plesiosaur-like.

paired. bearing possibly a like relation to the known Ichthyosauria Triassic. and clavicles. twenty two sacral. Perhaps when finally known they will be found to be incoherent. Palatal bones with teeth. here identified as the squamosal. an interclavicle. so far as known. 10. as here limited. feet in length. At least seven and a long. dorsal. with a single. is This order. slender. Cervical ribs. reptile lizards. that Glohidens does to the typical Pytho- nomorpha.THE SUBCLASS PARAPSIDA reptiles. or Quadrupedal. subaquatic reptiles one to six upper temporal opening between the parietal all and the temporal arch. upper temporal vacuity. arboreal. brae. the quadrate fixed. Pelvis primitive. a provisional one. slender Vertebrae amphicoelous with persistent intercentra. For the present they may be defined as families. at least. hol- Family Araeoscelidae. some of them imperfectly known. Lac- rimal vestigial or absent. The dermosupraoccipital is apparently large. leaping or . arboreal or leaping. Coracoid and scapula closely fused. Middle Omphalosaurus Merriam. single-headed. All cranial bones cervical vertetail. which cannot be placed in any other known order. bounded as in the It and a fixed quadrate. including several reptiles. Very low-boned legs reptiles of less tail. was a very slender. is Araeoscelis. single-headed. with long lateral and long The broad temporal region is formed ap- parently of a single bone. the dorsal more or less dichocephalous. 259 Somewhat doubtfully associated with those remains are others of ichthyosaur-Hke bones that the describer provisionally associated with the Ichthyosauria. Phalangeal formula primitive. from which they are distinguished by the absence of a lateral temporal opening. Humerus with both entepicondylar and ectepicondylar foramina. A parietal foramen. the type of the family. articulating with centra — a single Ribs in part or coracoid. Nevada. Most of them have hitherto been classified with the Rhynchocephalia. the earHest definitely known with a single. Calcaneum produced. Spitzbergen. ORDER PROTOROSAURIA terrestrial. than eighteen inches in length. Pessopteryx Wiman. the quadratojugal absent.

Skeleton oi Araeoscelis (Protorosauria). 260 .{\on oi Araeoscelis. Fig.Fig. Re&tora. 180. About one fourth natural size. 179.

arboricolous, insectivorous, lizard-like reptile from the


Lower Permian of Texas. Of Kadaliosaurus, unfortunately, the skull is unknown. Its slender bones were less hollow, and it has also numerous
parasternal ribs,

unknown in Araeoscelis. Lower Permian. Araeoscelis Williston, Texas. Credner, Germany.
Family Protorosauridae. Elongate


with long neck and

hind legs and hollow bones, from three to five feet in length. Skull
imperfectly known, probably with an upper temporal opening only.
Sclerotic plates in orbits.

Prevomers, palatines, and pterygoid with

small teeth.

Vertebrae amphicoelous, with persistent intercentra.

a long

cervicals, sixteen to eighteen dorsals,


single coracoid.


two or three sacrals, and more or less plate-like, with

Fig. 181.

Skeleton oi Protorosanrus (Protorosauria), modified from Seeley. About one tenth natural size.

probably a small pubo-ischiatic vacuity. Ribs single-headed, articulating with centrum, those of the cervical region very slender.


podials about as long as propodials, the hind legs


longer than

the front.

Humeri with ectepicondylar


foramen; nine or ten

carpals, seven tarsals

phalangeal formula primitive, the digits long.
ribs. fossil reptiles,

Numerous abdominal

Although the first-described

the protorosaurs are


in the details of their structure, especially of

the skull, pectoral, and pelvic girdles.

In the elongation of the neck

and the slender legs Protorosaurus very


resembles Araeoscelis,

and doubtless had similar habits, whether or not the structure of the skull was the same. The numerous known specimens of Protorosaurus differ so much from each other that it is not at all improbable that they represent different genera.


is still


of it are the trunk

more problematical, inasmuch as all that is and limbs. The limbs resemble those of



Protorosaurus in



ribs are described

and by Thevenin as




articulating intercentrally.

Lower Permian. ?Aphelosaurus Gervais, France. Upper Permian. Protorosaurus v. Meyer, Germany. The nares were described by Seeley as immediately
the orbits

— an

in front of




be a small antorbital foramen,


it is

Slender, terrestrial or subaquatic rep-

Family Saphaeosauridae.

about two

feet in length.

Skull with a single temporal opening,

the quadrate fixed and the lateral temporal region moderately broad.


postf rontals postorbitals large.


parietal foramen.


and dentaries edentulous, with cutting edges. Vertebrae procoelous without intercentra; twenty-three presacrals, two sacrals, and fifty or more caudals. Caudal vertebrae with splitting point (?). Ribs
single-headed, articulating with anterior part of centrum.


with two median emarginations. Interclavicle T-shaped, the clavicles

Parasternals numerous, composed of a median unpaired
lateral splint


and a

on each


Pubes and

ischia broadly



pubo-ischiatic opening, the ischia with a stout posterior



ectepicondylar foramen in humerus.

Manus and


pentedactylate, with primitive phalangeal formula.

Saphaeosaurus, usually called Sauranodon, has long been classed

The by both von Meyer and Lortet, has but a single temporal opening on each side, bounded externally by the postorbital and squamosal (tabular?) There is no lower temporal opening. The
as a representative of a distinct family of the Rhynchocephalia.
skull, as described

structure of the temporal region as described





probability the tabular, squamosal, and quadratojugal are



all its essential


it is

a Lacertilian with a primitively

fixed quadrate.


vertebrae, as figured

and described by Lortet,
of such in geological

are procoelous, perhaps the

known evidence




Meyer {Sauranodon Jourdan),

Family Pleurosauridae. Very

slender, snake-like, aquatic reptail,

with short neck, long body, very long flattened

and small

pentedactylate legs;

attaining a length of nearly five feet.





Skeleton of Saphaeosaurus {ProtorosAunz). One fourth natural size.

After Lortet.




elongate, pointed, the nares remote from end.
parietal foramen.




single temporal opening

bounded within by

the parietal, without by the postorbital and





Teeth pointed and recurved.



unknown. Five cervicals, forty or forty-one dorsals, two saand more than seventy caudals. Vertebrae amphicoelous, cerSquamata. Numerous
slender, parasternal ribs.

vical intercentra hypapophysial. Ribs single-headed, articulating as in the

Pleurosaurus, the only certainly

known genus

of the family,


long supposed to be a
also long

member of

the Rhynchocephalia, though

been known to have but a single upper temporal opening.
life is

remarkable adaptation to aquatic


in the elongated

head, posterior nares, short neck, very slender trunk, very small

and enormously elongated tail, with its long chevrons and spines, which in life was surmounted by a thin crest of scales. Acrosaurus is probably only the young of Pleurosaurus, as the author convinced himself by examination of specimens in the Munich museum. In consequence, the ordinal name once proposed for these
reptiles, Acrosauria, is inappropriate.


structure of the temporal



needs confirmation.

If there is
it is

but a single bone bounding

the temporal opening posteriorly,


probability the real






Meyer {Anguisaurus Miin?

Saurophidium Jourdan), Germany, France.



Meyer. Germany.



With a

single temporal vacuity

tabular, squamosal,

on each side, bounded by parietal, and postorbital, secondarily sometimes roofed

over or the arcade obsolete.


lower temporal opening or bar.

Quadrate movably articulated,

streptostylic, secondarily



supratemporals, dermosupraoccipitals, or quadratojugals.
articulate in front with the palatines, never with the

The pterygoids

prevomers. Paroccipitals fused with exoccipitals.

Interorbital septo

tum not


Teeth acrodont or pleurodont, often attached
Prearticular fused with articular.

palatine and pterygoid.


single-headed, articulating with centrum.



Suborder Lacertilia


Subvolant, arboreal, terrestrial, burrowing, subaquatic, or marine

from a few inches
or limbless;

about forty feet in length quadrupedal,


herbivorous, insectivorous, or carnivorous.

Brain-case in front of prootics more or less membranous. Lacrimals
small or vestigial.
usually united
Posterior arcade sometimes absent.



Vertebrae procoelous, except in the Geck-

onidae and Uroplatidae not more than two sacral vertebrae. Clavicles and interclavicle rarely absent. No entepicondylar, but usually an ectepicondylar foramen in humerus. This group is often given an ordinal rank, equivalent to the Ophidia or even to the Pythonomorpha, but the ultimate distinctions be-

tween them are almost

trivial, as will

be seen, and in



burrowing lizards the skull structure mimics that

of the snakes.

More than

eighteen hundred species are known, distributed widely

throughout the world, usually classed in about twenty families and

numerous genera.
of their


terrestrial habits,

but few remains

of lizards are found in the rocks, aside

from the more aquatic or

marine types.

Only about


genera of extinct forms have been

described and less than one hundred species, and the greater majority

most part fragmentary and incomplete, so much and provisional. Doubtless they have had a long and abundant geological history from very remote times, but of the true land lizards almost
of those are for the

so that their systematic positions are very often uncertain



known throughout

the Mesozoic.

But few

positive char-

acters are distinctive of the group, though


negative ones are.

The mandibles

are usually suturally united in the middle, but a few

forms have them ligamentously attached. The presence of legs


though at

least a vestige of the pectoral girdle remains.

The more

or less open brain-case in front

perhaps the most

diagnostic, only partially enclosed

optics ("alisphenoids," "postorbitals").

baenia even this character
tinct descending plate


more or less vestigial postHowever, in the Amphisdoubtful, and in the mosasaurs a disthe


from the parietals resembles that of the snakes,

see C. L.

[For a very comprehensive morphological and taxonomic revision of the Lacertilia, Camp, " Classiiication of the Lizards," Bulletin, Amer. Mus. Nat. Hist., 1923,

— Ed.]



but does not reach the basisphenoid. The jugals, squamosals, and tabulars may be more or less vestigial, and even the quadrate may be
secondarily fixed and immovable.

Tribe Kionocrania
Terrestrial, burrowing, subaquatic, or subvolant.


slender epi-

pterygoid articulates with parietal and pterygoid; no descending
plates of the parietals.

Palate with large openings, usually with teeth

on palatines or pterygoids or both. Feet when present usually pentedactyl, with the primitive phalangeal formula, the fifth metatarsal


or less hook-shaped proximally. Eight cervical vertebrae.

Family Geckonidae. Vertebrae amphicoelous,^ notochordal, with
persistent intercentra.

Quadrupedal. Jugal





Parietals paired.
of small lizards

Clavicles perforated near mesial end.

A family

widely scattered over the earth, compris-

ing nearly three hundred species and about fifty genera.



of interest because of the persistently primitive condition of the


early Mesozoic times, but no species are

They must have had a long independent history from known as fossils.

Family Euposauridae. Small

from two to four inches in

length, of doubtful position; referred to the Anguinidae

by Boulenger.


relatively large

and broad,

orbits very large, the temporal

openings said to be closed.

Structure poorly known, twenty-three



Euposaurus Lortet, France.

Family Agamidae.

A parietal

foramen. ^


Temporal and postorbital arches complete. dermal ossicles [on back]. Teeth acrodont.

This exclusively Old-World family includes about two hundred


some of them attaining a members are the Flying Dragons {Draco), small lizards with an extraordinary development of the ribs to support a parachute membrane. Chlamydosaurus, one
species of about thirty genera,

length of three feet. Perhaps the most noted

of the largest of the family,

has an extraordinary


about the neck

[Rarely procoelous. See G. K. Noble, 1921, Amer. Mus. Novitates, No.


[Except Liolepis.

— G. K. N.]

— Ed.]

Arboreal. much like a "Horned Toad" in appear- ance. ossicles beneath corneous Temporal opening roofed over by dermal bones. of the ways in which terrestrial reptiles Eocene. scales. body and skull more or No less parietal fo- covered by dermal An ossified. France. Spines of vertebrae sometimes elongate. reaching a length of six feet. 267 are subaquatic in Some The Moloch lizard. But one genus and two species of this family are known. Teeth pleurodont. surface of and frontals fused. habit. or Body covered with dermal parietal foramen. Iguanas. Miocene. Upper ossicles. France. terrestrial. — the Basilisc lizards. or sub- aquatic. Oligocene. Family Anguinidae. rhynchus. the Poisonous. the fa- . Herbivorous and insectivorous. perhaps one acquired water habits. . Amblyits a noteworthy herbivorous. Family mal Iguanidae. burrowing. Europe and America. Pleistocene. With well-developed. France [Agama]. North America. Iguanavus Marsh. ossifications. arch. terrestrial lizards with grooved. A postorbital but no temporal squamosal absent. subfrontal. slender. Diploglossus. A parietal foramen." The large Galapagos lizard. A This family. returning to the land for safety when pressed by enemies. Zygosphenes sometimes present. in distribution. including our lizards. No der- Temporal and orbital arches complete. rhinencephalic chamber. About largest three hundi'ed species and fifty genera are known of this family. Proiguana Filhol. limbs vestigial. has long dermal spines. Anguis. Chlamydosaurus Australia. Quadrupedal. almost exclusively and some is of American our most common etc. Family Helodermatidae." with wholly without them. pleurodont teeth. "Horned Toads. comprises about Most noteworthy are the "Glass Snakes" and the to vestigial limbs or common "Slow Worms.THE SUBCLASS PARAPSIDA supported by the elongated hyoid bones. prefrontal and postfronto-orbital in Parietals contact over orbits. pentadactyl hmbs. Teeth pleurodont. ramen. fifty species. aquatic form that seeks food in shallow water.

terrestrial. like the Cnemidophorus common throughout the United A . Lower Cretaceous (Neocomian). crushing organs. cosmopolitan in feet. terrestrial lizards. Glyptosaurus Marsh. They are thickset. K. Upper surface of skull with numerous dermal bones. Tejus. The teeth of Dracaena are large oval. Family Lacertidae. None is large and some are common throughout England. N. Family SciNCiDAE. Thinosaurus Marsh. family of American lizards including about one hundred species. North America. Asia. The large family of skinks comprises about four hundred li\dng species. North America. ' [A postorbital arch is present. mous "Gila Monsters" with a club-like tail. Cadurcosaurus Filhol. or subaquatic lizards at- taining a length of three feet.^ A No dermal ossicles. Oligocene.268 THE OSTEOLOGY OF THE REPTILES of Arizona. Helodermatoides Douglass. A parietal foramen. slow lizards feet. some. Zygosphenes sometimes present. terrestrial. Family Tejidae. The family of Lacertidae comprises about one hundred species restricted in distribution to Europe. Uppermost Cretaceous.] . or limbless. Temporal arch complete. Oligo- cene. Teeth pleurodont. or burrowing. reaching a length of about two of the suborder. ? Chamops Marsh. Quadrupedal. is the only reptile known to occur in Ireland. North America. Miocene. subaquatic. Temporal openings roofed over by dermal bones. France. one. bipedal. Some is attain a length of about two tail Trachysaurus of AustraUa scales of the peculiar in its stumpy and very large body. Ardeosaurus Meyer. No postorbito-squamosal arch. Pleurodont. its distribution. and Africa. — G. the arches complete. Lacerta. Temporal opening roofed over by the postfrontal extending back between parietal and squamosal. Arboreal. parietal foramen. Quadrupedal. Placosaurus Gervais. the only known poisonous members Eocene. Cyclodus has spherical crush- ing teeth. France. France. Lacerta vivipara. States. Eocene. Body also covered by dermal ossicles beneath the corneous scales.

Didosaunis Giinther. with about thirty living species. and Asiatic. Raven. Australia. Megalania Owen. Proganosaurus Portis. that it would seem very probable they is origin in early Cretaceous times. so like that of the following forms of the Dolichosauridae. Pliocene.THE SUBCLASS PARAPSIDA Oligocene. C. nasals. includes but one genus. Dracaenosaurus Gervais. about thirty feet (Megalania) Skull more or less elongate. This family. Slender aquatic two or three feet in length. vertebrae. none more than seven feet in length. the nostrils rather far back. [Megalania. the largest of all known ter- Unlike most lizards. twenty dorsal. of the Dutch East Indies. India [and Pleistocene of Australia]. France. with the primitive phalangeal formula. and Postorbital arch incomplete. Family Dolichosauridae. Varanus. present. Sacrals Family Varanidae. India. An imperfect joint between angular and splenial. cervical. long 1 neck of thirteen thir- teen feet. exclusively [Australian]. Varanus. they have a long protrusible Paleovaranus Filhol. Protrachysaurus France. broadly open. pentadactylate. reaches a length of H. in which they tail. India. of Terrestrial or subaquatic. seeking the water. Premaxillae. reaching a length . Pleistocene. Eocene. African. all had a common from the Pliocene of Megalania. Descending plates from the frontals enclose a rhinencephalic chamber. Epipterygoid and parietal foramen present. tongue. North America. Large palatal openings. with a relatively small skull. and especially the Aigialosauridae. Nine parietals unpaired. 269 Stefano. Pliocene.] lizards.^ Some are subaquatic in habit. Girdles complete. swim with freedom by aid Their structure of the long flattened is to escape their enemies. No dermal bones. Saniva Leidy. Tribe Platynota Terrestrial or subaquatic lizards from two or three feet to about Feet thirty in length. restrial lizards.] — . [Varanus komodoensis Owens.

Subaquatic lizards from three to six Skull large. England. has a short neck. Opetiosaurus Kornhuber. Pontosaurus Kramberger. have been thought by some to be ancestrally related to the snakes but this is very doubtful. with the same of vertebrae number were reptiles found in some mosasaurs. since their flattened tail shows a distinct adaptation to water life and it is improbable that the snakes ever passed through an aquatic stage in their evolution. slender cylindrical vertebrae. ent. terrestrial. Feet not hyperphalangic. size. they are the only known swimming reptiles with both neck and tail elongated. Family Aigialosauridae. Aigialosaunis Kramberger. tail. the body elongated. Dolichosaurus Owen. Aside from the Proganosauria. The dible in skull of the aigialosaurs is almost identical in structure with that of the mosasaurs.ES two vertebrae. The neck is shortened. . flattened. including the remarkable joint in the man- between the angular and splenial. Zygosphenes pres- Legs relatively small. Adriosaurns Seeley. Lower Cretaceous (Neocomion). body of twenty-one vertebrae. body. but not directly ancestral to any later forms. In probability the doli- chosaurs were a side branch from the common varanoid ancestral stock of the aigialosaurs and mosasaurs. The doHchosaurs. Pleurodont. feet in length. ? Mesoleptos CornaUa. however.270 THE OSTEOLOGY OF THE REPTn. Two sacrals. Acteosaurus Meyer. Pleurosaurus. and their ligamentous union front. and the small legs must have been all of peUing. frequenting the shallow waters for food. body of twenty-six or twenty-seven and a long flattened tail. Carsosaurus Kornhuber. no proan allied reptile of similar form. Europe (Dalmatia). The limbs. probably webbed. and but little other. Legs of nearly equal the propodials somewhat shortened. Because of the snake-like sinuosity of the neck. the front ones smaller than the hind. Doubtless the were amphibious in habit. Just what habits were subserved by this structure is a problem. with their greatly elongated neck and body. Neck of seven vertebrae. tail long. with only slight aquatic adaptations. use in the water. mosasauroid. Upper Cretaceous. Europe (Dalmatia). Lower Cretaceous (Neocomion). sacrals.

Skeleton of ^(/r«0Jfl«r«j (Lacertilia). 271 . After Seeley. Three fourths natural size. 183.Fig.

less well-ossified carpus and tarsus. calcified No clavicles.ossified carpus. the almost entirely cartilaginous mesopodials and highly developed hyperphalangy. Three types sals. A sternum. a well. a diving type. and only slight hyperphalangy. The mosasaurs distribution during Upper Cretaceous times. webbed. maxillae. The mosasaurs were clothed with small Fara«w5-like scales. hyperphalangic. zygosphenes. differing from them chiefly in the loss of the sacrum and the adaptation of their limbs to purely aquatic uses. with Platecarpus as a type. Palate with large openings. the tabular with a long process wedged in between paroccipital and prootic. perhaps rightly. were soft. a long. and greater hyperphalangy. In all probability they were descended from subaquatic lizards like the aigialosaurs in late Lower Cretaceous times. short. Nasals and premaxillae fused into a single bone. and greater size.272 THE OSTEOLOGY OF THE REPTILES Tribe Pythonomorpha (Mosasauria) Large marine lizards with more or less elongated head. dentaries. body with but twentytwo vertebrae. forty feet in length. sometimes a slender interclavicle. of which Tylosaurus is the best-known type. flattened tail with a more or less sub- terminal dilatation. less elongated trunk with but twenty-two more uniformly flattened tail. . recognized as distinct families. Seven cervicals. And these vertebrae. a deeper-sea type with proportionally shorter neck. a three groups have been. From six to about Temporal and postorbital arches complete. a relatively short neck. Vertebrae procoelous. rami of mandibles united by ligaments. a longer and much flattened tail. elongated body. pentadactylate. of which Mosasaurus and Clidastes are types. and pterygoids. doubtless impregnated in life with fat. A true joint between angular and splenial. Parietal and frontal unpaired. Teeth with osseous base inserted in shallow pits in premaxillae. of which impressions have been often found. and paddle-like extremities. heavy cartilaginous protections for the ears. The bones. with more elongated head. Legs paddle-like. of mosasaurs are recognized: the surface-swimming type with elongated trunk composed of as many as thirty-five dor- the tail with a pronounced subterminal dilatation. are a group of large marine lizards. a parietal foramen. No sacrum. especially of the deep-diving forms. of world-wide without claws. Sclerotic plates present sometimes with zygosphenes. shortened neck.

the quadrates fixed by the ptery- and tabulars indistinguishable. moving vertical rather than lateral undulations. Clidastes Mosasaurus Conybeare. Family Globidentidae. 273 Mosasauridae. Baptosaurus Marsh. Europe and North America. squamosals large. no postorbitals. . Russia). the The im- soUd skull with the palate firmly movable quadrates. Upper Cretaceous. Globidens. No parietal foramen. Mosasaurus Europe (England. fixed. burrowing lizards. PlioConybeare. and characters almost as far entire absence of arches. ?Sironectes Cope. North America. either legless or with short tetradactyl front limbs immediately back of the skull. Body with numerous rings and without scales. Upper Cretaceous. Platecarpus Cope. Belgium. Tribe Amphisbaenia Worm-like or snake-like. Teeth spheroidal. reaching a length of about one and one-half feet. together with the vestigial or absent limbs. Gilmore. France. Teeth conical. Palate without openings back of the nares. platecarpus Dollo. Stapes short and stout. rugose. No postorbital or temporal arch. A by curious group of burrowing Uzards.THE SUBCLASS PARAPSIDA Family pointed. are removed from the typical lacertiHan structure as are those of the snakes. Haino- saurus Dollo. Vertebrae procoelous. the nasals goids. Taniwha- New Zealand. lacrimals. Brachysaurus Williston. Macrosaurus Owen. saurus Hector. Imperfectly known. PrognatJwsauriis Dollo. Brain-case in front partly enclosed by plates from frontals. Cope. or jugals. the tail very short and blunt. Eyes minute.

vague. Eocene. and slender. Rhineura Cope. gia] . Europe. Chameleo (Leidy). . premaxillae small or vesti. two sacrals. With the characters Eocene. North America. . North America. France. Upper Cretaceous. confined to Mada- and Hzards. 3. Oligocene. fano. Pleistocene. though it is probable that some referred to it will eventually be found to have all the essential characters of the group. Family Chameleontidae. Saurillus South Africa. Diacium Cope.2 74 THE OSTEOLOGY OF THE REPTILES and seem to be as important in classification as those distinguishing the much more typically lizard-like mosasaurs. 4. the digits in opposable groups of two and three. palate with openings. ? ? Sedis Paliguana Broom. 3. No extinct lizards are certainly referable to this tribe. epipterygoids absent or vestigial. Cremastosaurus Cope. and for their Our paleontological knowledge of them is power to change color. formula 2. Hyporhina Baur (a postorbital arch). Saurospondylus Seeley. Enigmatosaurus Nopcsa (de Stefano). Africa. Platyrhachis Cope. ? Patricosaurus Seeley. long famous gascar. fifty living species A group composed of about India. prehensile tail. ? Tylosleus Cope. arboreal. England. North America. five cervicals. digital Clavicles absent or vestigial. of the group. Jura. North America. Jurassic. A curious group of insectivorous tree for their peculiar grasping digits. Notiosaurus Owen. the spines sometimes elon- gated. Arches complete. Aciprion Cope. Vertebrae procoelous. slender. Owen. England. the quadrate Postfrontals indistinguishable. 4. Prochameleo de Ste- Genera Incertae Triassic. Abdominal ribs present. Mesopodials much reduced. Family Amphisbaenidae. from eleven to fifteen dorsals. perching lizards. England. Tribe Rhiptoglossa Small. no septomaxillae parietals and f rontals unpaired no parietal foramen. With the characters of the group. Coniosaurus Owen. Naocephalus Cope.

The quadrate articulates loosely with the tabular only. not more than fifty or sixty species altoand of them with very few exceptions their fossil remains are few and fragmentary. probably the complete bony and there is never a vestige of a pectoral girdle. and caudal series. no parietal foramen. epiptery golds. the coronoids sometimes absent. to be evolved vipers are among the Reptilia. Anterior vertebrae. two or three without cervical. jugals. no squa- mosals. though several families have vestigial pelvic and hind limb bones. Parietals fused. sometimes with vestiges of hind limbs but never with front Umbs or pectoral girdle. There are no temporal arches. .THE SUBCLASS PARAPSIDA B. and sometimes no ectopterygoids. 275 Suborder Ophidia (Serpentes) Elongated. are numerous. And chiefly because of such upland habits they are very scantily represented gether. includes more than eighteen hundred Hving species widely distributed over the earth. most decisive. sometimes exceeding four hundred first in divisible into precaudal ribs. the dentaries loosely articulated. scarcely a single positive character to distinguish them the .case. but more or less of the caudals with a descending proc- on each side (lymphapophyses) This suborder. The mandibles are united in front by ligaments only the posterior bones are . The pterygoids and lae are small usually the palatines have teeth. and of the snakes the poisonous probably among the latest. lacrimals. The premaxil- and often edentulous. in a few instances even the tabular is absent (Uropeltidae) The brain-case in front is enclosed by descending plates from the parietals and frontals to the sphenoid. a others are aquatic. often fused. as has been mentioned. postoptics. from the latter sometimes in- terrupted by the coalescent optic foramina. maxillae rarely edentulous. legless reptiles of from a few inches to thirty feet in length. and still restrial in habit. sometimes to the caudals with a more or less prominent hypapophysis. Most snakes are purely terfew are burrowing. Teeth acrodont. The vertebrae number. and their taxonomic relations very doubtful. often considered an order. the Always procoelous and always with zygosphenes and zygantra. among fossils. No ess chevrons. Prootics largely visible. Like so there is many groups of organisms known is in many related forms. Probably the snakes are the latest group of equivalent rank closure of the brain.

England. They are burrowing in habit. Eocene. Boas. Cope. North America. of some them attaining a length etc. maxillae and mandibles edentulous. spines elongate. East India. Calamagras North America. Ogmophis Cope. Maxillae vertical. Botrophis Mercer. pres- Maxillae horizontal. — Ed. Ectopterygoids present. Vestiges of pelvis present. Heteropytlwn Miocene. Paleophis Owen.] . vertebrae with an Family Paleophidae. toothed. Family Viperidae. Protagaras Cope. . Tabular long. No illae coronoids. France. Oligocene. Limnophis Marsh. Paleophis Owen. or short and closely attached to the skull (Illysiidae) Vestiges of hind hmbs present. of nearly thirty feet. the single known vertebra is more probably that of a dohchosaur lizard. Lestophis Marsh. A single extinct form {Symoleophis Sauvage) from the Cretaceous of France (Senonian) has been referred here. imperfectly known.^ and copperheads) exclusively ' [Occur also in Asia and Malaysia. The Typhlopidae with but a single living genus and about one hundred species are widely distributed in the tropical regions. Python Daudin. Europe. Neural inferior ridge. Max- vertically erectile. Rochebrune. Paleopython Rochebrune. Ectopterygoid and coronoid ent. Poison fangs perforated. widely distributed. Family Boidae (Pythonidae). North America. Aphelophis Cope. excavated (Crotalinae) or not (Viperinae). Patagonia. About one hundred erectile fangs are living species of these poisonous snakes with known. Pterosphenus Lucas. No ectopterygoids or tabulars. A family of wide distribution comprising about sixty species. Pit vipers (rattlesnakes in America. France. Upper Cretaceous. reaching premaxillae. Pliocene. Large snakes. Eocene. Dinilysia Woodward. anacondas. pythons. Paleryx Owen. France. the latter with or without teeth. probably subaquatic. Boavus Marsh.276 THE OSTEOLOGY OF THE REPTILES Family Typhlopidae. articulating with prefrontal. Scytalophis Rochebrune. Scatophis Rochebrune. with sohd teeth.

THE SUBCLASS PARAPSIDA Uppermost Cretaceous. Elaphis Aldrich. They are practically unknown as fossils. comprises nearly wider sense including the cobras. Oligocene. Cope. present. Maxillae horizontal. living none attaining a is size of more than seven or eight feet. Tabular present. North America. Family Colubridae. Europe. Post- orbital not produced forward. Tamnophis Rochebrune. present. Miocene. Pleistocene. Their distribution world-wide. Periops Wagler. Ectopterygoids not erectile. deeply grooved or hollowed. Fylemophis Rochebrune. with solid teeth. Pleistocene. Germany. in its This family of highly poisonous snakes. Caudal hypophyses Laophis. Salonica. sea snakes. all This family of harmless snakes includes more than half of species. Eocene. Ectopterygoid sent. North America. Coluber Linne. their anterior teeth bifid. the coronoid ab- Maxillae horizontal. . but doubtfully. North America. Miocene. North America. ? ? 277 Coniophis Marsh. Helagras Cope. and the coral snakes of the southern United two hundred living species. Neurodromicus Family Elapidae. States. Cobras (Naja Laurenti) have been reported from the Pleistocene of France. Crotalus Linne. Viper a Laurenti. North America.. Europe and North America [ =] Bas- canion Baird and Gerard.

12. Skull short. Order "Eosuchia" Skull Family Younginidae. the birds. Youngina Broom. Upper Permian. Proterosuchus Broom. — Ed. no antorbital vacuity^. below. the lateral by the postorbito- squamosal arch above. tostylic. admits of Httle or no doubt. [An antorbital vacuity is present. ? Order Proterosuchia an antorbital vacuity.] 13. from the Upper Permian of Africa. Skeleton otherwise unknown. South Africa. At present the oldest known form referable to the subclass is Youngina. the Theromorpha. the postorbito-squamosal arch below. an intermediate type peculiar in its retention of certain skull bones lost in bers. probably with palatal teeth. by the simple separation parietal. all other It is. [Triassic. ability they In much prob- were derived from the single-arched type with the of the postorbital lateral opening. this and squamosal was confidently believed that the representative of the subclass was Paleoit In all probability. A single coracoid on each ischiatic opening. Doubtless memmany other forms from the Permian with these and yet other primitive characters await discovery. only known. tainty. the upper bounded by the parietal above.CHAPTER Two XII THE SUBCLASS DIAPSIDA temporal openings. according to Broom. side. if not cer- form did not have the upper temporal opening.] . or jugal and quadratojugal. Skull Skull elongate. yet very imperfectly known. with palatal teeth. with interparietals and tabulars (Psupratemporals). from the Until recently most primitive and oldest hatteria. and must therefore be included in the more primitive group. no cleithra. never strep- The phyletic unity of this great division of reptiles and their descendants. however. the jugal. Pelvis with puboQuadrate fixed or partly movable. South ^ Africa. from the Lower Permian.

. 279 SUPERORDER DIAPTOSAURIA all Teeth on some or the palatal bones. and edentulous beak. Except the living Sphenodon.three presacrals. as to the value of the groups. The differences between them seem hardly greater than among the Lacertilia with the inclusion of the Pythonomorpha. present. shore-dwelling and shell-eating reptiles from three to six feet in length. Palate primitive. Suborder rhynchosauria Skull more or less depressed and broad. No Nares undivided.THE SUBCLASS DIAPSIDA A. goidal opening. with teeth on some or girdle complete. The complete skull. most of the genera are yet incompletely known. parietal foramen. as usual. decurved. Several groups formerly placed under 14. Temporal openings relatively large. articulating in part or chiefly to centrum. and in the differences of opinion. Two or three sacral vertebrae. central space and The three groups of reptiles here considered suborders are by some authors given a family rank. acrodont or protacrodont. A. ORDER RHYNCHOCEPHALIA all Terrestrial or littoral lizard-like reptiles of small or moderate size. Parasternal In the absence of more complete information as to the structural details of some of the forms included under this group name. A small group of terrestrial. it are now relegated to other divisions. the bones. A small pubo-ischiatic vacuity. No antorbital opening. the tribe or superorder Diaptosauria has a present use. none. perhaps in some cases subaquatic. Palate with small interpteryDorsal intercentra absent or unknown. Dorsal ribs holocephalous. Pectoral Dorsal ribs holocephalous. no interparietals or tabulars. by others ordinal. Vertebrae amphicoelous. formed by the premaxillae. tail. their boundaries as in the Sphenodontia. with a strong. Phalangeal formula never reduced. two sacrals. articulating in interarch. Humerus without epicondylar foramina. About seven or eight cervicals and twenty. Fifth tarsale ribs absent. In Howesia a distinct and mesopodials are known in intermedium tarsus is figured if not an .

. After Woodward. However. where they occur on the pterygoids only. Five sixteenths natural size.28o THE OSTEOLOGY OF THE REPTH^ES only error. save in Howesia. 185. The palatal teeth are confined to the palatines in two or three rows. S^tXtton oi Rhynchosaurus (Rhynchocephalia). it is the known example among reptiles. the Fig.

Ardeosaurus is . dentaries. sometimes an ectepicondylar foramen. Hyperodapedon Scotland. Brachy- rhinodon Huene. Germany. A single row of acrodont teeth on maxillae. squamosal. Brachyrhinodon has two temporal arches. Homceosaurus. page 268 above. It has also no ectepisaid to condylar foramen present in Sphenodon. and the single-headed ribs of primitive type. and palatines. except that it has no uncinate process on the a character in which Spheno- don is almost unique among reptiles. Premaxillae with a decurved beak. South Africa. Howesia Broom. L. Pelvis with large pubo-ischiatic vacuity. can be located with certainty in this suborder.] Camp (1923). deeply amphicoelous vertebrae. postand postorbital. Stenometopon Boulenger. No lacrimals. Upper i[But Jurassic. The former. and still im- mediate relationships with the other genera are in doubt. but saurus the skull is poorly known otherwise. Triassic. Scotland. have teeth quite like those of Ardeosaurus which. Scotland. Suborder Sphenodontia (Rhynchocephalia vera) Upper temporal opening bounded by parietal. is a near relative of Acrosaurus. Ardeosaurus v.THE SUBCLASS DIAPSIDA anterior part of the skull of this genus is 281 its poorly known. the neck with not present. According to C. Upper Huxley. Germany.^ cf. B. more than eight. South Africa. ? Middle and Upper Eifelosaurus Jaekel. Meyer. also with vertebrae. usually with teeth. Rhynchosaurus Owen. Humerus with an entepicondylar foramen. Triassic. are Palacrodon and Opisthias is known only from mandibles. Palacrodon Broom. India. A parietal frontal. related to the geckos. however. Parasternal ribs Two genera only. Twenty-three to twenty-five presacral foramen. Frontals and parietals paired. according to Nopcsa. the living Sphenodon and the Jurassic Homoeo- saurus. ? Polyspheno- don Jaekel. Nor is the temporal region of Ardeosaurus as well known is as one could wish. as evidenced by the persistent dorsal intercentra. — Ed. Sphenodon has long enjoyed the reputation of being the most primitive of living reptiles. So far as known Homoeosaurus ribs. Carpus primitive. England. v. agrees closely. Meyer. Probably it has dorsal inter- centra. but this remains to be determined. Of Eifelo- wholly unknown.

Natural size. New Zealand.282 THE OSTEOLOGY OF THE REPTILES Opisthias Gilmore. Fig. Sphenodon Gray. Skeleton of //o»/aroJ/?«r« J (Rhynchocephalia). Wyoming. Recent. After Lortet. 186. Lowermost Cretaceous. .

otherwise rinthine teeth unknown and the absence rangement of the bones of the temporal region doubtful. those of the prevomers. The arThe legs are essentially terrestrial in structure. mosasaurs. the dentition intermediate between that of the Mosasauridae and that of the Globidentidae. The relationships between the known genera are very close. broad. Elongate. ? SUBORDER THALATTOSAURIA face. anterior. but the heavy flattened ribs and the elongate flat- tened tail decisively indicate bottom-crawling aquatic habits. articulating with centra. reptiles. amphicoelous without dorsal intercentra. This small group of water eight feet. Simosdosaurus Gervais^ France. Belgium. crowned. tail. Twenty-six presacral. is characters.THE SUBCLASS DIAPSIDA C. The upper opening The habits limbs. Teeth labyrinthine in structure. with small teeth on Vertebrae shallowly all No parietal foramen. A parietal foramen. subaquatic reptiles. Champsosaurus Cope Leidy). {Nothosaurops North America. ectepicon- Humerus with dylar foramen. with- out foramina. 283 SUBORDER CHORISTODERA palatal bones. openings. The . mandibular. sclerotic plates. and a long. Uppermost Cretaceous and Paleocene. so far as of the thalattosaurs known. flattened Dorsal ribs holocephalous. Premaxillary. two or three sacral. and heavy. Mesopodials imperfectly known. Humerus short. with but sUght aquatic adaptations. These small perfectly reptiles of is it known. and posterior part of maxillae and mandibles lownares. D. posterior[ly placed external] Marine reptiles with elongate and paddle-like extremities. with a very slender face. resemble those of the must have been similar to those of the mosasaurs. ribs intercentra chiefly unknown. and pterygoidal teeth conical. Dorsal holocephalous. is animals reaching a length of of interest because of the retention of several primitive in the Diapsida. terminal undivided nares. Parasternal ribs slender. Parasternals stout. especially the labyof a pubo-ischiatic opening. nor but three or four feet in length are still imquite certain that they have two temporal occupies a peculiar position. Internal nares posterior. Pelvis without pubo-ischiatic opening. Vertebrae rather deeply biconcave.

The epi- podials are long. The quadratojugal is Usually an and usually parietal fo[if] enters the border of the lateral temporal opening. Necto- saurus Merriam. by two articulations. the corocoid not elongate. Body usually with dermal armor.. Vertebrae amphicoelous or platycoelous. phalanges not reduced. slender. perhaps. SUPERORDER ARCHOSAURIA Dorsal ribs attached exclusively to the arch. and a large antorbital opening. and doubtfully. California. or [dermojsupraoccipitals. Clavicles and interclavicle present. ORDER PARASUCHIA pelvis. Typically a group of small. the cervicals to arch and centrum. 15. ever. the interparietals. A. Teeth thecodont. in the present sense The Parasuchia were long united with the str. Crocodilia as two suborders. at least anteriorly. sens. crawling or leaping reptiles. None of the forms referred to this suborder is completely known. tabulars. Roof bones of skull always paired. By some authors the three Sclerotic suborders here recognized are each given ordinal rank.284 THE OSTEOLOGY OF THE REPTILES Triassic. the Parasuchia. postfrontals present. as are also the antorbital openings. cHmbing or leaping reptiles with more or less elongated hind The external and internal nares are near the extremity of the more or less pointed skull. A A. plates are known in a single genus of Pseudosuchia. Vertebrae never notochordal. No ramen. well developed antorbital vacuity. but the marked differences in skull and pelvis justify their ordinal separation. the lateral orbits are large. the clavicles and interclavicle slender. character- ized especially by the normal absence of a secondary palate. some departing so widely. confined to jaws. rarely absent. and among the known forms there is a considerable diversity of structure. Middle and Upper Thalattosaurus Merriam. nor the dorsal intercentra persistent. that their location here Of the more typical. Parasternal ribs generally present. From small to rather large. . Scleromochlus has no dermal is provisional. Mesopodials imperfectly known. Suborder Pseudosuchia legs. and the Pseudosuchia or Aetosauria.

ings large. and the attachment the group is of the dorsal ribs are decisive. Dorsal scutes transversely elongate. covering the plates. the only member Ornithosuchus Newton. Connecticut. the epipodials longer than propodials. internal nares near extremity of triangular skull. South Africa. whole back. presacrals. accredited with an interparietal. three sacrals. two sacrals. No dermal armor. Palatines approximated or con- . Triassic. the latter has been With the inclusion of the doubtful forms there are but few con- stant characters to distinguish the group from the Rhynchocephalia typically. and Euparkeria alone has sclerotic plates. Euparkeria Broom. long and slender. coracoid short little and Legs very slender. Family Aetosauridae. the orbits relatively small. abdomen with small Triassic. Twenty-one Tubes Scapulae slender. Family Scleromochlidae. Premaxillae united. Slender parasternal ribs. Stegomus Marsh. Upper temporal opening not depressed below level of [parietals]. however. one wide genetic possibilities all and [may] have had a close genealogical relationship with the other members of the Archosauria. ? Erpetosuchus Newton. propodials. Twenty-five merus a little presacrals. Dyoplax Fraas.THE SUBCLASS DIAPSIDA accredited with an interparietal bone. Euparkeria of the group. B. Hu- longer than radius and ulna. Nearly every known genus has been ac- credited with family rank. Scleromochlus Woodward. Triassic. hind legs a half longer than the front. Scapula broad. expanded at extremity. Family Ornithosuchidae. 285 armor. slender. the absence of palatal teeth. Suborder Pelycosimia External and Antorbital open- Large. terrestrial or marsh reptiles. feet as long as tibia. each longer than broad. Aetosaurus Fraas. however. England. Sphenosuchus Haughton. coracoid long. calcaneum with tuberosity. the epipodials a longer than the Two is rows of dermal plates. Germany. and especially the Saurischia. heavily built. Eng- land. of As a whole.

Vertebrae platycoelous two sacrals. Suborder Desmatosuchia] by the probably secondary absence of Cervical and anterior dorsal bony [Large. India. the pectoral region and abdomen with bony scutes. Skull with large antorbital . Mesorhinus Jaekel. Legs short and rather stout. South Africa. Rutiodon Emmons (Rhytidodon) . ? Scaponyx Woodward. Feet probably webbed. Paleorhinus Williston. plates bearing long horn-like outgrowths. based almost excluis This group. without respiratory canal. the lateral. and the deep respiratory canal. Neck. New York. body. the posterior nares. Mystriosuchus Fraas. In the structure of the skull it some- what intermediate between the Pseudosuchia and the Phytosauria. compressed. Angistorhinus Mehl. South America. Triassic. Ilium with- Family Stagonolepidae. Skull rugose. Suborder Phytosauria characterized Large. pubis not dilated at extremity. cylindrical teeth on each. Carolina. especially characterized the upper temporal opening. temporal. sharply pointed. and is tiguous. Phytosaurus Jaeger. sively upon Erythrosuchus. . proposed as a separate order. Tip of premaxillae decurved. Lophoprosopus Mehl. IHum with postacetabular process. curved. formed by the underarching of the palatines. Family Phytosauridae. drical throughout. and antorbital openings large. England. A supracoracoid foramen. subaquatic reptiles. out postacetabular process. Stagonolepis Huxley. Rocky Mts. Connecticut. pubes dilated at extremity. Parasuchus Lydekker. with two or three very long. and tafl covered with four or more rows of strong dermal bones. separated. Germany. reaching a length of twenty or more feet. com- posed chiefly of the premaxillae. Tail long and flattened. Triassic. especially by the elongate face.286 THE OSTEOLOGY OF THE REPTILES Teeth compressed. c. [D. long-tailed reptiles reaching a length of perhaps sixteen feet. Erythrosuchus Broom. the supratemporal opening small and more or less depressed below the plane of the parietals. Triassic. crawling. or the posterior ones more or Teeth either cylinless flattened and smaller.

Nat. Some have face.walking gait. All are lizard-like in form. flattened tail. Both the carpus and tarsus are peculiarly modified. with conical the- codont teeth. Desmatosuchus Case. Amer. The verte- brae were platycoelous in Cretaceous. see numerous papers by C. osseous armor along the back and tail. very small in the broadfaced types. suggesting. Hist. Triassic. Two sacral vertebrae. Ace- tabulum formed by ilium and ischium only. — Ed. a primitive. snout not greatly produced. Von Huene refers Desmato- suchus to the Phytosauria. a relatively broad skull. Huene thinks. the so-called pubes (? prepubes) excluded and not meeting in a median symphysis. Teeth thecodont. The upper temporal openings in the modern forms are smaller. sometimes also on the under side. the external nares terminal. Distinguished from the Phytosauria especially by the absence of the upper temporal opening.THE SUBCLASS DIAPSIDA 287 opening and dorsal anterior nares. Mook. very predaceous. and in the later forms the pterygoids Premaxillae never much elongate. which may have been secondarily lost as in the caimans. maxillae and palatines. v. and the splenials meet in a symphysis. In such forms the nasals do not reach the external nares. Texas. ORDER CROCODILIAN [Loricata] Internal nares carried far back in the mouth by the union of the also. others a more or less elongated sometimes very slender. Mus. as in the ancient teleosaurs and the modern gavials. There versity of structure not a very great among the known forms. with a long. In size they than one foot to about all till fifty feet in length. 1921-.] 16. is Wealden times the bar more cylindrical 1 [For recent morphological and taxonomic treatment of the Crocodilia. The Crocodilia are at once distinguished from is all other reptiles by di- the structure of the palate and pelvis. The amphibious crocodiles have a strong dermal. and more or vary from less less water-loving in habit.] . Bulletin. Phalanges of fourth and fifth digits reduced. procoelous in about the beginning of the Lower all since the early part of the Eocene. C. since and more deeply placed. calcaneum elongate. more upright. In the early types the arch between the orbit and lateral temporal opening was covered immediately by the skin.

reach to the premaxillae. Mandible with an external vacuity Nine cervical vertebrae. Suborder Eusuchia many an- An cient antorbital opening primitively present but lost in and all posteriorly. and still by some authors. Internal Family Pholidosauridae. orbits. Vertebrae platycoelous. Head short.2 88 THE OSTEOLOGY OF THE REPTILES A. Suchodus Lydekker. Feet partly webbed. An antorbital not modified. Face long. Teleido- saurus Deslongchamps. probably extending on tail. amphicoelian forms comprised in the suborder Mesosuchia. Postorbital bar slender. broad. Jurassic. known Family Teleosauridae. Small reptiles from eight to sixteen inches in length. twenty-three or twenty-four sclerotic plates in orbits. Aeolodon Meyer. Dermal armor composed of two rows of quadrilateral plates. Family Atoposauridae. Vertebrae platycoelous. No Body with dermal bones. Until within recent years. It is now that the change in the form of the vertebrae was a relatively unimportant one and may have occurred in different lines of descent. the Eusuchia comprised only those crocodilians with procoelous vertebrae. Teleosauridae. Madagascar. Posterior nares not reaching pterygoids. tral scutes. Probably no ven- Tail long. modern forms. PhoUdosaunis Meyer (Macrorhynchus). presacrals. Steneosaurus Geoffroy. . Cretaceous. Vertebrae large. Europe. ? Teleorhinus Osborn. situated at posterior platycoelous. Teleosaurus Geoffroy. Upper Jura and Lowermost Cretaceous. Front feet much smaller than hind. Europe. Wyoming. dermal armor. . nares opening in palatines and pterygoids. Upper temporal opening large. Europe. From two to ten feet in length. not paddle-like. Internal nares Face very long and opening sometimes present. the nasals Upper temporal opening smaller than Front legs larger than in the Postorbital bar modified. Pelagosaurus Bronn. Dorsal and ventral armor present. Gnathosaurus Miinster. Upper temporal openings much smaller than orbits. Crocodilemus Jourdan. A nearly complete end of palatines. clawed. Pterosuchus Owen.

Posterior nares surrounded by pterygoids. six to forty or Sometimes an antorbital opening. Posterior nares elongated. Family Tomistomidae. broad. [Tomistoma. Face rather broad. Recent. procoelous. From ten to fifty feet in length. Dyrosaurus Pomel. Family Gavialidae. From length. India. Bernissartia Dollo. Coelosuchus WilHston. Florida. more feet in Upper Cretaceous. Goniopholis Owen. 289 Jourdan. Skull short. Nasals extend into nares. Europe. Upper temporal openings large. PaleosMchus Falconer and Cautley. Hylceochampsa Owen. Belgium. Europe. Nasals remote from nares. ?Heterosuchus Seeley. Family Dyrosauridae. South America. Atoposaurus Meyer. by pterygoids. Machimo- saurus Meyer. Lowermost Cretaceous. Africa. Tomistoma (?) [Gavialosiichus]. Vertebrae platycoelous. Vertebrae surrounded by pterygoids. Family Hylaeochampsidae. Lower Eocene. Wealden Cretaceous. not long. Pleistocene. Upper Cretaceous. Wyoming. Rhamphosuchus Owen. Xheriosuchus Owen. North and N annosuchus Owen. Thoracosaurus Leidy. Family Goniopholidae. gradually merging Postorbital bar subdermal. Face very slender. A dorsal armor of two or more rows of plates. Germany. internal nares Face very slender. Vertebrae platycoelous. Internal nares bounded by pterygoids and palatines. bar subdermal. South America. States. Postorbital bar subdermal. between palatines and pterygoids. . Dorsal but no ventral scutes. Internal nares surrounded England. From fifteen to eighteen feet in length. Vertebrae probably procoelous.THE SUBCLASS DIAPSIDA Upper Jurassic. Face less into cranium. Postorbital bar subdermal. Palate with large foramen between ectopterygoid and maxillae. Alligatorellus Alligatoriiim Lortet.] Eosuchus Dollo. Postorbital Africa. United Eocene. Vertebrae procoelous. Gavialis Oppel. Cynodontosuchus Woodward. Teleorhinus Osborn. Holops Cope. Notosuchus Woodward. Europe. Pliocene.

[?]. Africa. North America. Oligocene. Leidy osuchus Lambe. ? Polydectes Cope. single or divided. Postorbital bar subdermal. [Alligator]. Supratemporal openings Bones of skull smooth. 1 [Williston here includes the genera Alligator and Caiman under the Crocodilidae. large. Family Crocodilidae. North America.] Incertae Sedis. — Ed. protuberant. North America. less elongated. North America. Marine less crocodiles. Jacare. without claws. Limnosaurus Marsh. Upper temporal openings small. Pleistocene. Miocene. Osteoblepharon. Tomistoma. Eocene. Italy. Hungary. Jurassic. Dorsal plates in two or more rows. Face more or Prefrontals Nares at posterior end sclerotic plates. Orbits with Seven cervical. Leptorhamphus Ameghino. and with Umbs more or Vertebrae platycoelous. The nasals usually reach the external nares. South Dakota. Face never slender.] Crocodilus. [Recent.] . Tail long. Brachychampsa Gilmore. the ventral armor present or absent. ''Crocodilus'' Caimanoidea Mehl. From four or five to more than forty feet in length. Diplocynodon Pomel. Uppermost Cretaceous. Crocodilus Laurenti. and Jacare are more distinct from Crocodilus and its allies {Osteolae- mus. Osteolaemus. and places Tomistoma in a separate family. South B. with distal fin-like dilatation. Crocodilus Laurenti. Lower Africa. Borneo. North [Alligator.) has shown that Alligator. Recent. Bottosaurus Leidy [Agassiz]. North America. Notochampsa Broom. Caiman. anisodont. Oxydontosaurus Ameghino. Alligator. Argentina. No antorbital presacral. India. Posterior nares surrounded by pterygoids. twenty-five or mandibular openings. Osteoblepharon) than is Tomistoma. Europe. of palatines. Cai- man.^ Vertebrae procoelous. vertebrae. Crocodilus Laurenti. cil. Teeth stout. Deinosuchus Holland.290 Pleistocene. Suborder thalattosuchia without bony armor. Europe. large. Tertiary. but Mook {op. America. THE OSTEOLOGY OF THE REPTILES Tomistoma. modified as paddles.

Dollo. the Triassic. not recurved and somewhat thickened. Gressylosaurus Riitimeyer. Family Plateosauridae. bipedal in gait. 4. Hind digiti- more mesaxonic. with a distal dilatation. Front legs a formula little longer than the femora. Europe. Europe. though of rather heavy build. the hind feet more or less digitigrade. Huxley. the front legs more or less reduced.THE SUBCLASS DIAPSIDA Family Metriorhynchidae. 29 Quenstedt. Upper Plateosaurus Meyer. three sacrals. in a ventral symphysis. Astragalus without ascending process. 3. Jaekel thinks that the hind feet were purely plantigrade. Teeth less compressed. No No dermal bones. Geosaurus Cuvier. (?). Feet grade or semiplantigrade. Europe. 5. the first claw large. is thought by some to have an ancestral relationship with the Sauropoda. The reptiles were clearly bipedal in gait. One or more antorbital openings. More ischia or less upright. Teratosaurus Meyer. Europe. . the Euskelosaurus most primitive family of the Theropoda. Patagonia. Anterior vertebrae platycoelous twenty-three presacrals. Melriorhynchus Quenstedt. their phalangeal feet 2. Gryponyx Broom. ORDER SAURISCHIA The normal pubes and acetabulum perforated. A. ? Enaliosuchus [DINOSAURIA] 17. South Africa. Plateosaurus attained a length of about fifteen feet. Pubes meeting in a long ventral symphysis. but this was improbable since the mesaxonic structure distinctly indicates the elevation of the ankle from the ground. Pachysaurus Huene. their anterior and posterior borders denticulated. The characters drawn chiefly from Plateosaurus may not and probably do not apply to all the genera listed in the family.walking reptiles. the meet predentary or rostral bones. Upper Jurassic. This. . Neustosaurns Raspail. preaxonic. Dacosaurus Lowermost Cretaceous. first and fifth toes reduced. Suborder Theropoda More or less Carnivorous or secondarily herbivorous in habit.

less recurved.] B. including the Ornithomimidae. Skeleton oi Gorgosaurus (Saurischia).. Family Anchisauridae. containing several families and numerous genera of light-limbed saurischian dinosaurs. see papers . hyposphene-hypantrum Four or hve and with sacrals. part of jaws. (2) the Megalosauria group of the Fig. Africa. more or Astragalus without ascending process. vol. One thirty-sixth natural size. Postfrontal sometimes present. with long tail and small skull. in a single row. XLVi). Mus. Hist. Nat. with a thickened. semiplantigrade. Vertebrae amphicoelous. no predentary. No coronoid process to mandible. The sometimes all. and more or anterior. Megadactylus Hitchcock. Jurassic. I and (3) the Deinodont group of the Cretaceous. 187. Teeth compressed. Mus. Connecticut Valley. — Ed. spoon-shaped crown. CETIOSAURIA) neck and Quadrupedal. England.. II) for groups 2 and 3 see Matthew and Brown. After Lambe. Amer. Bd. herbivorous dinosaurs. Zanclodon Plieninger. more or less developed hollow. Europe. with a articulation. XLiii).292 THE OSTEOLOGY OF THE REPTILES Smaller and more slender theropods. SUBORDER SAUROPODA (OPISTHOCOELIA. [No MS. was found for (i) the Coelurosauria. twenty-six or . Massospondylus Owen. vol. Anchisaurus Marsh. lateral cavities in centra. Hist. Teeth less restricted to anterior subcylindrical. For group by Osborn 191 7 {Bulletin. Ammosaurus Marsh. the premaxillae with teeth. AustraUa. von Huene 192 1 {Acta Zoblogica. Upper Triassic. Amer. 1922 {Bulletin. Sellosaurus Huene. presacral vertebrae opisthocoelous. Thecodontosaurus Riley and Stutchbury. Nat. South Africa.

Rocky Mts. of dorsal vertebrae simple. Titanosaurus Lydekker. Madagascar. Brachiosaurus Riggs. Lower Cretaceous (Morrison). Scapulae narrow distally. Dystrophaeus Cope. Spines of dorsal vertebrae furcate. Cetiosaurus Owen {Cardiodon Owen). Chondrosteosaurus Owen. Ischia slender. Haplacanthosaurus Hatcher. ity. Atlantosaurus Marsh. More slender sauropods. Rocky Mts. Gigantosaurus Fraas (non Seeley). Rocky Mts. Eucamerotus Hulke. Europe.] . England. South Africa. Family Atlantosauridae. Front hmbs distinctly shorter than hind. From about fifteen to about ninety Family Cetiosauridae. feet in length. not distally. Front legs shorter than hind. 293 twenty-seven presacrals. Scapulae dilated Pubes not constricted. Carpals and tarsals reduced. Amphicoelias Cope. Bothriospondylus Mantell. Upper Jurassic. France.THE SUBCLASS DIAPSIDA ventral symphysis. Spines Front legs much shorter than hind. feet preaxial. India. Marsh. North America. Lower Cretaceous (Morrison). confined to anterior part of jaws. Spines furcate. of presacral vertebrae furcate. The pubes are massive and meet in a large Limb bones cancellous in structure. {} Brachiosaurus Riggs). Front limbs as long as the hind.^ Family Diplodocidae. Rocky Upper Jura and Wealden. remote from extremIschia not expanded distally. Ischia expanded at extremity. Europe. Sedis: ? Jurassic. — Osborn and Mook. Scapulae distally ex- Lower Cretaceous. Teeth slender. Pelorosaurus Man tell. Diplodocus Marsh. Titanosaurus Lydekker. Madagascar. Lower Cretaceous (Morrison). Family Camarasauridae. Ischyrosaurus ^ [Belongs in Diplodocidae. panded. Patagonia. the pubes constricted in middle. Upper Cretaceous. Spines of presacral vertebrae furcate. External nares near top of skull. Genera Incertae Mts. Pleurocoelus Camarasauriis Cope {Morosaurus Marsh). Apatosaurus Marsh (Brontosaurus Marsh). Chondrosteus Owen.

digitigrade. Anterior vertebrae opisthocoelous. especially characterized by the presence of a predentary bone in the mandible and by the structure of the pelvis. sis. usually long. — Lull. Lower Cretaceous (Morrison). Hypselosaurus Matheron. rarely with as many as four phalanges in any ally tridactylate. Antorbital opening face. Oplosaurus Gervais. Algaosaurus Broom. Of the size of a cat. Microcoelus Lydek- Lower Cretaceous. or less Hind Umbs usually functiondigitigrade.] . Nesodon Mousaye. Femur shorter than tibia.2 Elosaurus Peterson and Gilmore. Epanterias Cope. Rocky Mts. Teeth in a single row. Dinodocus Owen. South America. postpuhis. slender. Argyrosaurus Lydekker. Europe. Family Hypsilophodontidae. Upper Cretaceous. Ornithopsis Seeley. more or less spatulate. Suborder Ornithopoda Bipedal in habit. Front limbs always shorter than hind. Colorado. ORDER ORNITHISCHIA [Orthopoda. prepuhis. Antorbital openings small or absent. External nares near extremity of Postpubis complete. Vertebrae amphicoelous. more A. Femur longer than 1 [Belongs in the Diplodocidae. Bones very hollow. Vertebrae amphicoelous or amphiplatyan. Symphyrophus Cope. Aepysaurus Gervais.294 THE OSTEOLOGY OF THE REPTILES Hulke. Morinosaurus Sauvage. Premaxillae edentulous. Premaxillae rarely with teeth. divided. three sacrals. Ungual phalanges short digit. 18. the anterior ones often opisthocoelous. Teeth in a single row. Predentata] Quadrupedal or bipedal dinosaurs. ker. — Osborn and Mook. Lower Jurassic. Pubes not meeting in a median symphyand a posterior. and broad. underlying the ischium. never functionally pentadactylate. Europe. ? Astrodon Leidy. Barosaurus Marsh/ Caulodon Cope. Nanosaurus Marsh. compressed and pointed. Premaxillae with teeth.] 2 [Belongs in the Camarasauridae. South Africa. with an anterior. sometimes small. Family Nanosauridae.

Four or five sacral vertebrae. perhaps. with dermal armor of plates and spines. less dilated. Suborder Stegosauria [Quadrupedal. 295 Five sacral vertethe fifth vestigial. skull small. digits. Prosaurolophus Brown. Step hanosaurus Lambe. seven or Sub- Femur longer than tibia. bones solid. has been located plano-concave or with the Hypsilophodontidae. Gryposaurus Lambe. Kritosaurus Brown. forming a tessellated pavement in use. Manus with five.THE SUBCLASS DIAPSIDA tibia. Family Iguanodontidae. Jurassic to close of Cretaceous. the Laosauri- dae with platycoelous vertebrae.] Suborder Ceratopsia Secondarily quadrupedal dinosaurs. Laosaurus Marsh. (Hadrosauridae. find proper location in a distinct family. Claosaurus Marsh. c. Wealden). Hadrosaurus Leidy. Hypacrosaurus Brown. Family Trachodontidae. brae. though its teeth are unknown. England. three functional toes. Hypsilophodon Hulke. It is Its vertebrae are the latest of its known Ornithopoda and may eventually. and Iguanodontidae with opisthocoelous vertebrae. Teeth in a Anterior vertebrae platycoelous or opisthocoelous. Premaxillae edentulous. Trachodon Leidy. England. and the margin of a greatly extended "frill" or extension of the skull over . about Tail flattened. Upper Cretaceous. single row. postorbitals. No dermal ossifications. the Camptosauridae. located on nasal. Body covered with dermal ossifications. shorter than tibia. Corythrosaurus Brown. Extremity of face more or fifteen in Cervical vertebrae opisthocoelous. No MS. Premaxillae edentulous. The Scelosaurus. Femur longer or Four functional fingers. Lower Cretaceous (Morrison. Iguanodon Mantell. four functional fingers and three functional toes.) Teeth in many rows. with large skull. sclerotic plates in orbits. Wealden. but the differences seem to be of minor importance. number. armed with horns and protuberances. phalanges reduced. Cranium often with eight sacrals. aquatic in habit. Camptosaurus Marsh. Saurolophus Brown. Rocky Mts. B. crest. nearly amphiplatyan. Cheneosaurus Lambe. the pes with four. This family has been sometimes divided into three.

THE OSTEOLOGY OF THE REPTH^ES Lateral temporal openings small. but two carpaHa. Anchiceratops Brown. first tarsal Femora more or more or A. or in- Sternum large. Prepubes articulating with pelvis. ten or more dorsals. four to ten sacrals. First three fingers Hucarpus more for small. less less row fused with tibia. SUBORDER PTERODERMATA (RHAMPHORHYNCHOIDEA) Jaws with teeth. No teeth on premaxillae. Brachyceratops Gilmore. absent. the caudals amphicoelous. Antorbital opening distinct. with large lateral process. articulating with carpus. Free cervical ribs sometimes Tail long. the presacrals procoelous. Astragalus firmly united with tibia.296 the neck. a pteroid bone articulating with carpus. pneumatic skeleton. shorter than tibia. fifth toe vestigial. the first three or four cervicals coossiiied. Teeth with divided Vertebrae platy- roots in a single functional row. Western North America. the calcaneum free. Orbits with sclerotic plates. postacetabular process. present. posed of numerous vertebrae. terclavicle. Monoclonius Cope. parasternals present. Uppermost Cretaceous. . curved. Skull more or less pointed. Torosaurus Marsh. Neck elon- gate. fibula reduced or absent. Fibula present. ORDER PTEROSAURIA ossified. clavicles. Volant reptiles with highly elongated. than half the length of the forearm. Chasmosaurus Lambe. the external nares remote from the No parietal foramen. pentadactylate. Leptoceratops Brown. internal nares distinct. Ischium coelous. Prevomers and Metacarpals less Orbits large. the fifth toe reduced. feet long. and about twelve to forty caudals. Ceratops Marsh. or less reduced. Diceratops Lull. Carpus and tarsus reduced. the postpubis more or less vestigial. eight or nine cervicals. Sacrum comlUum with long preacetabular and slender. No supracoracoid foramen. merus shorter than forearm. lifth toe complete. well ossified. Triceratops Marsh. with claws. fifth digit fourth greatly elongated support of patagium. with a terminal dilatation. the upper jaws terminating in a distinct "rostral" bone. Agathaumas Cope. Eoceratops Lambe. Centrosaurus Lambe. Styracosaurus Lambe. tip. 19.

Skeleton oi Rhamphorhynchus (Pterosauria). i88.Fig. One third natural size. 297 .

Dorygnathus Orpel. Family Pterodactylidae. Upper ing. Pterodactylus Cuvier {Ornithocephalus SommerLy- Diopecephalus Seeley. All metacarpals articulating with carpus. Dermodactylus Marsh. Skull much long supraoccipital crest. First three metacarpals splint- Upper end of scapula articulating with notarium. Genera Incertae tell. Doratorhynchus Seeley. antorbital vacuity more Teeth in anterior part of jaws. Europe. but no occipital crest. fifth toe like. Nares and or less coalescent. Skull more elongate. Upper Cretaceous. Campylognathus Plieninger. Skeleton imperfectly known. very short. Eight-foot wing expanse. SUBORDER PTERODACTYLOIDEA longer or but little Wing metacarpal. . Jurassic. Lower Cretaceous. shorter than forearm. Kansas. antorbital opening. Cycnorhamphus Seeley. Smaller. and the upper end of scapula flat. A supra- Scapula with enlarged distal end articulating with notarium. fibula. No without phalanges. Family Pteranodontidae. Ditnorphodon Meyer. North America. Scaphognathus Wagner. Jurassic. A No Orbits small. Europe.298 THE OSTEOLOGY OF THE REPTH^ES Family Rhamphorhynchidae. Pre- pubes not band-Hke. Family Ornithocheiridae. occipital crest. Jaws with teeth in front. not articulating with notarium. Sedis. Cretaceous. Fifth toe more or less reduced. Anterior dorsal vertebrae not fused. Like the Pteranodontidae. Wyoming. England. toma (? Pteranodon) Seeley. Nyctosaurus {N yctodactylus) Marsh. Paleornis Man- England. Ornithodesmus Seeley. B. Wealden. Tail No cervical ribs. From twelve to twenty-five feet in expanse of wings. England. Rhamphorhynchus Meyer. elongated. Pteranodon Marsh. toothless. Family Nyctosauridae. Prepubes Ornithos- band-like. Ornithocheirus Seeley. Pterodracon dekker).

299 .Fig. One eighth natural size. Four thirds natural size. 190. Skeleton oi Nyctosaurus (Pterosauria). Fig. 189. Skeleton of P/(?ro(/(j<r/y/«j.

^ 30C .

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