LIBRARY UNIVERSITY OF CALIFOKNTAI Digitized by the Internet Archive in 2007 with funding from Microsoft Corporation http://www.archive.org/details/coalmeasuresamphOOmoodrich THE COAL MEASURES AMPHIBIA OF NORTH AMERICA By ROY LEE MOODIE Associate in Anatomy, University oj Illinois, Chicago Published by the Carnegie Institution of Washington WASHINGTON, 1916 LIBRARY UNIVERSITY OF CAL DAVIS CAKNEGIE INSTITUTION OF WASHINGTON Publication No. 238 Copies of this Book were first issued SEP 28 1916 PRESS OF J. B. LIPPINCOTT COMPANY PHILADELPHIA PREFATORY NOTE. The Carnegie Institution of Washington has already published several mono- graphs upon paleobiological subjects, written by its research associates, Hay, Wieland, and Case. Each author has dealt with the subject-matter of his particular field, but each has brought to bear upon his work common factors which have placed his labors upon a broader basis than the mere morphological descriptions of fossil forms of life. Case has published four monographs upon the morphology and taxonomy of the Permo-Carboniferous vertebrates of North America, and has followed these by a fifth, in which all the known factors bearing upon the development of the life were assembled in an effort to discuss the paleogeography of the period. In his conception paleogeography is a very broad term, involving not only a study of the distribution of land, water, and life in any one interval of time, but a consideration of all the factors in the extremely complex inter-rela- tions of organic and inorganic matter and causes which influence the development of each part. Geologists and paleobiologists have alike suffered in their interpretation of past conditions, because of their lack of knowledge of the work done by others. Stratigraphy may not be interpreted from the preserved fossils without a knowledge of biological laws, and the formations of the earth may not safely be rearranged to account for the present or past distribution of life without a knowledge of geo- logical processes. It is obvious that such work is beyond the possibilities of any one man; it is rather the work of a group of men, each broadly trained and each master of his own field and able to contribute to and criticize the work of his fellows. Nowhere could close cooperation of this kind be better accomplished than under a system such as the Carnegie Institution of Washington has developed, whereby the research associates of the Institution and others of its staff may call in the assistance of men in related fields. Already the value of this procedure is apparent in the results accomplished by cooperation. The following monograph, by Dr. Roy L. Moodie, adds an important link to the series of paleobiological publications of the Institution and is closely connected with the work already done upon the Permo-Carboniferous vertebrates, since it supplies a description of the life of the period immediately preceding. It is hoped that the volume will contribute in no small measure to an understanding of the broader problems of paleogeography and the recognition of the mutual problems of the paleobiologists and the geologists. E. C. Case. University of Michigan, March i$, igi6. PREFACE. The question of the origin of land vertebrates, which has appealed so strongly to students of fossil Amphibia, is by no means solved from the material furnished by the Coal Measures of North America. The Amphibia are, however, well known from several localities in the Coal Measures of this continent, where skeletons have been recovered which are sufficiently well preserved to afford a fair knowledge of their anatomy. The specimens rescued from the dumps of the coal mines are regret- tably few in comparison with the number that must have been burned as fuel, or carried down the slopes as silt. Yet scanty as is the material thus collected, it is of great importance, because it represents such an early period in the recorded history of the air-breathing vertebrates. The amphibian fauna in the Coal Measures of North America is represented by several hundred individual specimens, preserved in various museums. All of the collections have been available in the preparation of this memoir, with the exception of those species from Nova Scotia which are preserved in the Peter Redpath Museum of McGill University and in the British Museum of Natural History. The European material, which has been used in comparisons with the American forms, has been studied chiefly from the literature, although there have been available a series of specimens of Branchiosaurus amblystomus Credner, from Saxony, presented by the late Professor Credner, and a single specimen of Archegosaurus from Dr. von Huene, of Tubingen. The collection which has been of the greatest value is that at the American Museum of Natural History, chiefly assembled by Dr. J. S. Newberry from the dumps of the coal mines at Linton, Ohio, while he was in charge of the Ohio Geo- logical Survey (1869- 1884). This collection, a part of which is at Columbia Uni- versity, furnished Cope with the most of his type material for the "Synopsis of the Extinct Batrachia from the Coal Measures" (123).* This entire collection, including all of Professor Cope's types and representing many new and hitherto undescribed forms, was generously placed at the writer's disposal for a period of five years through the kindness of Dr. Bashford Dean and Dr. Louis Hussakof. Dr. Hussakof made a trip through the Linton region and his description of the place occupied by the "Old Diamond Mine" is given on page 16. An interesting collection of air-breathing vertebrates from the Coal Measures, representing 19 species, is in the U. S. National Museum (464). This is chiefly the collection of Mr. R. D. Lacoe and includes specimens from Mazon Creek, Illinois, from Kansas, and from Linton, Ohio. It is especially important in that it contains the skeleton (plate 20, fig. 3) of the oldest known reptile, Eosauravus copei Williston (Jour. Geol., xvi, 295). It contains also, besides many of Cope's types, new forms which have been described by the writer (464, 470, 471, 472, 473, 474, 478, 479). Dr. Stuart Weller first secured the use of this collection for me, and its continued use has been granted by Dr. C. D. Walcott. Mr. Charles Gilmore has called my atten- tion to several interesting specimens and has kindly loaned them for description. » The numbers in parentheses refer to the bibliography at the end of this volume. V VI PREFACE. A small but interesting collection of Mazon Creek Amphibia is that of the Pea- body Museum of Yale University. Through the courtesy of the officers of this museum the writer was permitted to study these specimens and was given a grant for their illustration. The results of that study are contained in a previous paper (478) and in the present memoir. Dr. Schuchert has offered suggestions as to the environmental conditions of the ancient Amphibia. A few specimens of Coal Measures Amphibia are at the Walker Museum, Uni- versity of Chicago. This collection includes the type of Micrerpeton caudatum Moodie, the first branchiosaur discovered in the western hemisphere, and a few specimens from Linton, Ohio. A single specimen of Amphibamus grandiceps Cope, very beautifully preserved, is in the possession of Mr. L. E. Daniel's, of Rolling Prairie, Indiana. This specimen has been studied and described by Hay (316) and by the writer (462, 469, 478). The works of Cope and Dawson, published between i860 and 1897, on the Amphibia from the Coal Measures, have been indispensable in the present study. It has been necessary to rely on the published descriptions and photographs of the interesting fauna from Nova Scotia, since it has not been possible for me to visit and examine the types preserved in the Peter Redpath Museum of McGill Uni- versity and in the British Museum of Natural History. It has been possible to check Dawson's work, to a certain extent, by a study of a series of excellent photo- graphs of the types of Coal Measures Amphibia collected by Dawson and Lyell and described by Dawson and Owen. The descriptions of these authors have been drawn on for the discussion of the Canadian forms. The descriptions given below have been made full and complete in the belief that in this way our knowledge of these interesting vertebrates may be advanced. Many of the species have been described elsewhere in scattered papers by various authors. These descriptions have been revised and verified and are collected here in monographic form. The work is a morphologic and taxonomic revision of the Amphibia from the Coal Measures of North America. Especial attention has been paid to the factors which have been most active in the evolution of the group, so far as these factors may be interpreted. It is the author's hope that this review may open up the field for many more workers, since we are just beginning to learn about the evolution of this group of vertebrates. The trustees of the Elizabeth Thompson Science Fund allotted a grant for the present investigation. This aid has enabled the writer to present his work in much better form than would have been possible otherwise. Dr. S. W. Williston has offered many suggestions and criticisms which have been gratefully adopted. It is with the greatest sense of pleasure that the author dedicates this memoir to his teacher and friend. After the manuscript was completed the author enjoyed a visit from Mr. D. M. S. Watson, of King's College, London, whose knowledge of the European and African forms enabled him to offer several very valuble suggestions. It is fitting also to express my indebtedness to the Carnegie Institution of Washington for the privilege of publishing my work in the series of monographs contributed by Dr. E. C. Case, dealing with the anatomy and relationships of the early land vertebrates of North America. Roy Lee Moodie. Prefatory Note Preface 9-22 23-36 CONTENTS. Page. iii \ vi List of Illustrations viii-x CHAP. I. The Prohi.em of the Amphibia from the Coal Measures ,-e II. History of the Discovery of Amphibia in the Coal Measures 6-8 III. Stratigraphic and Geographic Distribution of Amphibia in the Coal Measures of North America IV. The Morphology of the Coal Measures Amphibia V. The Amphibia of the Devonian and Mississippian of North America 37-38 VI. A History of the Classification of the Amphibia, with Especial Reference to the Species from the Coal Measures 39-45 VII. Classification of Amphibia Adopted in this Work, and a List of the Coal Measures Amphibia from North America 46-48 VIII. Definition of the Class Amphibia, the Subclass Euamphibia, and the Order Branchio- sauria 49-50 IX. The American Coal Measures Branchiosaurid/E 51-66 X. The Order Caudata 67-71 XI. The Order Salientia 72-74 XII. The Subclass Lepospondylia, the Order Microsauria, and the Group Aistopoda 75-77 XIII. The Microsaurian Family Hylonomid*, from the Coal Measures of Nova Scotia 78-84 XIV. The Microsaurian Family Tuditanid;e, from the Coal Measures of Ohio and Pennsylvania 85 in XV. The Mh kosaurian Family Stegopid/E, from the Coal Measures of Ohio 112-114 XVI. The Microsaurian Family Urocordylid.e, from the Coal Measures of Ohio 115 125 XVI I. The Microsaurian Family Amphibamid.e, from the Coal Measures of Ma/on Creek, Illinois 126-134 XVIII. The Microsaurian Family Nyraniid^, from the Coal Measures of Ohio 135-138 XIX. The Aistopodous Microsaurian Family Ptyoniid.e, from the Coal Measures of Ohio. . . 139-146 XX. The Microsaurian Family Molgophid^e, from the Coal Measures of Ohio and Mazon Crbbk, Illinois 147 154 XXI. The Microsaurian Family Sauropleurid/E, from the Coal Measures of Ohio 155-170 XXII. The Microsaurian Family Ichthycanthid.e, from the Coal Measures of Ohio 171-174 XXIII. Supposed Microsaurian Species of Uncertain Relationship 175-177 XXIV. The Temnospondylous Amphibia from the Coal Measures of North America 178-197 XXV. The Stereospondylous Amphibia from the Coal Measures of North America 198-201 Bibliography of the Fossil Amphibia, with Especial Reference to the Amphibia from the Coal Measures of North America 202-217 An Index to the Bibliography of Fossil Amphibia 218-219 Index 220-222 vii ILLUSTRATIONS. PLATES. Page. 1. Views along Mazon Creek, Illinois 12 2. Drawing of type specimen of Micrerpeton caudatum Moodie, from the Coal Measures of Mazon Creek. . 52 3. Specimens of Eumicrerpeton parvum, Erpetobrachium mazonensis, Erierpeton branchialis, Mazonerpeton longicaudatum, and Amphibamus grandiceps 58 4. (1 and 2) Vertebrae of Spondylerpeton spinatum Moodie. (3) Type specimen of Mazonerpeton costatuto Moodie. (4) Type skeleton of Cephalerpeton ventriarmatum Moodie. (5 and 6) The halves of the nodule containing a practically complete skeleton of Amphibamus grandiceps Cope 60 5. (1) A reconstruction of the Coal Measures branchiosaurian, Eumicrerpeton parvum Moodie, a small primitive salamander 64 (2) A restoration of the branchiosaurian, Mazonerpeton, based on two specimens 64 6. Dendrerpeton acadianum Owen. Mandibles, parts of anterior extremities, humerus, etc 68 7. Hylerpeton dawsoni Owen. Mandible, teeth, rib, and bones of anterior extremity. Bones of pelvis and posterior limb and bony scales 72 8. Fritschia curtidentata Dawson. Bones of skull and anterior extremity, bony rods of belly, of pelvis, and posterior extremity 76 9. Hylonomus lyelli Dawson. (1) maxillae and skull bones; (10) sternal bones; (2) mandible; (3) humerus, ribs, and vertebrae; (4) posterior limb; (5) pelvis; (6) caudal vertebrae 78 10. Hylonomus latidens Dawson. Skull, portion of skeleton, foot, scapular, and sternal bones, humerus and rib, believed to belong to this species. Erect tree, Coal formation, of Nova Scotia 80 11. Hylerpeton longidentatum Dawson. Mandible and other bones. Erect tree, Coal formation 82 12. Smilerpeton aciedentatum Dawson. Mandible, portions of skull, scales, and various bones. Erect tree, Coal formation 82 13. Dendrerpeton oweni Dawson. Skull, mandible, and bones of anterior limbs, posterior limb, pelvis, and bony scales 100 14. (1 and 2) Amphibamus grandiceps Cope, from the Mazon Creek shales 106 (3) Sauropleura (Colosteus) scutellata Newberry, from the Linton Coal Measures, the first known of the Ohio Coal Measures Amphibia; at first ascribed by Newberry to the fishes, but later correctly identi- fied by Cope 1 06 (4) Type of Diceratosaurus (Ccraterpeton) punctolineatus Cope, from the Linton Coal Measures 106 15. (1) Dorsum of skull of Diceratosaurus punctolineatus (Cope), from the Coal Measures of Linton, ( )hio. . 1 14 (2) Ventral surface of skull of Diceratosaurus punctolineatus (Cope), from the Coal Measures of Linton . . 114 (3) Pectoral girdle of Diceratosaurus punctolineatus (Cope), from the Coal Measures of Linton 114 (4) Cervical or anterior dorsal vertebra of Diceratosaurus punctolineatus (Cope), from the Linton Coal Measures 114 16. (1) Type specimen of Diceratosaurus punetolineatus Cope II.fi (2) Skull of Sauropleura longidentata Moodie, from the Coal Measures of Linton, Ohio 116 (3) Mandible of Sauropleura longidentata Moodie, from the Coal Measures of Linton, Ohio 116 (4) Type specimen of Sauropleura enchodus Cope, from the Coal Measures of Linton, Ohio 116 (5) Additional specimen of Diceratosaurus punctolineatus Cope, from the Coal Measures of Linton, Ohio. 1 16 17. Type of Saurerpeton lathithorax Cope 126 18. (1) Type of Erpetosaurus sculptilis Moodie, from the Cannelton Shales of Pennsylvania 132 (2) Skeletal elements of Eryops sp. indet., from the Pittsburgh Red Shale at Pitcairn 132 (3) Amphibian footprints, Dromopus aduncus Branson, from the Mississippian shales of Giles County, Virginia 1 32 (4) Type of Thinopus antiquus Marsh, amphibian footprint from the Devonian of Pennsylvania 132 19. Type of Ctenerpeton alveolatum Cope, from the Coal Measures of Ohio 134 20. (1) Skull of Erpetosaurus minutus Moodie, from the Cannelton slates of Pennsylvania 134 (2) Skull and anterior part of body of Ptyonius pectinatus Cope, from the Coal Measures of Linton 1 ^4 (3) Skeleton of Eosauravus copei Williston, from the Coal Measures of Linton. "The oldest known reptile from North America" and closely related structurally to the Microsauria 134 (4) Part of ventral scutellation and ribs of Sauropleura digitata Cope, from the Coal Measures of Linton . 134 viii »3- 24. 25- ILLUSTRATIONS. IX Mandible of Macrerpeton deani MocKlie, from the Linton Coal Measures 136 Portion of the skull of Macrerpeton deani Moodie, possibly of the same individual as the mandible. From the Linton Coal Measures 1 ie, Type of Cereariomorphus parvisquaniis Cope, from the Linton Coal Measures 136 An additional specimen of Cereariomorphus parvisquaniis Cope, from the Linton Coal Measures. . 136 Skull of Sauropleura scutellata Newberry. From the Coal Measures of Ohio 136 Tooth of Mastodonsaurus sp. indet. of the Carboniferous of Kansas 136 Tooth of Mastodonsaurus giganteus Jaeger, from the Triassic of Germany. Introduced for compar- ison with the tooth from the Kansas Carboniferous 136 Type of Leptophractus lineolatus Cope, from the Coal Measures of Linton. Portions of maxilla and mandible of left side with teeth 160 Type of Proterpeton gurleyi Moodie, from the Coal Measures of Illinois, near Danville. Cervical of an otherwise unknown amphibian 160 Amphibian phalanx from the Coal Measures near Breeze, Illinois, of an unknown species 160 Large rib of a stcrcospondylous stegocephalan, otherwise unknown 160 Type of Cope's species Tuditanus mordax referred by him to the cranium, on account of the sculp- turing of the elements, now known to be portions of the interclavicle and clavicles of Diceratosaurus punctolineatus 160 Skull of Baphetes planiceps Owen, from the Coal Measures of Nova Scotia 160 Ventral seutella; of Ctenerpeton alveolatum Cope, from the Coal Measures of Ohio 166 Left leg and pelvis of Ichthycanthus platypus Cope, from the Coal Measures of Ohio 166 Type specimen of Pelion lyelli Wyman, from the Coal Measures of Ohio. Supposed to represent the ancestral form of the Salientia • 172 Scales of Cereariomorphus parvisquamis Cope, a microsaur from the Ohio Coal Measures 172 Type specimen of Cereariomorphus parvisquamis Cope 172 Photograph of type specimen of Erpetosaurus (Tuditanus) radiatus Cope, from the Coal Measures of Linton 180 Photograph of type specimen of Erpetosaurus tabulatus Cope, from the Coal Measures of Linton. . . 180 Photograph of the impression of Stegops divaricata Cope, from the Coal Measures of Linton 180 Type and only known specimen of Micrerpeton caudatum Moodie, a branchiosaur from the Coal Measures shales of Mazon Creek 180 Type specimen of Erpetosaurus tuberculatus Moodie, from the Ohio Coal Measures 182 Type of Macrerpeton huxleyi Cope, from the Coal Measures of Ohio 182 TEXT-FIGURES. 1 . Map of the Coal Measures in North America 9 2. Distribution of Coal Measures Amphibia in North America 11 3. Topographical Map of Mazon Creek Region 13 4. Topographical Map of Linton, Ohio, Region 16 5. Fossil Tree Trunk in Position 21 6. Generalized Amphibian Skull 23 7. Alimentary Canal of a Carboniferous Salamander 26 8. Vertebra and Ribs of Coal Measures Amphibia 28 9. Ventral Scutellae of Micrerpeton 3° 10. Horny Armor of Hylonomus 3' n. The Skulls of two Microsaurians : A. Eoserpeton tenuicorne; B, Ceraterpeton galvani 33 12. Devonian Footprint 37 1 3. Restoration of Micrerpeton 53 14. Mazon Creek Amphibia: A, Eumicrerpeton parvum; B, Amphibamus thoracatus. 59 140. Skeleton of Mazonerpeton longicaudatum °2 146. Skeleton of Mazonerpeton costatum 64 15. A. Impression of Erierpeton branchialis °5 B. Eumicrerpeton parvum 65 C. Larger Specimen of Eumicrerpeton parvum (,5 D. Skeleton of Erpetobraehiutn mazonensis &3 E. Rib of Mazonerpeton costatum °5 15a. Type Material of Sparodus 6° 16. Obverse of Cocytinus gyrinoides ™ 16a. Nearly Complete Specimen of Cocytinus gyrinoides <*> 17. Pelion lyelli, supposed ancestral Salicntian "4 1 8. Skeletal Elements of Smilcrpeton aciedentatum ■* 26. X ILLUSTRATIONS. 19. Skull and Skeleton of Tuditanus punctulatus 87 20. Skull and Skeleton of Tuditanus longipes 90 2 1 . Skeleton of Tuditanus walcotti : A, Body ; B, Leg 94 22. A. Outline of Skull and Cranial Elements of Erpetosaurus minutus Moodie, from the Cannelton Slates of Pennsylvania 99 B. Outline of Skull and Cranial Elements of Erpetosaurus radiatus Cope, from the Coal Measures of Linton 99 C. Palate of Erpetosaurus (tabulatus?), from the Coal Measures of Linton, Ohio 99 D. Outline of Skull and Cranial Elements of Erpetosaurus acutirostris Moodie, from the Coal Measures of Linton, Ohio 99 E. Outline of Larger Part of Skeleton of Odonterpeton triangularis Moodie, from the Coal Measures of Linton, Ohio 99 F. Right Mandible of Erpetosaurus tabulatus Cope, from the Linton, Ohio, Coal Measures 99 G. Skull Elements and Lateral-line Canals of Erpetosaurus tabulatus Cope, from the Coal Measures of Linton 99 23. Skull of Stegops divaricata 113 24. Microsaurian Skulls from Linton, Ohio: A, Diceratosaurus lsevis; B, Diceratosaurus robustus 119 25. Restoration of Eoserpeton 1 24 26. Restoration of Amphibamus 128 27. Skeleton of Amphibamus grandiceps 129 28. Probable Appearance of Amphibamus 130 29. Skeleton of Cephalerpeton 133 30. Restoration of Ptyonius 140 31 . Restoration of ffistocephalus 144 32. Vertebrae of Molgophis brevicostatus 148 33. Fore-limb of a Member of the Molgophidas, Possibly Pleuroptyx 152 34. A. Interclavicle of Sauropleura pauciradiata 159 B. Left Clavicle of Sauropleura pauciradiata 1 59 35. Skull and Skeleton of Saurerpelon latithorax 164 36. Mandible of Leptophractus dentatus 169 37. So-called Interclavicle of Eury thorax subkevis 170 38. Skeletal Elements of Amblyodon 1 77 39. Vertebrae of Spondylerpeton spinatum 1 79 40. Mandible of Macrerpeton deani 184 41. Vertebras of Eosaurus acadianus: A, Oblique Lateral View; B, Oblique View; C, Posterior View; D, Trans- verse Section; E and F, Microscopic Sections 188 42. Skull and Mandible of Eobaphetes kansensis: A, Outer View of Mandible; B, Portion of Skull; C, Inner Surface of Mandible 191 43. Footprints of Dromopus agilis 200 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA CHAPTER I. THE PROBLEM OF THE AMPHIBIA FROM THE COAL MEASURES. The Amphibia from the Coal Measures of North America present the problem of the origin of the land vertebrates, since the air-breathing vertebrates in the Coal Measures of this continent are the earliest known in the western hemisphere. The difference in age between the chief amphibian-bearing deposits of North America and Europe is not great, although it has been asserted that Pholidogaster and its allied fauna, described by Huxley from Scotland (331), is much older, probably Mississippian. It is interesting to note that these earliest representatives of the Amphibia in Scotland are all temnospondyles, of which there are very few represen- tatives in the Coal Measures of North America. The forms so far described from the North American Coal Measures present a very high degree of development and differentiation, the earliest known species being already specialized and well adapted for various modes of life. As far back in geological time as the middle Coal Measures, when the first well-defined forms are known, environmental conditions had effected a wide diversity of structure within the group. Thus, early in the geological history of the land vertebrates, we have, among the Coal Measures Amphibia, various forms which had specialized into strictly aquatic, terrestrial, subterrestrial, and arboreal, or at least partly arboreal. Specialization had extended to the loss of limbs, ribs, and ventral armature in a few species, and to the acquirement of claws, running legs, or a long propelling tail with expanded neural and haemal arches in others. The forms range in size from small creatures less than an inch in length to large species which must have attained a length of several feet. A rather interesting parallel, though of no phy- logenetic significance, can be drawn between the Amphibia of the North American Coal Measures and the reptiles of to-day. The snakes are represented by the limb- less, snake-like forms, such as Ptyonius and Phlegethontia. The lizards find their counterpart in the Hylonomidae and the Tuditanida?. No known characters of these animals tend to ally them directly with any known group of fishes, except in the most general way. These facts all indicate a long antecedent history for the amphibian group or else a preceding period of greatly accelerated development of which we now know nothing. The Amphibia whose remains have been brought to light from the Coal Measures have hitherto been regarded as pertaining to a single order, the Stegocephalia, characterized by the' completely roofed-over cranium and a large parasphenoid. The writer (469) had previously assigned 5 suborders to the group : the Branchio- sauria, Microsauria, Aistopoda, Temnospondylia, and Stereospondylia. All of these groups are represented in the Coal Measures of North America. It has seemed inadvisable, in the light of our present knowledge of the Amphibia, to retain these 5 groups as suborders, and, in the revised scheme of classification which has been 3 4 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. published elsewhere (469), they are given the rank of orders — all excepting the Aistopoda, which are now regarded by the writer as specialized Microsauria. The recent Caudata are possibly represented in the North American Coal Meas- ures by forms which may be assigned tentatively to the Proteida. Such forms as Cocytinus gyrinoides, Hyphasma loevis, and Erierpeton branchialis possibly represent this group in the Pennsylvanian. This relationship is based chiefly on the structure of the hyobranchial apparatus and on the general structure of the species. The three above-mentioned species are, however, very insufficiently known, and the relationship can hardly be regarded as more than suggested by the characters which are at hand. The Salientia, or frogs, may possibly have their ancestral type in Pelion lyelli, the first known species from the Linton, Ohio, Coal Measures. Oddly enough, among the hundreds of specimens collected later from this horizon, not a fragment can be identified with this species. The type specimen is unique, and although incomplete its characters are suggestive. The Branchiosauria are represented in North America by four species: Mic- rerpeton caudatum, Eumicrerpeton parvum, Mazonerpeton longicaudatiim, and M. costatum. Three other genera which occur in North America have been placed (642) in this group, but they do not belong there, for reasons given below. The branchiosaurs were salamander-like in appearance. They were naked, with the exception of small ovoid scales on the back and the chevron-shaped armature of the ventral surface, the latter being almost universally present among the Paleozoic Amphibia. They were adapted for life in the water for at least the early part of their existence, as is shown by the possession of gills on many of the late Carboniferous and early Permian forms of Europe. The group is, without doubt, ancestral to the modern Caudata. No branchiosaurians have been described elsewhere from so low in the geological series as those here given and they are the first and only evidence of the occurrence of the group in the western hemisphere. The Microsauria are represented in the Coal Measures by numerous forms which are usually characterized as lizard-like animals with a well-developed ventral scu- tellation. Other characters, such as the possession of lateral -line grooves on the cranium, the arrangement of the cranial elements, and the condition of the ribs, will be discussed further on. The pectoral arch is well developed and is composed of five dermal bones plus the regular skeletal elements. The skeletal membrane bones are sculptured after the manner of those of the cranium. The bodies of the animals were, in a few cases, covered with scales; but most of them appear to have been completely naked, even the ventral armature being absent in some cases. The ventral scutellation was especially strong and highly developed in some of the forms; e.g., in the genera Saarerpeton and Sauropleura. The vertebrae are uniformly of the hour-glass or notochordal type. This is so generally the case that the characters of the vertebrae and ribs are taken as the chief diagnostic characters of the major groups. Various peculiarities are seen among the Microsauria, such as the development of horns in various genera which are, apparently, related. The order seems to have gone completely out of existence during the early Permian, PROBLEM OF THE AMPHIBIA FROM THE COAL MEASURES. 5 and if their descendants continued on as reptiles, as has been suggested (469), we do not know the intermediate stages. The Aistopoda are without doubt specialized microsaurs, and, in the opinion of the writer, are not entitled to separate rank. Some of these forms reached a high degree of specialization. One American species has the skeleton reduced to a long, slender head and a slender series of elongate vertebrae, all other parts of the skeleton, even the ventral armature, being absent. The proportions attained by this species, Phlegetliontia linearis Cope, recall those of the coach-whip snake, Zamenis flagellum Shaw, of the western plains. Some of the so-called Aistopoda have been credited by Fritsch with the possession of peculiar clasping organs, "Kammplatten." Newberry has written of the discovery of similar structures in the Ohio Coal Measures (498), but the statement of the actual association of these "Kammplatten ' ' needs confirmation. Dr. R. H. Traquair wrote to the author under date of April 28, 1909: "I maintain that the association of a bundle of 'Kammplatten' with a specimen of Ophi- derpeton in the Bohemian gas coal was entirely accidental. Of such pitfalls the paleontolo- gist has to beware or serious mistakes may be the consequence, as has happened more than once. I must, however, publish a short paper on the Kammplatten, for I think I know what they are now." Fritsch, however, has very clearly figured a nearly complete specimen of Ophi- derpeton (251, Bd. iv) as possessing the Kammplatten in place near the cloaca, where he suggests they may have served the function of accessory copulatory organs or claspers. The Temnospondylia are represented by scanty remains of species from Illinois, Pennsylvania, and Nova Scotia. The forms belonging to this group are all rela- tively large, and they had a wide geographical distribution during the Permian. This group contains two types of vertebras, known as the embolomerous and the rachitomous, both of which are present in the Coal Measures. Such forms as Eosa urns, Baphetes, Eobaphetes kansensis, Macrerpeton, and Dendrerpeton are regarded tentatively as temnospondyles, but there is no definite assurance that they are such. It is possible that Eosaurus is a stereospondyle, but the species is too incompletely known for a definite statement to be made. The close resemblance between the vertebras of Eosaurus and Anthracosaurus has been noted by Huxley (332). The Stereospondylia are very scantily represented in the Coal Measures, if at all. Eosaurus may belong here as indicated above. The tooth and cranial frag- ments discovered and described by Williston from the Coal Measures of Kansas may represent a stereospondyle as he states (608), but the evidence is incomplete. A fragment of a large rib (plate 22, fig. 4) of a species from Linton, Ohio, otherwise unknown, may be a stereospondyle. We would expect an early development for this group, but it is an interesting fact that no stereospondyles are known defi- nitely before the Triassic, during which period they had an extensive distribution. CHAPTER II. HISTORY OF THE DISCOVERY OF AMPHIBIA IN THE COAL MEASURES. Sir William Logan, in 1841, discovered in the Coal Measures of Horton's Bluff, Nova Scotia, some tracks of Amphibia which he carried to London and which Sir Richard Owen pronounced to be undoubted "reptilian" tracks. This fact was published in 1842 (380) and was the first recorded evidence of the occurrence of land vertebrates in the Carboniferous rocks of the world. To these tracks Sir William Dawson later gave the name of Hylopus logani. Two years later Dr. Gergens (291) wrote a letter to Professor Bronn, the founder and one of the editors of the "Neues Jahrbuch fur Mineralogie, Geologie und Pale- ontologie," in regard to an important discovery in the Carboniferous rocks of Ger- many. The letter is of such exceptional interest in connection with the history of the fossil Amphibia that it is given here : "In dem Brandschiefer von Munsterappel in Rhein-Bayern habe ich in vorigen Jahre einen Salamander aufgefunden und Hrn. H. v. Meyer in Frankfurt zur naheren Unter- suchung und Beschreibung tibergeben; — Gehort dieser Schiefer der Kohlen-Formation ? — in diesem Falle ware der Fund in anderen Hinsicht interessant." The form discovered by Dr. Gergens and described by Hermann von Meyer as an amphibian is a little puzzling as to its characters. Miall (449, p. 183) says that the remains are too imperfect for close definition. The form, as figured, resembles an immature branchiosaurian, as one is at once reminded, from an examination of Von Ammon's Branchiosaurus caducas (7, Taf. iv, fig. 1). In 1844 Dr. Alfred King (356) announced the discovery of "reptilian" footprints in the Carboniferous of Pennsylvania. The next announcement of fossil Amphibia was made by Goldfuss (296), who in 1847 described the famous Archegosaurus from the upper Carboniferous of Ger- many, from the remains which had as long ago as 1777 been regarded as a fish. Two years later Isaac Lea (371) announced to the British Association for the Advance- ment of Science, through Buckland, the discovery of footprints in the old Red Sandstone (Mauch Chunk) of Pennsylvania. These objects occur not rarely in the Mauch Chunk shales, which are of upper Mississippian age. Barrell (21, p. 460) records the finding of imperfect tracks in the same beds, and Rogers (Geology of Pennsylvania, pt. 11, 1856, p. 831) records three unnamed varieties from 2,200 feet below the top of the Mauch Chunk. Branson (50) has recorded the finding of other amphibian footprints from the Mississippian of Giles County, Virginia. Lyell and Dawson (396), in 1853, read a paper before the Geological Society of London, in which they announced the discovery of remains of Amphibia in the Coal Measures of North America, although Dawson had previously, in 1850, discovered the skull of Baphetes planiceps Owen, which was not described until the latter part of 1853 (509). The specimen had lain unnoticed in the collection of the Geological Society for more than two years. When, however, the announcement was made 6 HISTORY OF DISCOVERY. 7 by Lyell and Dawson of the discovery of Amphibia in the Coal Measures of Nova Scotia, so much interest was excited that the skull, now known as Baphetes plani- ceps, was brought to light by the president or secretary and was described (509) by Sir Richard Owen. The only other known evidences of land vertebrates in the Paleozoic of North America, up to this time, had been the footprints described by Lea and King from the Mississippian (Mauch Chunk) and Pennsylvanian of Penn- sylvania. The specimens presented to the Geological Society of London by Lyell and Dawson were found at the South Joggins, Nova Scotia, and consisted of scutes, a few limb bones, a fragment of a jaw, and a few vertebrae, a part of which were associated. The remains were found quite accidentally and unexpectedly by them in the petrified trunks of ancient Sigillariae which were exposed on the coast. Dr. Jeffries Wyman, of Harvard College, had examined these remains in the United States and had pronounced (638) them to be amphibian, comparing them with similar elements in Menobranehus. On the arrival of the specimens in England they were submitted to Sir Richard Owen, who suggested the name (514) Dendrerpeton acadianum and compared the remains with Archegosaurus. At the same meeting of the London Geological Society, Owen read a paper on a small amphibian (508) from the British Carboniferous which he named Parabatrachus. Subsequent dis- coveries have shown, however, that this form belongs among the fishes. At the meeting of the Geological Society held in the latter part of the same year Owen announced (509) further discoveries in the Nova Scotia coal beds. Hermann von Meyer (436), in 1857, described numerous stegocephalian remains from the upper Carboniferous of Germany. Dr. Jeffries Wyman, in the same year, described (639) a new form of amphibian from Linton, Ohio. This form he called Raniceps lyelli, but as the name Raniceps had been preoccupied by Cuvier for a genus of gadid fishes, Wyman later (1868) changed the name to Pelion. This was the first form to be described from the locality at Linton, which has since yielded the remains of half a hundred species. Dawson (204), in 1859, made a further contribution to the fauna of Nova Scotia by the description of Hylonomus and other species of Dendrerpeton from the South Joggins deposits. Huxley (331), in 1862, described the genera Loxomma and Plioli- dogastcr from the Carboniferous of Scotland. The same year Owen made a further contribution (514) to the fauna of the Nova Scotia beds, and Huxley (332) discussed the anatomy of Anthracosaurus from Scotland. Marsh (404), in the next year, described, as an enaliosaurian, the interesting Eosaurus acadianus from the Nova Scotia Coal Measures, basing the species on two vertebrae, apparently from the dorsal region. The vertebrae resemble the stereospondylous type, and Huxley (332) called attention to the similarity of these vertebrae to those of Anthracosaurus. Cope (105), in 1865, began his researches among the Coal Measures Amphibia of North America by the description of Amphibamus grandiceps from the Mazon Creek shales of Illinois. Ten years later (123) he published a complete synopsis of the Carboniferous Amphibia of North America, with especial reference to the Linton, Ohio, species, illustrating many of the forms now known from Linton. Between the years 1865 and 1897, Cope published numerous papers (105-177) on the Amphibia 8 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. of the Paleozoic, and to his researches is due a large part of our knowledge of these forms. Great credit is due Dr. J. S. Newberry (495, 498) for the enthusiasm and interest which his collections of Coal Measures Amphibia exhibit. He furnished Cope with the majority of the type material described by him, and it was through Dr. New- berry's instrumentality that the "Synopsis of the Extinct Batrachia from the Coal Measures" (123) was published. The material which Dr. Newberry had collected he took with him from Ohio to Columbia University, New York, and a part of his collection still remains in the geological collection of that institution, although the greater portion has been transferred to the American Museum of Natural History. The Newberry collection forms the basis for the larger part of this memoir. Between the year 1853 and the early nineties, Dawson continued (200-223) his researches on the Amphibia of the Coal Measures of Nova Scotia. His most notable single work (208) is "The Air-Breathers of the Coal Period," published in Montreal in 1863, in which he gives a complete account of the forms then known from Canada, attempting some restorations. Since his death there have been no new species described from Canada, and, so far as I can learn, there has been no further collecting at the South Joggins. Recently G. F. Matthew (409) has rearranged the classification of amphibian footprints from Nova Scotia. Jaekel (347) has described very fully the remains of Diceratosaurus punctolineatus (Cope) from Linton, Ohio, basing the new genus on a species described by Cope as a member of Ceraterpeton. Hay (316) has added to the knowledge of the anatomy of Amphibamus, his most interesting contribution being the detection of long, curved ribs in this form. This character excludes the species from the order Branchiosauria and shows the relationship of the form to the Hylonomidae and the Microsauria. Schwarz (540) has described the characters of the vertebrae and ribs of several genera of the Coal Measures Amphibia and has (541) offered his views as to the descent of the Amphibia, based entirely on his work on the vertebras of species from North America and Europe. Since 1908 the writer has published several contributions (457-489) on the Amphibia from the Coal Measures of North America. The results of these investi- gations are given in this work. CHAPTER STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION OF AMPHIBIA IN THE COAL MEASURES OF NORTH AMERICA. There are but four localities in North America which have furnished any notable remains of Amphibia in the Coal Measures. These are, in the order of their dis- covery, the deposits at the South Joggins, Nova Scotia; the Linton, Ohio, Coal Meas- ures; the Mazon Creek, Illinois, shales; and the Cannelton slates near Cannelton, Pennsylvania. There are, however, several other localities on the continent which have furnished evidences of Amphibia in the Coal Measures. The principal one of the latter localities is doubtfully of Coal Measures age, although recent discoveries would tend to show it is such. The deposits in question, those of the Clepsydrops shales of Vermilion County, Illinois, have, heretofore, been regarded as Permian, but the discovery of similar remains in rocks of undoubted Pennsylvanian age in Pennsylvania would seem to indicate that the Illinois deposits were contemporaneous with them. (a) The deposits in Vermilion County, Illinois, lie along the north bank of Salt Fork Creek, at the tip of the "Horseshoe Bend," about 2 miles south of Oakwood, Illinois. They were discovered by Dr. J. C. Winslow, of Danville, in 1 875. The remains discovered by him were forwarded to Pro- fessor Cope for identification. Later the deposits were thoroughly explored by W. F. E. Gurley, and the specimens collected by him are now preserved (86) in Walker Mu- seum, University of Chicago. In 1907, the writer, while working for the University of Chicago, in exploring the same locality, exhausted the beds so far as they could at that time be uncovered from the landslide which had overwhelmed them. The for- mation in which the bones occur is a soft gray or reddish shale, and it lies without any ap- parent stratigraphic break on shales of Penn- sylvanian age. Below these shales are sev- eral feet of limestone containing invertebrates of typical Pennsylvanian fades. There are indications of at least 3 species of Amphibia in the deposits. Case (86) has in- dicated with doubt a fourth species. The species are: Cricotus Iwteroclitus Cope, 9 Pig. i.— Map of Upper Pennsylvania!) showing land and water conditions under which the Coal Measures amphibian fauna lived. It will be noted that the chief deposits which have furnished amphibian re- mains are on the margins of the heavily shaded areas. (After Schuchert.) Explanation of symbols: Lands are white. Water areas are lined. Formation outcrops are black or dotted. Known shore-lines are solid lines; probable ones broken. Vertical lines in middle of continent incii- cate Gulf marine. 10 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Cricotus gibsonii Cope, Diplocaulus salamandroides Cope. The remains are very fragmentary, and consist for the most part of incomplete vertebrae, with a few small skull fragments. (b) In 1897 Dr. Williston (607) described some fragments of Cricotus from a deposit in Cowley County, near Winfield, Kansas. There has been some dispute as to the age of the deposit, but the consensus of opinion seems to be that the beds are of approximately the same age as those of Illinois and Pennsylvania in which similar remains are found, and those deposits are looked upon as Upper Pennsylvanic (Case (94), pp. 239-240). No new forms were described from Winfield, since only a few fragments were obtained. Williston referred the phalange, the fragment of a jaw, and the tooth to Cricotus heteroclitus Cope. (c) Later in the same year Williston (608) announced the discovery of a tooth of typical labyrinthodont structure from near Louisville, Kansas (plate 21, fig. 6). The tooth was accompanied by fragments of bone and was probably not far from the bed in which it was fossilized. Williston states that the remains were from the shales which are "nearly at the upper part of the Carboniferous, probably within one hundred feet of the Manhattan Limestone." (d) In 1894 Marsh (406) and earlier (1873) Mudge (490) described footprints of vertebrates from the stone-quarries near Osage City, Kansas. The stone in which they were found was a fine-grained limestone which occurs near the middle of the Kansas Coal Measures. (e) Two years later Marsh (407) announced the discovery of traces of the oldest known (Devonian) air-breathing vertebrate. The footprints of Thin opus aiitiuuuswere regarded by Marsh as "apparently amphibian." This still remains the oldest geo- logical evidence of air-breathing vertebrates, although Lohest some years ago (381) called attention to remains from the Devonian of France which he thought might be amphibian. The footprint described by Professor Marsh was "found in the town of Pleasant, one mile south of the Allegheny River, Warren County, Pennsylvania, by Dr. Charles E. Beecher, who presented it to Yale Museum, and also furnished the information in regard to its geological position. , . . The geological horizon is near the top of the Chemung, in the upper Devonian. In the same beds are ripple marks, mud cracks, and impressions of rain drops, indicating shallow water and shore deposits." (f) Among the collections of the American Museum there is an impression of a small amphibian foot obtained from Phoenix Tunnel, Pennsylvania. The impres- sion is in hard black slate very similar to the slate of the Cannelton region. It is possible that the specimen may have been obtained from the Cannelton beds, since they would be expected to occur at Phoenix Tunnel. The impression is rather small. It is the footprint of a 5-toed animal, probably of the right foot, since no amphibian (465) so far is known from the Coal Measures with 5 digits on the hand. The first digit is short and thick, with a large ball at its base. The foot measures from the posterior edge of the palm to the tip of the longest digit 12 mm. The length of the first digit is 7 mm. The impression differs in some respects from the impressions so far known from the Coal Measures, but no attempt will be made to assign it to a STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. I I species. It may have been made by either a branchiosaurian or a microsaurian, but more probably the latter, since we do not know of any of the former animals from the Cannclton beds, or in fact from any of the Coal Measures beds excepting the Mazon Creek shales. The specimen is No. 2872 of the American Museum. 2. 3- 4- 5- Fie;. 2. — Distribution of Coal Measures Amphibia in North America. I. Linton, Ohio, near Yellow Creek P. O., Jefferson County, Ohio, on the banks of Yellow Creek, near the Ohio River, 16 miles north of Steubenville. Mazon Creek shales, Grundy County, Illinois, near Morris. "Clepsydrops shales," Salt Fork Creek, Vermilion County, Illinois, near Oakwood, on Tate farm, 8 miles west of Danville, Illinois. Danville, Illinois, coal where the type of Prolerpelon gurleyi Moodie was found. Breeze, Illinois, where Dr. J. A. Udden, in 1907, found a fragment of an amphibian phalange on the dump of the Cooper- ative Coal Company. 6. Pitcaim, Pennsylvania, 15 miles east of Pittsburgh. 7. Cannelton, Pennsylvania, Beaver County, Cannelton slates, Kittanning formation, 45 miles northwest of Pittsburgh. 8. Fairfield, Iowa, where Dr. J. A. Udden found remains attributed by Dr. Eastman to Pkuroptyx davatus Cope. 9. Louisville, Pottawatomie County, Kansas, where Dr. S. W. Williston discovered remains at Maslodonsaurus in the Coal Measures. Washington County, Kansas, source of type of Eobapheles kansensis Moodie, from the Coal Measures. Osage City, Osage County, Kansas, amphibian footprints from the Coal Measures. Winfield, Kansas, source of Cricotus material. Lander, Wyoming, in Wind River Carboniferous. Pictou, Pictou County, Nova Scotia, 84 miles northeast of Halifax. Source of Baphetes planiceps Owen. Joggins (Joggins Mines), Cumberland County, Nova Scotia, 4 miles from River Hebert. Source of HyUrpcton and Den- drerpeton faunas. 16. South Joggins, Nova Scotia, source of the Eosaurus acadianus Marsh. 10. 11. 12. 13- 14. 15. (g) Dr. J. A. Udden, in 1907, discovered a fragment of a phalanx of some amphibian (plate 22, fig. 3) on the dump of the Cooperative Coal Company, a mile east of Breeze, Illinois. It was obtained from below the Shoal Creek limestone and somewhere above the (Illinois) Coal No. 6, according to Dr. Udden's notes. The maximum width of the phalanx is 10 mm. and it probably had a length of 16 mm. 12 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. (h) Mr. N. H. Brown, in 191 4, discovered in the Carboniferous rocks to the east of the Wind River Mountains, near Lander, Wyoming, a single fragment of an amphibian. The writer was accompanying Mr. Brown at the time of the discovery and there can be no doubt that the fragment was amphibian ; the location of the beds was such that no later age than the Coal Measures can be assigned to them. (i) Dr. J. A. Udden (577), in 1912, announced the discovery of an amphibian in the Des Moines formation of Iowa. These remains were identified by Dr. Charles Eastman as Plenroptyx clavatus Cope. Since the Des Moines is probably nearly contemporaneous with the Mazon Creek shales of Illinois, the discovery does not extend the geological range to any extent, but is of interest as it adds another note to our knowledge of the geographical distribution of the Amphibia in the Coal Measures. (j) The Gurley collection of the University of Chicago possesses a single cervical vertebra of some amphibian ( ?) . The vertebra is unlike anything previously described and represents a new form (plate 22, fig. 2) which may be designated Proterpeton gurleyi, new genus and species. The material was collected near Danville, Illinois. (k) Deposits have been discovered in Pennsylvania in which are found the remains of amphibians and reptiles, very similar to those from Vermilion County, Illinois, Cowley County, Kansas, and the Texas Permian. The remains (plate 18, fig. 2) were found in a thin stratum below the "Ames" limestone, and are therefore in the Coal Measures, fairly well below the top. The fossils, as described by Case (94) , consist of fragments which he ascribes to pelycosaurian reptiles and to temno- spondylous amphibians. The genus Eryops (94) is recognized in several fragments and a nearly complete dorsal vertebral centrum. Other types of Amphibia are like- wise represented. (I) The ironstone nodules, in which the Mazon Creek fossils (plate 1) occur, are found in the shale which forms the roof of the Morris or "No. 2" Coal of Illinois, which "lies probably somewhat lower than the horizon of the Lower Kittanning Coal of Pennsylvania" (599). "The nodules of iron contained in the Coal shales on the banks of Mazon Creek near Morris, Illinois, generally contain organic nuclei, and thousands of beautiful specimens have been obtained there. They are usually fragments of fern fronds, but are sometimes shells, crustaceans, myriapods, scorpions, spiders, cockroaches, . . . fishes" (498, p. 214), and amphibians, of which 10 species are at present known. These species have been arranged zodlogically according to the following plan : Class Amphibia Linne', 1758. Subclass Euamphibia Moodie, 1909. Order Branchiosauria Lydekker, 1889. Family Branchiosanridce Fritsch, 1879. Micrerpeton caudatum Moodie, 1909. Eumicrerpeton panmm Moodie, 1910. Mazonerpeton longicaudatum Moodie, 191 2. Mazonerpeton costatum Moodie, 191 2. Order Caudata Dum£ril, 1806. Suborder Proteida Cope. Family Cocytinidw Moodie, 191 2. Erierpeton branchialis Moodie, 191 2. MOODIE VIEWS ALONG MAZON CREEK, ILLINOIS. 1. A nodule weathering out of the shale, at the head of the hammer. Most of the nodules at the so-called "lower beds" contain specimens of Xeuropteris. I. The nodules in the creek bed at the "upper beds." Many of them have been cracked open by the frost. 3. Looking south at the "upper beds." The nodules found in the background are non-fossiliferous. 4. Nodules may be seen through the clear water embedded in the shale. Neuropterid insects in the water. 5. Looking for nodules at the "upper beds." The uppermost reaches of the .ossiferous beds correspond with the extreme background of the picture. Nodules in the stream bed at the "lower beds." Many of these a,e cracked open by the frost and good specimens are sometimes found in the noduU>. S. STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. 1 3 Class Amphibia Linne, 1758— Continued. Subclass Lepospondylia Zittel, 1887. Order Microsauria Dawson, 1863. Family Amphibamidce Cope, 1875. Amphibamus grandiceps Cope, 1865. Amphibamus thoracatus Moodie, 191 1. Cephalerpeton ventriarmatum Moodie, 19 12. Family Molgophidm Cope, 1875. Erpetobrachium mazonensis Moodie, 191 2. Subclass Stegocephala Cope, 1868. Order Temnospondylia Zittel, 1887. Suborder Embolomeri Cope, 1885. Family Cricotidm Cope, 1884. Spondylerpeton spinatum Moodie, 19 12. It will be seen from the above arrangement that nearly all of the orders of Amphibia are represented in the Mazon Creek fauna. These animals are the oldest known land vertebrates of North America. Contour interval SOfeet Fig. 3. — Portion of the "Morris sheet" of the U. S. Geological Survey, to show topography and situations of the exposures of fossil-bearing shales at Mazon River, a, the "Bartlett place," the so-called "upper beds"; b, "lower beds." The writer was able, during July 191 1, to spend a week studying the fossil beds (479) at Mazon Creek. The object of the visit was primarily to collect Amphibia, but although several thousand nodules were examined, not one contained an amphibian nor a fragment of one. Mr. J. C. Carr, of Morris, Illinois, who has collected at Mazon Creek for more than 30 years, has never collected an amphibian. These facts interested me in making the following comparison: If we take 100,000 nodules as a basis for computation of the rarity of the various forms, something like the following will be the approximate result of the investigation : 14 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Of 100,000 nodules, 20,000 will be barren or contain only indeterminate fragments ; 68,500 will contain plants; 7,500 will contain insects, Crustacea, myriapods, scor- pions, spiders, and other arthropods; 3,900 will contain fish coprolites or scales; 95 may contain fish or fragments of fish ; 4 may contain mollusks ; and 1 may contain an amphibian or a fragment of one. Perhaps even 100,000 is low as a basis of estimate. Mr. Carr was of the opinion that 1 nodule in every 500,000 might contain an amphibian. The beds from which the nodules are usually collected occur along both banks and in the bottom of the creek, in two localities. One locality known as the Bartlett place is situated 8 miles southeast of Morris, in Grundy County, Illinois, Wauponsee Township, N.W. quarter, section 30, Township 33, Range 8, the land being now owned by Mrs. Emma Akerly, of Wilmington, Illinois. The fossil-bearing nodules occur throughout 6 to 8 feet of shale along both banks of the creek at the "upper beds" (plate 1 , fig. 3) , as the Bartlett place is called. They may also be seen in the bed of the creek, when the water is low (plate 1, fig. 4), still embedded in the shale. With a potato fork the shale is easily turned and the nodules come out like potatoes. One sometimes finds a "pocket" of nodules from which as many as a peck may be secured. Nearly every nodule has a fossil at the "upper beds," but all of the fossils are not well preserved, possibly only 1 or 2 out of every 10 being worth carrying to the museum. The nodules crack best when wet, and it requires some skill to crack them evenly. They seem quite light and, in one place where the stream curves, are piled in a long windrow. On this were found, in nodules cracked by the frost, several good crustaceans and many good plants. Table of Pennsylvanic Formations. gerjc Northern Appalachian. Bituminous, Pennsylvania-Ohio. Illinois. Monongahela or Upper Productive Coal Measures {Break"" Fork'i- Conemaugh or Lower Barren Ames limestone near middle: (*) (Pitcairn) Coal No. 6. . — . I Kreeport_ Allegheny or Lower Coal Measures t c 3 w o « c £ > >>< i o c 0 c (U H< i w 1) £3 £*| (*) (Cannellon) Coal No. 2 (*). Kittanning (*). (Morris?) (Mazon Creek). (Linton). Clarion. Homewood. Mercer. Conoquenessing. Sharon. (*) marks the position of the Amphibian-producing horizons in these regions. (After Schuchert.) The fossils at the "upper beds" are localized into special strata. At one place in the upper part of the deposit, in a reddish shale, one finds that insects are more abundant than they are lower down. The Crustacea seem to come from appar- ently the same shale. At the lower end of the deposit certain definite species of Pecopteris are localized. It is an interesting fact that one seldom finds a Neurop- teris at the "upper beds." The most abundant fossils are the various species of Pecopteris and Aiimilariu. When specimens of Neuropteris are found they are usually discovered at the lower end of the exposures. In one place behind the "island" very blue nodules, hard and flinty and with sometimes well-preserved STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. 1 5 specimens of Pecopteris, are found quite definitely localized. These nodules are apt to assume an irregular shape. These localizations of the fossils are, of course, what we would expect from our knowledge of the recent fauna and flora. There is, to be sure, more or less intermingling of the species. The myriapods, so far as they have been found, are also localized. Mr. Carr found 3 within a space of a few feet, but again these are found widely scattered. The exposures at the "upper beds" are about a quarter of a mile long. They disappear under a heavy ledge of sandy limestone. At the ' ' lower beds' ' (plate 1 , fig. 6) , those further down the creek, conditions are quite different from those just described, although of the same horizon; the banks of the creek are higher and almost perpendicular, so that the chances of collection from the shales are fewer. The bed of the creek, however, is wider and there are more nodules washed out. The most abundant fossil at this place is Neuropteris. The nodules at the upper end of the exposure are all, almost without exception, barren of fossils. The exposures here are of about the same extent as the "upper beds," though the species are not so varied. Judging from the collections made while there, Arthropoda are more abundant at the "lower beds." Bradley (Geol. Surv. Illinois, iv, 196, 1870) mentions the occurrence of these nodules at or near Morris. Other than these places the nodules have been thrown out of a coal mine near Braidwood, Illinois. Doubtless close search would reveal other localities where the shale is cut through in mining. The beds at both places are slightly folded. This is true especially of the ' ' upper beds," where a conspicuous fold caused the beds to disappear in the bed of the creek and to reappear farther down stream. This is directly across the large "ox-bow" bend of the creek. The beds at Mazon Creek were first explored in 1857 by Mr. Joseph Evans, who sent his specimens to Berlin, Germany, where they excited great interest. It was he who collected the type specimen of Amphibatnus grandiceps Cope. Since the time of Mr. Evans many have collected at Mazon Creek, and without doubt the fossil-bearing nodules from this locality are more widely scattered in the museums of the world than are organic remains from any other one horizon. So far as we know there was no upland vertebrate life at that time. The forms at present known were confined to the water or the margins of the water. The absence of knowledge of upland and terrestrial deposits of this time doubtless accounts for the absence of known vertebrates. It is, however, especially interesting to speculate on the ancestral types of the land vertebrates, and it must be admitted that the Coal Measures Amphibia as at present known throw the ancestry of land- living vertebrates far back into geological time. (w) The Cannelton slates of Beaver County, Pennsylvania, have furnished 3 species of Amphibia and fragments of other species are represented in the U. S. National Museum (462). The species so far known are: Tuditanus minimus Moodie, Erpetosaurus sculptilis Moodie, Erpetosaurus minutus Moodie. They are the first evidence of the occurrence of amphibian remains in these deposits. The Cannelton specimens are found in a thin stratum of slate which forms part of the Middle Cannelton Coal. The Cannelton slate, in which the fossils occur, forms the roof of the Middle Kittanning Coal, which is only 20 to 30 feet above the 16 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Lower Kittanning bed (I.C. White), so it becomes evident that the deposition of the Cannelton slates was at only a slightly later period than that of the shales in which the Mazon Creek nodules occur, since the Mazon Creek shales form the roof of the Morris, which "is probably somewhat lower than the Lower Kittanning of Pennsyl- vania." From the Cannelton slates are known the remains of plants, insects, Crus- tacea, especially "Eurypterids found in shale immediately below the Darlington (Upper Kittanning) Cannel Coal, near Cannelton, Darlington Township, Beaver County, Pennsylvania, Horizon, Allegheny River Series" (Hall, 1884). In these shales occur also fishes and the 3 species of amphibians referred to above. The Amphibia known from this region are small, the largest of them not exceeding 6 inches in length. Contour interval 200 fret Fig. 4. — Portion of "West Virginia-Ohio-Pennsylvania, Wellsville Quadrangle" of the U. S. Geological Survey, to show topography and situation of Linton coal mines. Some fossil amphibians doubtless came from across the line in Columbiana County. («) The Linton, Ohio, beds outcropped near Linton post-office, which was for- merly located at the mouth of Yellow Creek, a few hundred yards from the present station, Yellow Creek, Salem Township, Jefferson County, in the valley of Yellow Creek, near the Ohio River, and thus near the Pennsylvania state line. In regard to the exact location of the town of Linton, which has long since been abandoned, I quote from a letter from Dr. Louis Hussakof, who visited the locality : ' ' The locality appears to have been known as Yellow Creek for many years past. That is the name used in the Geological Map of Ohio published by Orton in 1888 and which was based on the earlier maps of Newberry (1869 and 1879). When I visited the place in 1905, and asked for Linton (which I had not been able to locate on any map then available to me), hardly anyone knew of such a locality. Only one old man in Steubenville, Ohio, recalled that Yellow Creek was identical with Linton. STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. 1 7 "Yellow Creek is not a village, but only a R. R. station (on the Pennsylvania R. R.), and marks a spot where once was an active and prosperous mine. Probably at a former day there was a small post-office somewhere in the neighborhood known as Linton. I did not take any photographs, as I was not certain of the spot, or the mine, from which the fossils had come. There are some cement mines within a few minutes' walk of the station, but no coal appears to be mined at present at Yellow Creek. 'Smith's Pit,' the coal mine best remembered by the younger men, is not worked. "Now as to the question whether some of the Amphibia might have come from localities in Columbiana County. I believe it very probable that they did. I walked along the road from Yellow Creek (Jefferson County) to Wellsville (Columbiana County), a distance of about 2 or 2.5 miles, and the country seemed quite the same. Everywhere one sees outcrops of coal in the cuts along the road. Furthermore, I inclose a copy of a page in an old note- book of Professor Newberry from which you will see that Coal Measure fossil localities were known not only at Yellow Creek, but also from near Wellsville. There can be hardly a doubt that most of the specimens you have are from Yellow Creek; and quite a number are those collected by Sam Huston." Newberry says, in regard to the fauna of the Linton Coal: "The Linton locality is especially interesting and instructive. It has already (1889) yielded more than 20 species of fishes and nearly 40 species of aquatic amphibians, all inhabitants of the same body of water. These were found in a thin stratum of cannel which, over a limited area, underlies a thick bed of cubical coal (No. 6 of the Ohio reports), of which the place is near the top of the Lower Coal Measures. At Linton, ... we have evidence that the great marsh in which the peat accumulated that formed coal No. 6 was for a time a lake or a lagoon, inhabited by the fishes and amphibians to which I have referred. . . . Many of the fishes and the amphibians were highly carnivorous and powerful, as we leam from their teeth and coprolites. The largest of the amphibians must have been 8 or 10 feet in length, having strong jaws, set with numerous lancet-shaped teeth an inch or more in length. . . . After a sufficient time had elapsed for many generations of fishes and aquatic salamanders to live and die, the lake was filled by the extension of its peaty shores into it — just as so many lakelets are filled and obliterated at the present time — and after- ward over the cannel was formed a mass of peat, which has now become a stratum of cubical coal 7 feet in thickness. ' ' In the Linton cannel are buried fragments or entire individuals of all the inhabitants of this body of water which had hard parts, bones, scales, spines, or teeth, capable of preser- vation. Hence we get a locally complete picture of the life of the Carboniferous age, and we find it to be unexpectedly rich and varied. In that age fishes and amphibians were the highest forms of animal life, and the amphibians were comparatively newcomers on the earth's surface. Yet they had multiplied and differentiated until this little pool contained millions of them, varying in length from 6 inches to 10 feet and curiously diversified in their forms, their scales, and spines, and in the ornamentation of their enamel-covered heads" (498). "To the paleontologist there are few places in the world more interesting than the Dia- mond mine, at Linton, since here we get such a view of the life of the Carboniferous age as is afforded almost nowhere else, and of the great numbers of species found there, not more than three or four have been met with elsewhere" (497). On page 18 is a list of the Amphibia which are thus far described from the Lin- ton deposits. They all belong, so far as known, to the Microsauria, the reference of any of the species to other orders being doubtful. The larger Amphibia seem to be indicated by a large rib which resembles very much that described by Huxley in 1863 for Anthracosaurus. 1 8 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Amphibia from the Linton Beds (51 Species). Brachydectes newberryi Cope. Fragment of a skull. Cercariomorphus parvisquamis Cope. Impression of body. Cocytinus gyrinoides Cope. A skull and anterior dorsal vertebra?. Ctenerpeton alveolatum Cope. Large portion of skeleton, no skull. Diceratosaarns Icevis Moodie. Complete skull. Diceratosaurus punctolineatus Cope. Anterior vertebrae, part of skull, with ribs and portion of ventral armature. Diceratosaurus robustus Moodie. Incomplete cranium. Eoserpeton (Ceraterpeton) tenuicome Cope. Incomplete skull. Erpetosaurus acutirostris Moodie. Complete skull. Erpetosaurus obtusus Cope. Incomplete skull. Erpetosaurus radiatus Cope. Incomplete skull. Erpetosaurus tabulatus Cope. Incomplete skull, with clavicles. Erpetosaurus tuberculatus Moodie. Incomplete skull. Eurythorax subkevis Cope. A single interclavicle. (Operculum of lung fish, Sagcnodus.) Hyphasma laevis Cope. Incomplete skull and anterior vertebras. Ichthycanthus ohiensis Cope. Portion of dorsal region. Ichthycanthus platypus Cope. Posterior portion of body. Leptophractus dentatus Moodie. Mandible. Leptophractus lineolatus Cope. Incomplete skull. Leptophractus obsolctus Cope. Portions of skull. Macrerpeton deani Moodie. Mandible and part of skull. Macrerpeton huxleyi Cope. Part of cranium. Molgophis brevicostatus Cope. Part of vertebral column with ribs. Molgophis macrurus Cope. Vertebral column. Molgophis wheatleyi Cope. Part of skull with 2 5 vertebrae. Odonterpeton triangularis Moodie. Skull and anterior part of body. GLstocephalus rectidens Cope. Part of mandible. CEstocephalus remex Cope. Skull and anterior part of body. Pelion lyelli Wyman. Cranium, fore part of body, hind limb. Phlegethontia linearis Cope. Skull and anterior part of body. Phlegethontia serpens Cope. Series of 22 dorsal vertebras. Pleuroptyx clavatus Cope. Part of vertebral column and limbs. Ptyonius marshii Cope. Part of skull and anterior vertebrae. Ptyonius nummijer Cope. Skull and greater part of vertebral column. Ptyonius pectinatus Cope. Many specimens, some nearly perfect. Ptyonius serrula Cope. Nearly complete skeleton. Ptyonius vinchellianus Cope. Skull and anterior vertebra;. Saurerpeton latithorax Cope. Skull and fore part of body. Sauropleura digitata Cope. Greater part of body minus skull. Sauropleura (Anisodexis) enchodus Cope. Part of jaw. Sauropleura joveata Cope. A single interclavicle with impression. Sauropleura longidentata Moodie. Incomplete skull with mandible. Sauropleura newberryi Cope. Two incomplete skulls with vertebras. Sauropleura pauciradiata Cope. Elements of a pectoral arch. Sauropleura scutellata Newberry. Imperfect skeleton. Stegops divaricata Cope. Nearly complete skull. Thyrsidium fasciculare Cope. Dorsal vertebrae. Tuditanus brevirostris Cope. Skull and anterior vertebrae. Tuditanus longipes Cope. Part of vertebral column with limbs. Tuditanus punctulatus Cope. Skull and anterior part of body. Tuditanus walcotti Moodie. Skull and portions of body. Besides the above-listed species there are others indicated by fragments too poorly preserved to be worthy of specific designation. The Linton Amphibia are all apparently confined exclusively to that locality. Species from the Cannelton slates have been assigned, however, to genera which occur at Linton, i.e., Erpetosaurus and Tuditanus. This reference may be due to lack of knowledge, as the forms are insuf- ficiently known. A single Linton species has been assigned to Ichthyerpeton, a genus STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. 19 known otherwise only from the Coal Measures of Kilkenny, Ireland. Cope referred species from Linton to the genus Ceraterpeton of Huxley, from Kilkenny, Ireland, but Jaekel (347) and the writer (462) have shown that the species were incorrectly assigned to the genus Ceraterpeton, and that in fact they represent widely distinct genera. A single species has been identified by Eastman from the Des Moines limestone of Iowa as identical with one from Linton, Pleuroptyx clavatus Cope. The Linton fauna is distinct from that of the Mazon Creek beds, and also from that of South Joggins, Nova Scotia. (0) The deposits in Nova Scotia have been correlated with the Coal Measures strata of the United States (Bell, Summ. Rpt. Geol. Surv. Canada, 1912, 1914, 360- 371). They are very near the same age as the Linton beds and come in near the base of the Allegheny River series. The exposures are at the South Joggins, along the sea- coast. Here in strata of clay interstratified with coal are found the erect stumps of the Sigillariae, and it was in the rock within these stumps that Lyell and Dawson, in 1853, discovered the remains of the amphibians which they termed "reptiles." "The bones of Dcndrcrpeton hitherto found, as well as those of the smaller species, have been obtained from the interior of erect Sigillariae, and all of those in one of the many beds which, at the Joggins, contain such remains. The thick cellular inner bark of the Sigillaria was very perishable ; the slender woody axis was somewhat more durable ; but near the sur- face of the stem, there was a layer of elongated cells, or bast tissue of considerable dura- bility, and the outer bark was exceedingly dense and indestructible. Hence an erect tree, partly imbedded in sediment, and subjected to the influence of the weather, became a hol- low shell of bark. When they remained open for a considerable time, they would constitute pitfalls into which animals walking on the surface might be precipitated. When the sur- face was inundated all such remains would be covered and imbedded in the sediment. These seem to have been the precise conditions of the bed which afforded these remains." (Dawson, 223, 1894.) Fifteen species have been described from the Joggins deposits. Two are known from the Albion mines, south Nova Scotia, where were obtained the remains of Baphetes planiceps Owen and B. minor Dawson. The following 17 species of Amphibia are known from the Carboniferous of Canada : Amblyodon problematicum Dawson. Teeth and fragments. Baphetes minor Dawson. An incomplete mandible. Baphetes planiceps Owen. An incomplete cranium from Albion. Dendrcrpeton acadianum Owen. A jaw, limb bones, and fragments. Dendrerpeton oweni Dawson. Phalangeal bone and fragments. Eosaurus acadianus Marsh. Two dorsal vertebrae. Fritschia curtidentata Dawson. A mandible, vertebras, ribs. Hylerpeton dawsoni Owen. Mandible, teeth and incomplete maxilla. Hylcrpeton intermedium Dawson. Mandible and portions of skull. Hylerpeton longidentatum Dawson. Fragments of mandible and skull. Hylonomus lattdens Dawson. Mandible and teeth. Hylonomus lyelli Dawson. Incomplete skeleton and part of skull. Hylonomus multidens Dawson. Fragments of skull. Hylonomus wymani Dawson. Mandible and vertebrae. Platystegos loricatum Dawson. Incomplete skull, vertebrae. Smilerpeton aciedentatum Dawson. Teeth, ribs, fragments. Sparodus sp indet. Teeth, scales. 20 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. (p) All the remains representing the above species were collected by Sir J. Wil- liam Dawson at the South Joggins and at the mines of Albion, with the exception of Eosaurus, which was collected by O. C. Marsh. The collections of Dawson are now in the Peter Redpath Museum of McGill University in Montreal and in the British Museum of Natural History at South Kensington, London. The history of the discovery of the deposits and their amphibian fossils at the South Joggins is so interesting that it was thought worth while to reproduce in large part Dawson's paper "On the Mode of Occurrence of Remains of Land Animals in Erect Trees at the South Joggins, Nova Scotia," published in 1891 in the Transactions of the Royal Society of Canada, section iv, p. 127: "The remarkable section of coal-formation rocks at the South Joggins, in Cumberland County, has long been known as one of the most instructive in the world; exhibiting as it does a thickness of 5,000 feet of strata of coal-formation in a cliff of considerable height, kept clean by the tides and waves, and in the reefs extending from this to the shore, which at low tide expose the beds very perfectly. It was first described in detail by the late Sir W. E. Logan (Report Geol. Surv. Canada, 1844), and afterwards the middle portion of it was still more detailed by the author (Dawson), more especially in connection with the fossil remains characteristic of the several beds and the vegetable constituents and accom- paniments of the numerous seams of coal (Jour. Geol. Soc. Lond., x, p. 1, 1853). It was on occasion of a visit of the author in company with Sir Charles Lyell, and in the pursuit of these investigations, that one of the most remarkable features of the section was disclosed in 185 1. This is the occurrence, in the trunks of certain trees imbedded in an erect position in the sandstones of Coal-mine Point, of remains of small reptiles, which with one exception, a specimen from the Pictou coal-fields, were the first ever discovered in the Carboniferous rocks of the American continent, and are still (1801) the most perfect examples known of a most interesting family of coal-formation animals, intermediate in some respects between reptiles proper and batrachians, and known as Microsauria. With these were found the first-known Carboniferous land snails and millipedes. Very complete collections of these remains have been placed by the author with his other specimens in the Peter Redpath Museum and in the British Museum. "A forest or grove of the large ribbed trees known as Sigillariw was either submerged by subsidence or, growing on low ground, was invaded with the muddy waters of an inun- dation, or successive inundations, so that the trunks were buried to the depth of several feet. The projecting tops having been removed by subaerial decay, the buried stumps became hollow, while their hard outer bark remained intact. They thus became hollow cylinders in a vertical position and open at the top. The surface having then become dry land, covered with vegetation, was haunted by small quadrupeds and other land animals, which from time to time fell into the open holes, in some cases nine feet deep, and could not extricate themselves. On their death, and the decomposition of their soft parts, their bones and other hard portions remained in the bottom of the tree intermixed with any vege- table debris or soil washed in by rain, and which formed thin layers separating successive animal deposits from each other. Finally, the area was again submerged or overflowed by water, bearing sand and mud. The hollow trees were filled to the top and their animal con- tents thus sealed up. At length the material filling the trees was by pressure and the access of cementing matter hardened to stone, not infrequently harder than that of the contained beds, and the whole being tilted to an angle of 200, and elevated into land exposed to the action of the tide and waves, these singular coffins present themselves as stony cylinders projecting from the cliff or reef, and can be extracted and their contents studied. The sin- gular combination of accidents above detailed was, of course, of very rare occurrence, and. STRATIGRAPHIC AND GEOGRAPHIC DISTRIBUTION. 21 in point of fact, we know only one set of beds at the South Joggins in which such remains so preserved occur; nor is there, so far as I am aware, any other known instance elsewhere. Even in the beds in question, only a portion of the trees, about 15 out of 30, have afforded animal remains. We have, however, thus been enabled to obtain specimens of a number of species which would probably otherwise have been unknown, being less likely than others to be preserved in properly aqueous deposits. Such discoveries on the one hand impress us with the imperfection of the geological record; on the other, they show us the singular provisions which have been made in the course of geological time for preserving the relics of the ancient world, and which await the industry and skill of collectors to disclose their hidden treasures. "There is evidence in coprolitic matter on one of the surfaces within the trunks, and also in certain trails on these surfaces, that some of the imprisoned animals lived for a time in their subterranean prisons; that they crept around their walls in search of a way of escape, and that the larger animals fed on smaller species entrapped along with them." After the discovery of these en- tombed amphibians Sir William Dawson was given a grant of £50 from the Gov- ernment Fund by the council of the Royal Society of London, to aid in the extrac- tion of these trees and the collection of their contents. The trees were carefully taken out and their contents examined ; the portions containing the animal re- mains were carefully boxed to be taken to Montreal for final cleaning and study. Erosion goes on rapidly at the South Joggins, but no one has paid any atten- tion to the occurrence of Amphibia along the coast of Nova Scotia within recent years. (q) A deposit which will be of un- doubted interest in connection with the occurrence of Amphibia in the Coal Measures is that which outcrops along the banks of Rock Creek in the south- western part of Douglas County, Kansas, Fig. 5.— Dawson's tree No. 13 at the South Joggins, Nova Scotia. Upper part, in situ, in the reef after it had been exposed by blasting. (After Dawson, based on a photograph.) in Marion Township (Township 14 south, Range 18 east, SW. and SE. quarters of section 7), about 2 miles from the now-abandoned post-office of Twin Mounds, so called from the two flat-topped, elongated mounds of Oread limestone to the west of the town. The interest in these beds is not due to the discovery of Amphibia in them, but the possibilities of such discoveries. This is indicated by the occurrence of fossils, in nodules similar to those obtained from Mazon Creek, which are identical gener- ically, and in most cases specifically, with the Mazon Creek animals and plants. 22 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The fossils so far collected from this interesting locality are : Insecta (Identified by Dr. E. H. Sellards). Spiloblattina maledicta (Scudder). The basal half of a front wing. Etoblattina sp. The hind wing of a cockroach. Arachnida. Anthrocomartus. Impression of the body. Prestwichia dance M. & W. Nearly complete specimen. Crustacea. Acanthotelson stimpsoni M. & W. Three nearly complete individuals. Plants (Identified by Mr. J. C. Carr, of Morris, III). Pecopteris sp. Sagittaria reticulata Lesquereux. Annularia longijolia Lesquereux. Annularia inflata Lesquereux. Pecopteris villosa Brongniart. Neuropteris decipiens Lesquereux. Pecopteris serpulifolia Lesquereux. By far the most abundant plant is Pecopteris. The fossils occur in definite strata of nodules immediately above a io-inch coal seam which is worked for local consumption. The coal lies near the base of the exposure in the more western portion of the outcrop, but it is raised by an anticlinal fold to near the top of the creek-banks by the bridge across Rock Creek, along the banks of which the shales are exposed. Nodules containing fossils are found most abundantly at the western exposure on the McKinzie place, only a few having been found near the bridge. Below the coal-seam, nodules of various shapes and sizes occur, but they seldom contain fossils and never good ones. Occasionally, as at Mazon Creek, fragments of plants adhere to the outside of the incrusting shale. A single nodule may have adhering to it fragments of 4 genera of plants. The fossiliferous nodules all occur above the coal and are most prolific and abundant immediately above the seam, within the first 10 inches. In the same stratum of shale with the nodules are found abundant impressions of plants in the shale, often perfect fronds being uncovered. (See, in this connection, Twenhofel and Dunbar, 1914, "Nodules with Fishes from the Coal Measures of Kansas," Amer. Journ. Sci., xxxvm, pp. 157-163.) G. F. Matthew (408-413) has described numerous genera and species of foot- prints, presumably amphibian, from the Carboniferous of Canada. The impressions indicate small creatures for the most part. Other imprints have been described by Logan, Dawson, Lyell, Marsh, Mudge, and Lea. Since the present work is intended largely for a morphological review, only passing notice can be given to the ichnites. The literature on the " Ichnites" has been brought together in Hay's "Bibliography and Catalogue of Fossil Vertebrata of North America," pp. 538-553. References since the publication of Hay's catalogue (317) will be found in the bibliography at the end of this work. Footprints are of interest in that they are the only evidence we have of the occurrence of land vertebrates in the Devonian and Mississippian of North America. CHAPTER IV. THE MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. The anatomy of the Coal Measures Amphibia presents many primitive types of structure. Their organization represents a stage passed through in the ontogeny of higher vertebrates. The animals are similar in a general way, yet so diverse are the modifications which they have suffered under different environmental conditions, that close scrutiny is needed to discern the exact relationship of the forms. Our knowledge of this relationship is based on the structures preserved, which are largely skeletal, since little is known of the soft anatomy (471) of the air-breathing vertebrates of the Coal Measures. The pubis is ossified in the Paleozoic Amphibia a. com. Fig. 6. — Generalized figure of dorsum of an early amphibian skull to show position of elements and terminology adopted m this work. The outline is based on that of Eryops, but is in no way intended to indicate that form. a. com, anterior commissure of lateral-line canals; com, commissural communication between infra- and supra- orbital lateral-line canals; fr, frontal; inf, interfrontal; inn, internasal; info, infraorbital lateral-line canal; it, intertemporal; jl, jugal lateral-line canal; j, jugal; lar, lacrimal; mx, maxilla; n, nasal; oc, occiput; occ, occipital cross-commissure of the lateral-line system; or, orbit; par, parietal; pof, postfrontal; pmx, premax- illa; pf, prefrontal; po, postorbital; pp, postparietal ; q, quadrate; qj, quadratojugal ; spo, supraorbital lateral-line canal; sq, squamosal; spt, supratemjxjral ; /, temporal lateral-line canal; tab, tabulare. - Jl later than the ischium and ilium; the carpus and tarsus are cartilaginous; the verte- brae consist of a pleurocentrum and two neurocentra, thus paralleling conditions in modern mammalian embryos. (a) The skull of the Coal Measures Amphibia has (fig. 6) essentially the same structure in the different groups. It is largely formed of bones of intramembranous origin, representing the face bones of the mammalian skull. The skull in life was doubtless a chondrocranium with the membrane bones laid down upon the carti- laginous box containing the sense-organs, as in the sturgeon (Acipenser), where the surface bones of the face were probably originally scales, which later became consol- idated into large bony scutes. The membrane bones of the early Amphibia may have been originally derived from scales, but at present nothing is known of this origin ; doubtless the elements had an intramembranous origin in the ancestors of 23 24 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. the group. Judging from Credner's studies on the series of specimens of Branchio- saurus amblystomus Credner (187), the skull bones do not ossify completely until relatively late in the life of the individual. The skull in the youngest individual figured by Credner {op. cit., Taf. xvi, fig. 1) seems to be largely cartilaginous, with beginnings of separation into the skeletal elements. The manner and time of devel- opment and ossification of the skull seems to proceed much as it does in modern amphibians. The condition found in the skull of Cryptobranchus allegheniensis or Necturus maculosus will represent pretty accurately the condition of most of the Coal Measures Amphibia. The face bones in certain forms were sculptured and cut by lateral-line canals. A median suture divides the skull into two equal regions dorsally. On the sides of this median suture lie pairs of elements which are common to all higher verte- brates. These elements are: the premaxillae, nasals, frontals, parietals, and post- parietals. All of these elements vary somewhat in shape and slightly in arrange- ment, but always occupy the same relative positions. To the side of these elements lie the prefrontal, the postfrontal, the supratemporal, the squamosal, and tabulare, and occupying the margin of the skull are the maxilla, the jugal, the quadratojugal, and possibly the quadrate in a few forms. The parietal foramen occurs usually within the parietal bone, but its position is subject to slight variations and it may occur on the suture between the frontal and the parietal, or even far posterior near the postparietal. The nostrils often lie well forward and are included by the premaxillae, nasals, and prefrontals. The orbit is usually well posterior, but it may occur far forward. It is bounded by the prefrontal, the frontal, the postfrontal, the post- orbital, and the jugal. Sometimes the lacrimal is present and has been clearly identified on the anterior margin of the orbit in a few cases. (b) Sclerotic plates often occur within the orbits, and are not confined to any par- ticular group, though they are quite constant among the Branchiosauria. They are usually delicate, thin, broad plates which evidently overlap and operate as in mod- ern animals. The number varies, as many as 30 occurring within the orbit of one branchiosaur. Between the margin of the orbit and the sclerotic plates there often occur, in the Branchiosauria (186) particularly, small scale-like particles which were doubtless embedded in the heavy skin above the orbit during life. (c) The palate of the skull is very incompletely known, being indicated in a very few cases. These specimens, however, show that the characters of the palate were quite similar, if not identical, in essential respects with the palate among the Euro- pean species of the same or slightly later time. A large cultriform parasphenoid occupies the posterior portion of the palate, on either side of which in some species lies the posterior palatine foramen. On the sides of the anterior prolongation of the parasphenoid lie the vomers (186), membra- nous bones often bearing minute tubercular teeth, apparently adapted for crushing. The vomers and the maxillae, with sometimes the palatine, surround the anterior palatine foramen, which is almost always present; sometimes, however, quite small. The transverse or ectopterygoid unites the pterygoid, a broad plate of thin bone, with the maxilla and jugal. MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 25 (d) The teeth of the Coal Measures Amphibia (194) are remarkably similar in the various forms. They are always slender, pleurodont denticles arranged in a single row on the jaws or as tubercular eruptions on the palate bones, with a large pulp- cavity and the enamel often striated. The food of the creatures must have been small Crustacea, worms, insects, and succulent vegetation, such as is the food of the modern Amphibia. (e) The occiput is formed of partially ossified (465) ex- and basi-occipitals, though these elements are never firmly united by ossific union. Often a pair of condyles occur, one on either exoccipital. The occiput was usually, however, car- tilaginous and no trace of its structure is preserved. (/) The mandible is usually as long as the skull and is slender. It is composed of 6 elements so far as known (465) ; these are the articular, the surangular, the angular, the coronoid, the dentary, and the splenial. Other elements may be present, but the anatomy of this portion of the animals is not very completely known. The bones are sculptured and cut by lateral-line canals (458) in a few forms. Whether the articular operated on an osseous or cartilaginous quadrate is unknown, though certain specimens seem to indicate an osseous condition for that element. The anterior symphysis was doubtless ligamentous, the halves always separating before fossilization. The dentary always bears a single row of pleurodont teeth, which may vary greatly in size and number. (g) The hyoid apparatus is well preserved in a few forms (123). Doubtless it was present in all of them, though it has seldom been preserved. The condition of the hyobranchial apparatus in Cocytinus gyrinoides (text-fig. 16) from the Coal Measures of Linton, Ohio, seems to indicate that the species was a perennibranchiate salamander (123). It is well known from the studies of Credner that the Euro- pean Branchiosauria, in the young, possessed external branchiae (187) supported by lateral basibranchials. The gill-arches seem to have been slightly calcified or ossified in a few cases, and they supported denticle-like projections which bore the gill-filaments. When the Branchiosauria had attained a length of 100 mm. or more they lost their gills (187). This change was accompanied by the reduc- tion of the tail, expansion of the pelvis, and increase in ossification of the skull and skeletal elements. Gills have not yet been detected among the American Branchiosauria. (//) The eye in a few species had a large amount of black pigment, as indicated by the blackening of the stone in the Mazon Creek nodules. Professor Cope (107) thought that this would indicate a nocturnal and crepuscular habit for these verte- brates, since the pigmentum nigrum of the choroid is largely developed. Other than this suggestion nothing is known of the soft parts of the head. (I) The alimentary canal (text-fig. 7) is beautifully preserved as a cast in three specimens of the American branchiosaur species Eitmicrcrpcton parvum Moodie (471) from the Mazon Creek beds. The nodules which contain these interesting little fossils measure less than 3 inches in long diameter. The fossil salamanders, about 30 mm. in length, are preserved on their backs and occur as nearly as is possible in the center of the nodule. 26 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. If it were not for the fact that the oesophagus became loosened and dropped from its place shortly after death, the alimentary canal would be in place and would immediately recall a freshly dissected specimen of a recent salamander. The ante- rior end of the oesophagus lies obliquely across the chest region with its tip pointing slightly downward. The length of the oesophagus proper, in one specimen, is only about 3 mm. As it is preserved, the oesophagus lies in a semi-sigmoid curve with the convexity anterior, and enters the cardiac portion of the stomach by a gradual con- striction. The stomach is clearly preserved as a distinct sac-like organ with two lobes which correspond to the cardiac and pyloric limbs. It measures about 7 mm. in length by 2 mm. in its greatest diameter. The muscular constriction which divides the organ into pyloric and cardiac divisions occurs at a distance of 4 mm. from the upper end. The pylorus is designated by a rather pronounced constric- tion which may be partly accidental, although it recalls the Is' pylorus of modern frogs. From this constriction, which lies >j>, on the left side of the fossil, as it is preserved, the duodenal ^ portion of the intestine makes a straight course posteriorly to near the anal region, where it takes a sharp bend and curves back to run parallel with itself for the distance of 4 mm. In its upward course the intestine enlarges, and practically the same enlargement continues throughout the remainder of the course to the anus. At a distance of 1 mm. from the anal end, the rectum dilates probably 0.125 mm. to form the cloaca. After the intestine has continued its parallel course for the 4 mm., as above stated, it turns abruptly to the right for a distance of 2 mm. It then runs posteriorly for a short distance, then bends back and under itself to again make a double sigmoid curve, when at a dis- pIG. tance of 6 mm. from the anus it assumes a straight course, which it continues to the end. The anus lies at a level which is approximately that of the lower end of the femur, which is preserved as an impres- sion on the left side of the fossil, thus agreeing in its position with that found in modern Caudata. Lying inside the curve of the stomach and partly inclosed by the oesophagus is a smooth area which may possibly represent the impression of some of the accessory digestive glands, such as the liver. Occurring in this smooth area are numerous fine lines which possibly represent the impressions of blood-vessels; but they are so imperfectly preserved that one can not be sure. Representatives of several genera of the modern Caudata have been dissected in order to make a direct comparison of the fossil alimentary canal with that of the recent forms. The alimentary tract of Desmognathus fuscus Raf. from the vicinity of Ithaca, New York, resembles in a marked degree that of the fossil form. The nearest approach to the condition there represented is found, however, in an imma- 7. — Alimentary canal of a Coal Measures salamander as illustrated by the smaller speci- men of Eumicrerpeton parvum Moodie, from the Mazon Creek shales. X 3. Original in Yale University Museum. a, anus; dd, duodenum; in, intestine; /, impression of liver(?); oes, oesophagus; si, stomach. MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 27 ture branchiate individual, some 47 mm. in length, of Diemyctylus torosus Esch., from a fresh-water pond on Orcas Island in Puget Sound. The presence of this species on the island is very suggestive. It is of extreme interest, too, that the con- dition represented in the alimentary tract of the fossil branchiosaurs should resem- ble so closely that of an immature rather than a mature form. (j) The vertebral column is clearly and readily separable into cervical, dorsal, sacral, and caudal regions. The neck is always short, with from 5 to 10 vertebra?, cervical ribs often present. The dorsal region is not long, but varies from 20 to 30 in the constituent vertebrae. There is a single sacral vertebra not always to be readily distinguished from those of the dorsal and anterior caudal series. The tail may be very short or extremely long, with neural and haemal spines elongate and flattened into an oar-like appendage. The distal caudals are in some species car- tilaginous, apparently always so in the Branchiosauria. (k) The atlas and axis are unknown among the American specimens, but we are able to infer from the structure of the other vertebrae what this must have been; and our inferences are partly confirmed by the conditions existing in the European forms (187). The atlas, apparently, consisted of a pair of neurocentral plates which are partly ossified, partly embedded in cartilage, judging from the edges of the plates which have been preserved. The centrum seems not to have been present in the atlas, or if present it was only very slightly developed and quite free from the neural pieces and largely embedded in cartilage. A fairly accurate picture of the condition of the atlas and axis may be seen on examining a cow, pig, or chick embryo (378) in the early stages of vertebral development, which has been cleared by the Schultze method (Amer. Journ. Anat., vil, No. 4, 1908). (/) The dorsal vertebrae, as well as those of the other series, present a primitive character (fig. 8) in the persistence of the notochord (540). Among the Branchio- sauria the notochord was not at all or but slightly constricted intravertebrally, but among the Microsauria it was constricted so far that the notochordal remnants in each centrum resemble an hour-glass. The structure of the vertebrae among American forms agrees fully with that outlined by Credner, Fritsch, and others for the European species. The details of structure are so fully given by Zittel (642, pp. 346~353) and by Schwarz (540, 541) that it will not be necessary to state more here as to their structure, since there is nothing new to add concerning the American species. The temnospondylous vertebra of the same nature and type as exhibited by the Permian forms has its representatives (94, 478) in the Coal Measures. Spondyler- peton spinatum Moodie (478) (plate 4, figs. 1, 2) and Eryops sp. (plate 18, fig. 2) indicate the embolomerous and rachitomous types of vertebral structure. The occurrence of these widely different types of vertebral structure indicates a long his- tory for the group prior to the Coal Measures. This history is further indicated by footprints in the Mississippian and Devonian of this continent. (m) The ribs (fig. 8) are very diverse in structure and in their mode of articula- tion (541) with the vertebral column. The characters of the ribs and vertebrae con- stitute the best means of classification of these animals so far discovered. In the 28 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Branchiosauria the ribs are always straight, heavy, and short, and articulate intra- vertebrally upon a large and strong transverse process. They occur throughout the vertebral column. There is a single pair of sacral ribs which are not to be clearly distinguished from the pre-sacral and post-sacral series. The cervical and caudal ribs are shorter than the dorsal series. The branchiosaurian rib is composed almost entirely of perichondral ossification. It presents the same condition as does the humerus of the cow embryo of 2 to 3 inches in length. The ribs. of the branchio- Fig. 8. — Vertebrae and ribs of Amphibia from the Coal Measures of Linton, Ohio. Originals in Geol. Inst. Berlin. (All after Schwarz.) A. Caudal vertebra of CRstocephalus remex Cope. Lateral view. X 4. B. Caudal vertebra of Ptyonius vinchellianus (?) Cope. Lateral view. X 6. C. Dorsal vertebra of Ptyonius peclinalus Cope. Lateral view. X 9. D. Notochordal cones and spinal canal of Thyrsidium fasciculare Cope. X 3. E. Rib of Molgophis sp. Cope. X 1.75. r = capitulum; / = tubercuium. F. Vertebra of Molgophis sp. Cope, from ventral side. X 2. G. Dorsal vertebra of Ptyonius pectinatus Cope. From above. X 8. H. Dorsal vertebra of Thyrsidium fasciculare Cope, from below. X 2.5. I. Vertebra of Phlegethontia linearis Cu\x-, from alxive. X 5.5 J. Rib of (Estocephalus remex Cope, from posterior dorsal region. X 5. K. Dorsal vertebra of Thyrsidium fascicidare Cope, from above. X 1.5 L. Anterior dorsal vertebra (cervical?) of Thyrsidium fasciculare Cope. Lateral view. X 1.5. M. Vertebra of Phlegethontia linearis Cope, from side. X 5. saurs are identical in every way with the ribs of modern salamanders and form one of the strong arguments in favor of the relationship of the Branchiosauria to the Caudata. Among the Microsauria the ribs are always long, slender, curved, and intercentral. They may be either single or double headed, but usually the former. They resemble in their characters the ribs of some of the early reptiles and an attempt has been made to relate the Microsauria (469) to the primitive reptiles on this basis. The ribs of the other groups are still unknown. Indeed, representatives of the Temnospondylia and the Stereospondylia are very scanty in the American MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 29 Coal Measures. One large rib (plate 22, fig. 4) may represent a labyrinthodont, but nothing is known of the species to which the rib belonged. (h) The pectoral girdle (187) is a very simple and uniform structure, although the details of the association of the elements still remain to be determined. A single, median, usually large and elongate interclavicle occupies the ventral line of the chest. This is morphologically the same element which occurs in the middle line of the chest of the lizards. It is a dermal bone and is usually, especially among the Microsauria (462), highly sculptured. It varies considerably in size and shape, but is remarkably uniform throughout the various groups. Lying anterior to the interclavicle and overlapping its antero-lateral margins lie the two clavicles, which are usually diamond-shaped and are sculptured, dermal bones. The position of the coracoid is still uncertain, and in fact its clear association in the pectoral girdle of these species is still a question. It seems to be constant in the European (186, 251) species and is usually represented by a small rounded plate of bone, which in life no doubt had a large amount of cartilage to form its borders. A cleithrum (285) has been ascribed to one of the Linton, Ohio, species (plate 15, fig. 3) by Jaekel (347), but this needs confirmation. An osseous scapula is usually present, resembling the scapula of modern salamanders, in that it was largely embedded in cartilage. The position of the pectoral girdle is largely a matter of doubt, especially for the American spe- cies. After death and before fossilization the girdle was always moved by post- mortem shifting, so that its exact relation to the ribs and vertebral column is still in doubt. Credner (186) has restored the pectoral girdle close behind the head, which would cause an amount of rigidity in the body which probably did not exist. (0) The arm consists of the humerus, radius, ulna, and 4 digits. The characters of the arm-bones are such as is constant among primitive animals and developing mammals. The osseous portion is perichondral. Epiphyses are totally lacking and it is doubtful if the endochondrium was at all ossified. The digits are often termi- nated by ungual phalanges, although usually the terminal phalanx was merely embedded in the web of the foot ; and among the terrestrial forms a claw was well developed. An osseous carpus is not known in the species from the Coal Measures. Its impression indicates a broad hand, well adapted for swimming. (p) The pelvic girdle consists uniformly (462) of the ilium and ischium. A small rounded pubis is present in some of the later forms of Amphibia; it is, however, totally absent from the Coal Measures species. The condition of the pelvis is paralleled by the partially grown pelvis of mammalian embryos in which the elements ossify in the order of ilium, ischium, and pubis. The ilium is always the larger of the elements. It supported or was attached to the sacral rib by means of a liga- mentous union. The ischium did not ossify completely until the animal was nearly mature. The union between the elements of the pelvis was probably of a loose, membranous sort or else the whole mass was embedded in cartilage; of the two hypotheses the former is the more probable. The pubis is indicated as a calcified quadrangular plate in a specimen olAmphi- bamus grandiceps Cope (478) from the Mazon Creek shales, and it is present as a rounded osseous element among some of the Permian forms. 3° THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. (q) The leg (fig. 21, B) is composed of the femur, tibia, fibula, and 5 digits. The tarsus is usually cartilaginous, a single osseous tarsus (483, 484) being known (plate 23, fig. 1) from America. The distal phalanges may or may not be clawed, depending on the habits of life of the animal. The elements of the leg are ossified in a similar manner to those of the arm. (r) The ventral scutellation (fig. 9), so commonly present among all groups of Amphibia in the Coal Measures, consists of a series of ossifications or calcifications in the myocommata. Among modern amphibians they occur as thin perpendicular planes of connective tissue which are sometimes cartilaginous, especially in Necturus, regarded by Wilder (Memoirs of the Boston Society of Natural History, vol. v, No. 9, p. 400, fig. 6, 1903) and by Wiedersheim (605, p. 58) as a homologue or predecessor of the sternum, although Wiedersheim says: ' ' The sternum appears for the first time in Amphibians in the form of a small variously shaped plate of cartilage situated in the middle line of the chest. It arises as a paired cartilaginous plate in the inscriptiones tendineae of the rectus abdominis muscle, and therefore may be looked upon as corresponding to a pair of 'abdominal ribs.' Such cartilaginous abdominal ribs must have been present in greater numbers in the ancestors of existing Urodeles." This supposition is fully sustained by the anatomy of FlG the Branchiosauria (459), which must be looked upon as the actual ancestors of the Caudata. Wilder says of these structures in Necturus (op. cit., p. 400): "The several cartilaginous rudiments which represent this part {i.e., sternum) in Nec- turus are somewhat difficult of detection and thus entirely escaped the attention of the earlier investigators. They consist of a number of thin cartilages found in several suc- cessive myocommata of the pectoral region and confined mainly to the area covered by the overlapping epicoracoids. " The homologue of the ventral scutellce is found in plesiosaurs, crocodiles, Sphenodon, and other reptiles in the "abdominal ribs," and the same myocommatous ossifica- tions undoubtedly go to the formation of the chelonian plastron. What the causes were which produced the development of the ventral scutellae to such a high degree among the primitive land vertebrates is uncertain, but they are certainly more highly developed among the primitive reptiles and amphibians than among the later members of those classes. Among the Amphibia of the Coal Measures they attained, in some forms, a high degree of development and differentiation. They are present in all families so far known, except the Tuditanidas, in which the myo- commata may have been cartilaginous. The Sauropleuridae present the highest development of these structures among the American forms, in which the scutes are large and osseous. Among the Branchiosauria they are calcified or partially ossi- fied and are always arranged en chevron on the belly, chest, arms, and throat, their arrangement and direction of the chevron being modified according to the myomeres of the various regions. The ventral scutellse of the European Branchiosauria are figured and described fully by Credner (192, p. 21, figs. 4 to 11). 9. — Ventral scutelke of Micrer- pclon caudatum, a Coal Meas- ures salamander from Mazon Creek. X 5. /.femur; /(, hu- merus; Is, lines of scutes; v, vertebral column. MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 31 (s) Scales (fig. 10 and plate 24, figs. 2 and 3) are present on the body of (462, 485) several species. It is a matter of regret that their preservation is so imperfect that nothing can be found out as to their structure. The Linton species, which pos- sess scales, are, of course, carbonized and hence impracticable for microscopic study, and in the Mazon Creek species of Amphibamus and Micrerpeton the scales have been replaced by kaolin. The bodies of two species (Cercariomorphus parvisquamis a Fig. 10. — Horny armor of back of Hylonomus. a, imbricated scales; b, horny plates; c, horny spines or tubercles; d, small imbricated scales. (After Dawson, based on a photograph.) and Ichthyerpeton squamosum) of the Linton Coal Measures Amphibia were com- pletely scaled. The scales in the Branchiosauria (462), so far as they have been observed, are slightly imbricated; rounded, with concentric markings after the manner of the modern cyprinoid fish-scale. They are extremely minute, and whether or not they covered the entire body of the animal is unknown. On the body of Cercariomorphus the scales have the appearance of being tubercular without imbrication, and they apparently covered the entire bodily surface of the animal. 32 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Among the Paleozoic Amphibia from Nova Scotia as described by Dawson and Owen (193, 201, 485) scales are well developed and frequent, although the details as to their occurrence on the bodies of the animals are still unknown, since the Nova Scotian species are all based on very fragmentary remains. Dawson (208, p. 34) has given a detailed discussion of the discovery and anatomy of the various types of scales possessed by the species from the Coal Measures of Nova Scotia. Suffice it to say here that none of the scales appear to be bony, but have a cuticular appear- ance with concentric markings. Some of them are tubercular, and Dawson thought that a few specimens indicated that some of the species possessed scaly lappets and a dorsal nuchal fringe of scaly skin along the back. He has indicated these facts in his restorations of the forms. The scales were all carbonized and burned readily with a strong flame. A section of the scale shows a thick upper corium with a vas- cular body (208, pi. iv, fig. 29) much like a fish-scale. Fragments of the skin were also preserved with the scales. Dawson says of the skin: "One of my specimens is a flattened portion of cuticle two and a quarter inches in length. The greater part of the surface is smooth and shining to the naked eye, and under the microscope shows only a minute granulation. A limited portion of the upper and, I suppose, anterior part is covered with imbricated scales, which must have been membra- nous or horny, and generally have a small spot or pore near the outer margin, some having in addition smaller scales or points on their surfaces" (208, pi. iv, figs. 22 and 25). (t) Muscle tissue (fig. 21) is preserved in a single specimen, previously described by the writer (464, p. 17, pi. 7, fig. 1). The carbonized muscles show a myomeric arrangement and the portions preserved probably represent one of the recti muscles of the abdominal wall. (u) The lateral-line system in the Coal Measures Amphibia will be best under- stood from a comparative review of the occurrence of these organs among all extinct Amphibia. Since all the orders of Amphibia are represented in the Coal Measures, such a review will not be out of place here. The preservation of the lateral-line system among ancient Amphibia is due to the fact that the skull of many forms (especially the later and larger) are grooved and marked by a regular series of furrows and pits, in which the sense-organs of the lateral-line system were contained (see fig. 6), as well as by the preservation of a series of clearly marked scales on the sides of the tails and bodies of others. The grooves are never arched over as in the Macropetalichthyidae, where "in favorable specimens each is shown to be covered by a delicate roof perforated by two lines of minute openings" (Dean, N. Y. Acad. Sci. Mem., vol. 11, pt. in, p. 115). They are always widely opened canals, either with perfectly smooth bottoms and sides or roughened with large pits, or even becoming a series of well-marked pits. An attempt has been made (458) to homologize the organs with those of fishes. The nomenclature adopted here for the canals does not depart from that used by Allis for Amia (Journ. of Morphology, vol. II, 1889). The supraorbital and infra- orbital canals are readily correlated with those of the same name in fishes, where they are very clearly marked. The anterior commissure is also homologous with that of the fishes, as is also the canal here called the " antorbital commissure. " The MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 33 others arc not so readily homologized. The upper canal (see fig. 6) in the posterior part of the cranium is here designated the temporal canal. It is, however, clearly a part of the infraorbital of the fishes. Its relations in the Stegocephala are such that a new name is deemed necessary. The jugal canal is, I believe, a new formation in Amphibia. The transverse canal of the amphibian skull is homologous with the "occipital cross-commissure." The figure (see fig. 6) is a composite picture of the lateral-line system of the higher or truly stegocephalous Amphibia. The outline of the skull is based on that of Eryops. All of the canals do not exist on any one skull or in any one order, but all are found somewhere in the group. Fig. ii. A. Skull of Eoserpeton tenuicorne Cope, showing arrangement of cranial elements. X 2. fr, frontal; j, jugal; mx, maxilla; n, nasal; or, orbit; par, parietal; pof, postfrontal; pmx, premaxilla; po, post- orbital; pp, postparietal; qj, quadrat ojugal ; sq, squamosal; spt, supratemporal; tab, tabularc. B. Outline of skull of Ceraterpeton galvani Huxley from the Carboniferous of England. Heavy broken lines show the distribution of lateraldine canals. X I. (After Andrews.) fr, frontal; par, parietal; or, orbit; po, postorbital; pp, postparietal; spt, supratemporal; tab, tabularc. The canals have been described in all known orders of fossil Amphibia and the system is found likewise in all the living orders, including the Gymnophiona, which have "a strong line of lateral sense-organs" (Gadow). In the Branchiosauria, the earliest of the true Amphibia (Euamphibia) and ancestral to the modern Caudata, the lateral-line system is known on the tails of two genera (462, 478) from the Mazon Creek, Illinois, shales — Micrerpeton and Eumicrerpeton. The system as there defined has been fully discussed in the description of the anatomical details of the species, to which reference may be made for further data (pp. 52-60) . Suffice it to say here that the system of sense-organs there preserved is identical with that of the larval Necturus; the lines arising as a median from the tip of the tail and a dorsal springing from the median at a distance of a few millimeters from the tip of the tail. The lines are more evident on account of the fact that the lateral-line sense-organs were located under specialized pigmented scales. The significance of the close simi- 34 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. larity between the arrangement of the lateral-line systems on the tail of Necturus, Micrerpeton, and Eumicrerpeton is doubtless of genetic (459) importance, indicating the origin of the caudate Amphibia from the Branchiosauria by a degenerative or recessive evolution in other structural characters. This system of sense-organs has been described in no other branchiosaurian. The Microsauria (458) are exceedingly interesting in possessing a very peculiar type of lateral-line system. It is known in a few forms and in one specimen espe- cially well {Erpetosaurus tabulatus) (fig. 22, G). In this species, which is represented by a single imperfect skull, there are evidences of a nearly complete lateral-line sys- tem of canals and pits. The occipital cross-commissure is represented on the pos- terior border of the skull by a row of elongate pits such as Andrews described for Ceraterpeton (8). I fail to find in American species the pores described by Andrews. The temporal canal forms with the jugal canal a complete ring, much as it is in Tre- matosaurus, only in Erpetosaurus tabulatus the temporal canal does not touch the tabulare. I think there are indications of a connection of the temporal canal with the supraorbital. The temporal canal cuts the supratemporal, the squamosal, and jugal. The jugal canal lies for the most part on the supratemporal and quadrato- jugal, and joins the infraorbital on the jugal. A portion only of the infraorbital canal is preserved. There is also a portion of the supraorbital canal. It seems not to be connected with the temporal canal, although there is a possible indication of this connection. The supraorbital crosses the frontal, prefrontal, and a part of the nasal. The squamosal element is peculiar in Erpetosaurus tabulatus in that it is excluded from the parietal by the extension of the tabulare and postorbital. This condition is found in several other species of the Microsauria. It will be noticed that with the changed condition of the squamosal the temporal canal has changed also, and this is further proof of the close connection between the cranial elements and the lateral-line canals, as Allis has maintained for Amia. (See in this connection C. J. Herrick, Journ. Comp. Neurol., vol. 11, p. 224, 1901.) The Diplocaulia, an amphibian order allied to the Branchiosauria (477) and through them to the Caudata, have the lateral-line system apparently well-devel- oped. The skulls are always crushed flat, so that the canals are nearly obliterated. On the mandible, however, the canals are clearly distinct and apparently run the entire course around the mandibular rami. On a well-preserved skull of Diplocaulus magnicornis Cope there are indications of three lateral-line canals (477, pi. 1). The infraorbital is clearly marked as a well-defined groove just below the orbit. The supraorbital is indicated only for a short distance, and there are indications of the temporal canal. The operculo-mandibular canal has its course, for the most part, near the middle of the rami, but as it approaches the posterior angle of the mandible it suddenly changes its course and drops down to the lower edge, only to rise again and to come out strongly marked near the median plane on the posterior angle of the mandible. The Temnospondylia, as represented by Eryops, Cricotus, and Archegosaurus, possess well-developed lateral-line canals (458). H. von Meyer (428) many years ago made out the course of the canals in Archegosaurus. The greater part of the fol- MORPHOLOGY OF THE COAL MEASURES AMPHIBIA. 35 lowing description is based on Eryops megacephalus Cope from the Texas Permian. The entire surface of the cranial elements in Eryops, as in other of the Stegocephala (458), is covered with coarse pits. The fossae are present even in the bottoms of the grooves which represent the lateral-line system, and are more marked in the mem- bers of the Temnospondylia than in the Stereospondylia. The occipital cross-commissure occurs in a well-developed form in Eryops. It is short and ends abruptly within the limits of the tabulare. Its ends are occupied by large pits. The commissure, as in Amia, grooves the postparietal and the tabu- lare elements. There is no evidence of an anterior commissure. I think there is evidence of a temporal canal on the left side of the skull, but am not sure. The jugal and infraorbital canals are well developed and strongly connected. The jugal canal starts far back on the supratemporal, and after curving around over the quad- ratojugal joins the infraorbital, or rather becomes a part of that canal, somewhere on the jugal. There is nothing unusual in the infraorbital. The antorbital commis- sure is well developed. It is longer and better developed than in any other known form. The supraorbital canal starts in the region of the orbit, and after curving downwards to meet the antorbital commissure, ends abruptly anterior to the nostril. There are faint traces of a lateral-line canal, the operculo-mandibular, on a poorly preserved mandible of Eryops. It does not differ greatly from that described below for Anaschisma. Although Archegosaurus has been known for more than a century, we have had no adequate discussion of the manner of occurrence of the lateral-line canals. Bur- meister (80) gave a figure of the canals as he thought they occurred on the cranium, but H. von Meyer (428) states that the representation is inaccurate, and they seem to be based largely on Trematosaurus. The lateral-line canals occur in well-developed form on the skulls of the Stereo- spondylia. The sutures between the elements of the skull are usually clearly marked by smooth, narrow grooves. The lateral-line canals can always be distinguished from the sutural grooves by the shape of the bottom, being U-shaped in the former and V-shaped in the latter. The lateral-line canals also at times have their bottoms roughened by pits occurring in them ; the sutural grooves always have smooth bot- toms. The lateral-line canals are usually rather shallow and sometimes broad, with the edges of the grooves more or less perpendicular, but in Metoposaurus the canals are deep and the borders are sharply incised. The temporal canals in Anaschisma from the Triassic (49) of Wyoming are rep- resented by broken furrows. The portions preserved exhibit the usual downward tendency to unite with the infraorbital on the postorbital element. In its course forward from the epiotic the temporal canal cuts the squamosal. The supraorbital canal has an unusually deviating course in Anaschisma, but aside from the minor twists and curves it does not differ essentially from the same canal in other forms. It ends abruptly on the anterior end of the muzzle. In its course it gives off the ves- tige of an antorbital commissure which tends to join a vestige from the infraorbital canal. The jugal canal begins broadly at the very posterior edge of the skull as though it were continued, as it undoubtedly was, to the body of the animal. In its 36 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. course forward it joins the infraorbital canal on the jugal. The course of the infra- orbital is not unusual in any respect. There is no anterior commissure on the skull, nor is the occipital cross-commissure developed on either skull of the genus at hand. There are distinct evidences of an operculo-mandibular lateral-line canal on the mandibles. The canal enters the mandible on the surangular and passes forward around the mandible as described for Diplocaidus (477). Other members of the Stereospondylia, such as Mastodonsaurus, Metoposaurus, and Trematosaurus possess well-developed lateral-line canals, but the above descrip- tion fits, in a general way, the condition in all genera; and for our present purposes that will suffice. CHAPTER V. THE AMPHIBIA OF THE DEVONIAN AND MISSISS1PPIAN OF NORTH AMERICA. Evidences of the earliest land vertebrates are exceedingly scanty in the strata between the close of the Silurian and the opening of the Coal Measures, being repre- sented solely by footprints. In the Devonian our knowledge of the group is con- fined to a single footprint, and in the Mississippian to series of footprints from several localities. These have been described by Lea (371), Rogers (Geology of Pennsylvania, pt. 11, 1856, p. 831), Barrell (21), Dawson (223), and Branson (50). The last-named author has described a new species from the Mississippian of Giles County, Virginia. His description of the footprints, with a photograph of one of the series, are published herewith (plate 18, fig. 3). Branson (50) has given a resume of the knowledge of Mississippian Amphibia in North America. Thinopus antiquus Marsh, 1896. MARSH, Am. Jour. Science, H, p. 374, Nov. 1896, with figure. Type: Specimen No. 784, Yale University Museum. Horizon : Near top of Chemung, in the upper Devonian. [The] "specimen shows one vertebrate footprint in fair preservation, and with it part of another of the same series. These impressions are of much interest, both on account of their geological age and the size and character of the footprints themselves. The one best preserved [fig. 12] is nearly 4 inches in length, 2.25 inches in width, and was apparently made by the left hind foot. On the inner side in front of the heel, a portion of the margin is split off, and this may have contained the imprint of another toe. The other footprint was a short distance in front, but only the posterior portion is now preserved in the present specimen. It is probably the imprint of the forefoot. "The specimen [plate 18, fig. 4] . . . was . . . found in the town of Pleasant, one mile south of the Allegheny River, Warren County, Pennsylvania, by Dr. Charles E. Beecher, who presented it to Yale Fie 12-Copy of Mush's drawing of footprint of rv , • ,-,, • Thinopus antiquus, from the Devonian of Penn- Museum, where it still remains. sylvania. X 'A. "The geological horizon is near the top of the Chemung in the upper Devonian. In the same beds are ripple marks, mud cracks, and impressions of rain drops, indicating shallow water and shore deposits. Land plants are found in the same general horizon. Marine molluscs also occur, and one characteristic form (Nuculana) is preserved in the footprint slab" (Marsh). This still remains after nearly 20 years the only evidence of air-breathing verte- brates in the Devonian of the world. Dromopus ad uncus Branson. Branson, Jour. Geol., .win, No. 4, pp. 356-358, fig. '. «910- Type and other specimens in Oberlin College Museum. Horizon and type locality: Near the bottom of the Hinton formation in Giles County, Virginia. (Plate 18, fig. 3.) 37 38 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The following description of the shales and footprints are from Dr. Branson's paper cited above: "The Hinton shales, like the Mauch Chunk, seem to have been subaerial in origin and are made up for the most part of variegated shales interbedded with thin layers of argil- laceous, fine-grained sandstone. The footprints occur in fine-grained sandstone, and remains of land plants are not uncommon in the same beds. ' ' Twenty- two footprints made by one animal walking in a straight course were collected in a slab. They give the impression of having been made by a bipedal animal for part of the distance, but after the fourth print of the right foot impressions of the forefeet appear. The hindfeet had 5 digits, the middle digit being longest and the 2 inside of it being only slightly shorter and lying close together. Their outer ends were slender and flexible and usually curved inward toward the middle toe. The 2 outer digits formed wide angles with the middle one and were shorter than the inner ones. The second toe was webbed to within 8 mm. of the tip, the third toe to within 23 mm. of the tip. The impression of the web is well preserved in only one impression of the hindfoot. "The forefeet had 4 digits. The 3 inner digits were subequal in length, the 2 inner being more flexible and incurved near the ends. The outer digit is two- thirds as long as the second. The webbing extends about half the length of the digits. The heel impression is broader than that of the hindfoot. "Measurements of Dromopus aduncus Branson. Tip of toe to tip of toe in first prints 21 cm. After appearance of forefeet the impressions are the following distances apart : 165 mm., 40 mm., 85 mm., 70 mm., 80 mm., 40 mm., 150 mm., then back to 20 and 21 cm. mm. Length of hindfeet 60 Width of hindfeet 20 to 25 Length of forefeet 45" CHAPTER VI. A HISTORY OF THE CLASSIFICATION OF THE AMPHIBIA, WITH ESPECIAL REFERENCE TO THE SPECIES FROM THE COAL MEASURES. It has been necessary, in the course of the present study, to review thoroughly the classifications which have been proposed for the group. A classification of some sort is necessary for the proper grouping of the species which have been recovered from the Coal Measures deposits of this continent, and my reason for publishing this relatively dry material is that the classifications formerly proposed (469), as well as the one here given, may have a proper historical background. The review of the proposed systems of classification has been much facilitated by the discovery, in the University of Chicago, of some notes by the late Dr. George Baur on the " Stegocephali." The notes were not discovered until after the literature had been pretty thoroughly covered, and it was a source of some gratification, on comparing notes with those of Dr. Baur, to find but few omissions. Whether Dr. Baur had ever contemplated a work on the Stegocephala or not I have been unable to learn, but it is certain that he carefully and laboriously went through the litera- ture on the subject and copied by hand the classifications of each author from 1842 to 1895, together with other notes of interest on the structure, distribution, and phylogeny, including many tracings. The classifications given below are taken, in part, from his notes, although all references have been verified with the original sources. The first attempt to combine in classification the knowledge of the extinct and recent amphibians was made by Johannes Jacob von Tschudi in 1839 (574). Pre- vious to that time Goldfuss (295) and von Meyer (418) had described various species of salamanders and frogs from the Tertiary deposits of Switzerland, and these Tschudi considered in his following classification: A. Ranae. B. Cceciliae. a. Hylcc. a. Cacilue. b. Cystignatlu. C. Salamandrinae. c. Ranee. a. Pleurodcles. d. Ceratophrydes. b. Salamatidrce. c. Bombinatores. c. Tritones. f. Bufones. d. Tritonides. g. PipcB. D. Protoideae. Although the remains of Mastodonsaurus had been known and widely com- mented on for several years before Tschudi proposed this scheme, he does not include this genus in his classification of the Amphibia, for the reason that for nearly a quarter of a century after the discovery of the labyrinthodonts they were regarded as reptiles, even so eminent an authority as von Meyer (423) including them in his "System der fossilen Saurier." The view that the labyrinthodonts were 39 40 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. reptiles was at times disputed, but no one seemed to pay any attention to the argu- ment of Quenstedt in 1850 that "Die Mastodonsaurier im grunen Keupersand- stein Wurtemburgs sind Batrachier" (527), nor to the contention of Vogt (581) in 1854 that " Archegosaurus und alle Labyrinthodonten sind Amphibien, nicht Rep- tilien." In 1842 von Meyer (420) proposed to include all the early forms allied to the Mastodonsaurus in the " Labyrinthodontes. " His definition of the group follows: Labyrinthodontes : Saurier deren Zahn-Struktur jener ahnlich ist, welche in den nach prismat- ischer Art gebauten Saugethier-Zahnen wahrgenommen wird, u.s.w. I. Mastodonsaurus Jaeger. (Salamandroides Jaeger, Batrachosaurus Fitzinger, Labyrinth- odon Owen.) M. Jaegeri Meyer. II. Capitosaurus Miinster. C. arenaceus Miinst. C. robustus Meyer. III. Metopias Meyer. M. diagnosticus Meyer. Three years later von Meyer (423) proposed his "System der fossilen Saurier," where the extinct Amphibia are treated as follows: Labyrinthodontes. 1. Prosthopthalmi (Augen-hohlen in der vordern Halfte der Schadel-Lange) Metopias Meyer Keuper. 2. Mesopthalmi (Augen-hohlen in der mitte der Schadel) Mastodonsaurus Jaeger — Keuper, Muschelkalk. 3. Opisthopthalmi (Augen-hohlen in der hintern Halfte der Schadel-Lange) Capitosaurus Miinster Keuper. 4. Labyrinthodonten ungewisser Stellung. Labyrinthodon Owen Keuper. Xestorrhytias Meyer Muschelkalk. Odontosaurus Meyer Bunter Sandstein. Trematosaurus Braun Bunter Sandstein. No other classification was proposed for the extinct Amphibia for 15 years, when Owen (512) in 1859 proposed the new order Ganocephala and retained von Meyer's Labyrinthodontes under Labyrinthodontia. Owen's classification is as follows: Class — Reptilia. Order I. Ganocephala. Genera: Archegosaurus, Dendrerpeton, Raniceps. Order II. Labyrinthodontia. Genera: Mastodonsaurus, Anisopus, Trematosaurus, Metopias, Capitosaurus, Zygosaurus, Xestorrhytias. In his Paleontology published in 1861, Owen gives the same classification, but adds new genera. Huxley in 1863 (332) did not accept Owen's Ganocephala, but instead proposed the following: Labyrinthodontia. A. Archegosauria. Archegosaurus, Pholidogaster . B. Mastodonsauria. Mastodonsaurus, Labyrinthodon, Capitosaurus, Trematosaurus. In the same year Dawson proposed (208) the term Microsauria to include the genera Hylonomus, Dendrerpeton, and Hylerpeton, all known from the Carboniferous HISTORY OF CLASSIFICATION OF AMPHIBIA. 4 1 rocks of Nova Scotia. Two years later Cope proposed the new order Xenorhachia (105) for the reception of the form Amphibamus grandiceps from the Coal Measures of Illinois. He gave as the characters of this order cartilaginous vertebras and the absence of ribs. In 1866 Owen proposed (516) the most elaborate and comprehensive scheme of classification which had thus far been offered. His classification is as follows: Subclass Dipnoa. Order Ganocephala (extinct). Genera: Archegosaurus, Dendrerpeton, etc. Order Labyrinthodontia. Genera: Labyrinthodon, Rhombopholis, etc. Order Batrachia. Suborder Ophiomorpha. Family: Ccecilidae. Suborder Ichthyomorpha. Family: Proteidae, Salamandridae. Suborder Theriomorpha (Anura). Family 1. Aglossa. Family 2. Ranida. Family 3. Hylidce, Family 4. Bufonidce. Haeckel the same year proposed (312) an entirely different scheme of classifica- tion and in some respects more acceptable than Owen's. Haeckel's classification is as follows : Class Amphibia. Subclass I. Phractamphibia. Ordnung 1. Ganocephala. Genera: Archegosaurus, Dendrerpeton, Raniceps. Ordnung 2. Labyrinthodonta. Genera: Baphetes, Zygosaurus, Mastodonsaurus, Trematosaurus, Capitosaurus. Ordnung 3. Peromela. Subclass II. Lissamphibia. Ordnung 1. Socobranchia. Genera: Siren, Proteus, Menobranchus, etc. Ordnung 2. Sozura (Caudata). Genera: Cryptobranchus, Triton, Salatnandra. Ordnung 3. Anura (Ecaudata). Families: Aglossa, Bufonidce, Ranidce. This classification is further elaborated in the edition of 1895. Cope in 1868 proposed (no) the scheme of classification which was in use for some time, although it has since suffered some change. His classification follows: Batrachia. Order 1. Trachystoma. Order 2. Proteida. Order 3. Urodela. Order 4. Gymnophiona. Order 5. Stegocephali. Suborder Xenorhachia. Amphibamus grandiceps Cope. Suborder Microsauria. Genera: Pelion Wyman, Hylonomus Dawson, Pariostegus Cope, Dendrerpeton Owen, Hylerpeton Owen, Brachydectes Cope, Sauropleura Cope, CEsto- cephalus Cope, Molgophis Cope. Suborder Labyrinthodontia. Genera: Dictyocephalus Leidy, Centemodon Lea, Baphetes Owen, Eupelor Cope. 42 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Huxley in 1869 proposed (335) the following classification, which does not differ essentially from that proposed in 1863: Amphibia. Order 1. Urodcla. Order 2. Batrachia. Order 3. Gymnophiona. Order 4. Labyrinthodontia. Suborder Archegosauria. Suborder Mastodonsauria. The next classification of the extinct Amphibia of any importance was that devised by the committee (450) for the British Association in 1874. This committee was formed of Huxley, Harkness, Henry Woodward, Thompson, and Brigg, with Miall as secretary. This classification is, however, too cumbersome and has never come into general use, and indeed none but English authors have paid it a great deal of attention, although the contribution was a valuable one. The group Aistopoda, which was the ninth group proposed by the committee, has generally been accepted as the group represented by the snake-like forms. The committee's classification follows: Labyrinthodontia. Section I. Euglypta. Genera: Mastodonsaurus, Jaeger; Capitosaurus, Miinst. ; Pachygonia, Huxley; Tremato- saurus, Braun; Gonioglyptus, Huxley; Metopias, von Meyer; Iuibyrintliodon, Owen; Diadetognathus, Miall; Dasyceps, Huxley; Anthracosaurus, Huxley. Section II. Brachyopina. Genera: Brachyops, Owen; Micropholis, Huxley; Rhinosaurus, Waldheim; Bothriceps, Huxley. Section III. Chauliodonta. Genera: Loxomma, Huxley; Zygosaurus, Eich.; Melosaurus, Meyer. Section IV. Arthroodonta. Genera: Batrachiderpeton, Hancock and Atthey; Pteroplax, Hancock and Atthey. Section V. An uncharacterized group. Genera: Pholidogaster, Huxley; Ichthyerpelon, Huxley; Pholiderpelon, Huxley. Section VI. Archegosauria, von Meyer. Genera: Archegosaurus, Goldfuss. Section VII. Heleothrepta. Genera: Lepterpeton, Huxley. Section VIII. Nectridea. Genera: Urocordylus, Huxley; Keraterpeton , Huxley. Section IX. Aistopoda. Genera: Ophiderpeton, Huxley; Dolichosoma, Huxley. Section X. Microsauria, Dawson. Genera: Dendrerpeton, Owen; Hylonomus, Dawson; Hylerpeton, Owen. The schemes of classification used for the next 10 years did not depart in any appreciable degree from those already given. In 1875 Cope, in his Check-list of North American Batrachia and Reptilia (120), with a systematic list of the higher groups, published the following classification as being "adopted provisionally by the Smithsonian Institution": Class Batrachia. Order Anura. (Anura, Dumeril; Salientia, Merrem, Gray.) Raniformia. (Raniformia, Cope, Nat. Hist. Rev., v, 114, 1865.) Families: Ranidce, Colostheidce. Firmisternia. (Bufonoid Raniformia, Cope, Jour. Acad. Nat. Sci., Philadelphia, n.s., vi, 190, 1867.) Families: Dendrobatidee, Phryniscidce, EngystomidcB, Breviceptidce. HISTORY OF CLASSIFICATION OF AMPHIBIA. 43 Class Batrachia — Continued. Order Anura — Continued. Gastrechmia. (Gastrechmia, Cope, Jour. Acad. Nat. Sci., Philadelphia., n.s., vi, 198, 1867.) Family: Hemisidce. Bufoniformia. (Bufoniformia, Dumeril et Bibron, partim; Cope, partim.) Families: Rhinophrynidw, Bufonida, Batrachophrynidw. Aglossa. Family: Pipidoe. Odontaglossa. Family: Dactylethrida. Arcifera. (Arcifera, Cope, N. H. Rev., v, 104, 1865.) Families: Cystignathidce, Hemiphractidce, Hylidce, Scaphiopidce, Pelodytida, Astero- phrydidce, Discoglossidee. Order Stegocephali. (Stegocephali Cope, Proc. Acad. Nat. Sci., Philadelphia, 1868, 209.) Labyrinthodontia . Families: Baphetidce Cope, Anthracosauridw Cope. Ganocephala. Family : Colosteidce Cope. Microsauria. Families: Phlegethontiidce Cope, Molgophidw Cope, Ptyoniida; Cope, Tuditanidx Cope, Peliontidce Cope. Order Gymnophiona. (Gymnophiona Muller.) Family: Caciliida Gray, 1850. Order Urodela. Families: Pleurodelidce Gray, 1858; Salamandridw Gray, 1858; Hynobiidce Cope, 1866; Desmognathidce Cope, 1866; Thoriidm Cope, 1869; Plethodontidce Cope, 1866; Amblystomidee Cope, 1866; Menopomidm (Protonopsidce Gray, 1850), Amphiumidm Cope, 1866; Cocytinidce Cope. Order Proteida. Family: Proteida Gray, 1850. Order Trachystomata. Family: Sirenidw Gray, 1850. In 1885 Cope proposed 2 new orders, which he arranges with the other orders already known, as follows: Order I. Rachitomi. Order V. Urodela. Order II. Embolomeri. Order VI. Trachystomata. Order III. Stegocephali. Order VII. Anura. Order IV. Proteida. Batrachia. The new order Rachitomi was to include forms like Eryops and the new order Embolomeri was to include forms like Cricotus; the other orders were as they had been given before. Zittel in 1888 proposed (642) the next classification of any note in his "Hand- buch der Paleontologie," where it stands as follows: Classe Amphibia. Ordnung 1. Stegocephali. Ordnung 3. Urodela. Unterordnung 1. Lepospondyli. Unterordnung 1. Ichthyoidea. Familie 1. Branchiosauridce Fritsch. Familie 1. Phanerobraiwlua. 2. Microsauria Dawson. " 2. Cryptol>ra>ulihi. " 3. Aistopoda Miall. Unterordnung 2. Salamandrina. Unterordnung 2. Temnospondyli. Ordnung 4. Anura. Genera: Archegosaurus, Eryops, etc. Unterordnung r. Phaneroglossa. Unterordnung 3. Stereospondyli. Familie 1. Ranidw. Familie 1. Gastrolepidoti. " 2. Bufonida. 2. Labyrinthodonta. " 3- Cystignathida Cope. Ordnung 2. Cceciliae. " 4- PelobaXidm Boul. " 5. Discoglossidee Cope. " 6. Palaobatraihidce Cope. 44 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Lydekker (393) in the next year proposed a system of classification which did not depart widely from that proposed (450) by the committee of the British Associ- ation for 1874. Lydekker's classification is as follows: Class Amphibia. Order I. Labyrinthodontia. Order II. Apoda. Suborder 1. Branchiosauria. Order III. Caudata. Family Protritonidce. Family Hylceobatrachid.ee. " Apateonidce . " Sirenidw. Suborder 2. Aistopoda. " Proteidoe. Family Dolichosamatidce . " Amphiumidce. Suborder 3. Microsauria. " Salamandridce. Family Urocordylidce. Order IV. Ecaudata. " Limner petidce. Family Discoglossidce. " Hyloplesionidce. " Pelobatidce. " Microbrachidcz. " Palceobatrachidce. Suborder 4. Labyrinthodontia vera. " Cystignathidce. Family Archegosauridce. " Ranidce. " Diplospondylidce. " Nyraniidce. " Dendrer petidce. " Anthracosauridce . " Mastodonsauridce. Uncertain family, Eosaurus. In 1890 Doederlein proposed a scheme of classification which is notable on account of the peculiar relations which it expresses between the groups — relations which, in reality, do not exist. His classification is as follows: Class Amphibia. Ordnung I. Stegocephali. A. Microsauria. Unterordnung 1. Branchiosauri. Genera: Branchiosaurus, Dawsonia, Melanerpeton, Pelosaurus. Unterordnung 2. Sauromorphi. Familie 1. Hylonomidce. " 2. Nectridce. " 3. Aistopodidce. B. Ganocephala. Unterordnung 1. Rhachitomi. * 2. Embolomeri. " 3. Labyrinthodontia. Ordnung II. Urodela. Ordnung III. Gymnophiona. Ordnung IV. Anura. In 1890 Lydekker used the same classification, with minor changes, which he had used in his Paleontology. Credner, who wrote at about the same time (193), fol- lowed Zittel's classification. Zittel in 1895 merely repeated his former classification. In 1898 appeared Smith Woodward's Paleontology, where the following scheme is adopted : Class Batrachia. Order I. Stegocephalia. Order II. Gymnophiona. Suborder 1. Branchiosauria. Order III. Caudata. 2. Aistopoda. Order IV. Ecaudata. " 3. Microsauria. 4. Labyrinthodontia. HISTORY OF CLASSIFICATION OF AMPHIBIA. 45 Hay's Catalogue of Fossil Vertebrata of North America contains the next scheme for the classification of the Amphibia which pays especial attention to the extinct forms. His classification is as follows: Class Batrachia Macartney, 1802. Order Stegocephali Cope, 1868. Suborder Microsauria Dawson, 1863. Family Protrilonidm Lydekker, 1889. Genera: Amphibamus Cope, Pelion Wym. Family Molgophidw Cope, 1875. Genera: Phlegethontia Cope, Molgophis Cope. Family Hylonomida Fritsch, 1883. Genera: Hylonomus Dawson, Smilerpeton Dawson, Hylcrpeton Owen, Fritschia Dawson; Brachydectes Cope. Family Ptyoniida Cope, 1875. Genera: Keraterpeton Huxley, Gistocephalus Cope, Piyonius Cope, Cienerpeton Cope. Family Tuditanida Cope, 1875. Genera: Tudilanus Cope, Cocytinus Cope. Family Diplocaulidce Cope, 1881. Genus: Diplocaulus Cope. Lepospondylous Genera of uncertain position: Amblyodon Dawson, Hyphasma Cope, Eurythorax Cope, Thyrsidium Cope, Pleuroptyx Cope, Cercario- morphus Cope. Suborder Apoecospondyli Hay, 1902. Family Dendrerpetontidce Fritsch, 1889. Genera: Dendrerpeton Owen, Baphetes Owen, Platystegos Dawson. Family Sauropleuridce Hay, 1902. Genera: Sauropleuta Cope, Leptophr actus Cope. Family Archegosauridce. Genera: Trimerorhachis Cope, Dissoropkus Cope. Family Cricotidce Cope, 1884. Genera: Cricotus Cope. Family Anthracosaurida. Genus: Eosaurus Marsh. Family Eryopidce Cope, 1882. Genera: Eryops Cope, Ichthycanthus Cope, Zatrachys Cope, Anisodexis Cope, Acheloma Cope. Family Mastodonsaurida Huxley, 1863. Genera : Mastodonsaurus Jaeger, Eupelor Cope, Pariostegus Cope, Dictyocephalus Leidy. Order Urodela. Genera: Scapherpelon Cope, Hemitrypus Cope. Order Salientia Laurcnti, 1768. Family Ranidce. Genera: Rana Linne, Eobatrachus Marsh. This classification given by Hay is only for the forms which occur in North America. CHAPTER VII. CLASSIFICATION OF AMPHIBIA ADOPTED IN THIS WORK, AND A LIST OF THE COAL MEASURES AMPHIBIA FROM NORTH AMERICA. A few words of explanation will be necessary for an understanding of the follow- ing classification. The term Amphibia Linne, 1758, is, according to Stejneger (550), the correct term for the entire group, and this term is adopted. The term Stego- cephala, formerly used as a group name for the entire Carboniferous, Permian, and Triassic Amphibia, regardless of structure, has been retained as a third subclass. The choice, so far as priority is concerned, between Labyrinthodontia and Stego- cephala, is not easy. The terms, however, imply different structures. The laby- rinthodonts proper have stereospondylous vertebrae, while the Stegocephala have either temnospondylous or stereospondylous vertebrae; so the latter term has been adopted. The ordinal terms are those which have been used previously as subordinal, sectional, or family names, with the exception of the new ordinal term "Diplocau- lia" (477). The term Branchiosauria is well-established, and it is here retained with the definitions previously given. The same may be said for the Microsauria, although Dawson first (208) regarded it as a family, though giving the term an ordi- nal form. The Aistopoda are not entitled to consideration as a group for reasons which are given subsequently. The Temnospondylia and the Stereospondylia, the Embolomeri and the Rachitomi may or may not be good group names, but they have priority, so far as our knowledge of structure goes. They have been retained in their original meanings. They have been variously regarded as sections, superf ami- lies, groups, and subfamilies. The following list of species is arranged according to the proposed scheme of classification : Class Amphibia Linne, 1758. Devonian to Recent. Subclass Euamphibia Moodie, 1909. Coal Measures to Recent. Order Branchiosauria Lydekker, 1889. Coal Measures to Permian. Family Branchiosauridce Fritsch, 1879. Micrerpeton caudatum Moodie, Mazon Creek. Eumicrerpeton parvum Moodie, Mazon Creek. Mazonerpeton longicaudatum Moodie, Mazon Creek. Mazonerpeton costatum Moodie, Mazon Creek. (?) Sparodus sp. indet. Dawson, Nova Scotia. Order Caudata Dumeril, 1806. Coal Measures to Recent. Suborder Proteida Cope, 1868. Coal Measures, Eocene, and Recent. Family Cocytinidoe Cope, 1875. Cocytinus gyrinoides Cope, Linton, Ohio. Erierpeton branchialis Moodie, Mazon Creek. Hyphasma laevis Cope, Linton, Ohio. Order Diplocaulia Moodie, 191 2. Coal Measures to Permian. Family Diplocaulidce Cope, 1881. Diplocauhts salamandroides Cope, Salt Fork, Illinois. (?) Order Salientia Laurenti, 1768. Coal Measures (?) to Recent. Family Peliontida Cope, 1875. Pelion lyelli Wyman, Linton, Ohio. 46 CLASSIFICATION OF AMPHIBIA AND LIST OF COAL MEASURES SPECIES. 47 Class Amphibia — Continued. Subclass Lepospondylia Zittel, 1887. Coal Measures. Order Microsauria Dawson, 1863. Coal Measures. Family Hylonomidce Fritsch, 1883. Hylanomus latidens Dawson, Nova Scotia. Hylonomus lyelli Dawson, Nova Scotia. Hylonomus multidens Dawson, Nova Scotia. Hylonomus wymani Dawson, Nova Scotia. Smilerpeton aciedentatum Dawson, Nova Scotia. Hylerpeton dawsonii Owen, Nova Scotia. Hylerpeton intermedium Dawson, Nova Scotia. Hylerpeton longidentatum Dawson, Nova Scotia. Fritschia curtidentata Dawson, Nova Scotia. (f) Amblyodon problematicum Dawson, Nova Scotia. Family Tuditanidce Cope, 1875. Tuditanus punctulatus Cope, Linton, Ohio. Tuditanus brevirostris Cope, Linton, Ohio. Tuditanus minimus Moodie, Cannelton, Pa. Tuditanus longipes Cope, Linton, Ohio. Tuditanus walcotti Moodie, Linton, Ohio. Erpetosaurus acutirostris Moodie, Linton, Ohio. Erpetosaurus minutus Moodie, Cannelton, Pa. Erpetosaurus obtusus Cope, Linton, Ohio. Erpetosaurus radiatus Cope, Linton, Ohio. Erpetosaurus sculptilts Moodie, Cannelton, Pa. Erpetosaurus tabulatus Cope, Linton. Ohio. Erpetosaurus tuberculatus Moodie, Linton, Ohio. Odonterpeton triangularis Moodie, Linton, Ohio. Family Stegopidw Moodie, 1009. Stegops divaricata Cope, Linton, Ohio. Family Urocordylidce Lydekkcr, 1890. Diceratosaurus punctolineatus Cope, Linton, Ohio. Diceratosaurus IcBvis Moodie, Linton, Ohio. Diceratosaurus robustus Moodie, Linton, Ohio. Eoserpeton tenuicorne Cope, Linton, Ohio. Family Amphibamida Moodie. Amphibamus grandiceps Cope, Mazon Creek. Amphibamus thoracatus Moodie, Mazon Creek. Cephalerpeton ventriarmatum Moodie, Mazon Creek. Family Nyraniidce Lydekker, 1889. Ichthyerpcton squamosum Moodie. Linton, Ohio. Cercariomorphus parvisquamis Cope, Linton, Ohio. Family Ptyoniida: Cope, 1875. Ptyonius nummifer Cope, Linton, Ohio. Ptyonius marshii Cope, Linton, Ohio. Ptyonius vinchellianus Cope, Linton, Ohio. Ptyonius pectinatus Cope, Linton, Ohio. Ptyonius serrula Cope, Linton, Ohio. CEstocephalus remex Cope, Linton, Ohio. Qistocephalus reciidens Cope, Linton, Ohio. Thyrsidium jasciculare Cope, Linton, Ohio. Family Ichthycanthidce Moodie. Ichthycanthus ohiensis Cope, Linton, Ohio. Ichthycanthus platypus Cope, Linton, Ohio. Family Molgophidee Cope, 1875. Molgophis macrurus Cope, Linton, Ohio. Molgophis brevicostatus Cope. Linton, Ohio. Molgophis wheatleyi Cope, Linton, Ohio. Erpetobrachium mazonensis Moodie, Mazon Creek. Pleuroptyx clavatus Cope, Linton, Ohio. Phlegethontia linearis Cope, Linton, Ohio. Phlegethontia serpens Cope, Linton, Ohio. 48 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Class Amphibia — Continued. Subclass Lepospondylia — Continued. Order Microsauria — Continued. Family Sauropleuridas Hay, 1902. Sauropleura digitata Cope, Linton, Ohio. Sauropleura newberryi Cope, Linton, Ohio. Sauropleura scutellata Newberry, Linton, Ohio. Sauropleura pauciradiata Cope, Linton, Ohio. Sauropleura longidentata Moodie, Linton, Ohio. Sauropleura foveata Cope, Linton, Ohio. Sauropleura (Anisodexis) enchodus Cope, Linton, Ohio. Sauropleura sp., Linton, Ohio. Ctenerpeton alveolatum Cope, Linton, Ohio. Saurerpeton latithorax Cope, Linton, Ohio. Leptophraclus obsoletus Cope, Linton, Ohio. Leptophractus dentatus Moodie, Linton, Ohio. Leptophraclus lineolalus Cope, Linton, Ohio. Order Temnospondylia Zittel, 1887. Coal Measures and Permian. Family Cricotidas Cope, 1884. Spondylerpeton spinatum Moodie, Mazon Creek, 111. Family Eryoptdce Cope, 1882. Eryops sp. indet., Pitcairn, Pa. Family Macrerpetidce Moodie, 1909. Macrerpeton huxleyi (Cope), Linton, Ohio. Macrerpelon deani Moodie, Linton, Ohio. Family Anthracosauridce Cope, 1875. Eosaurus acadianus Marsh, Nova Scotia. Eobaphctes kansensis Moodie, Kansas. Baphetes planiceps Owen, Nova Scotia. Baphetes minor Dawson, Nova Scotia. Dendrerpeton acadianum Owen, Nova Scotia. Dendrerpelon oweni Dawson, Nova Scotia. (f)Platystegos loricatum Dawson, Nova Scotia. Genera and species of uncertain position : Amblyodon problematicum Dawson, Nova Scotia. Proterpeton gurleyi Moodie, Illinois. Brachydectes neivberryi Cope, Linton, Ohio. Order Stereospondylia Zittel, 1887. Coal Measures (?) and Triassic. Family Mastodonsauridm Huxley, 1863. Mastodonsaurus sp. indet., Kansas. The above list of species is interesting in that it shows the diversity of structure among the Coal Measures Amphibia. There are at present 88 species known, many of them incompletely, divided among 1 7 families and 5 orders. The majority of the species are from the Linton Coal Measures, there being 50 species described or indi- cated from these beds. The Mazon Creek shales have furnished 10 species; Nova Scotia 18 species; the remainder of the species are from various localities. CHAPTER VIII. DEFINITION OF THE CLASS AMPHIBIA L1NNE, 1758, DEVONIAN TO RECENT. (World-wide distribution.) Cold-blooded vertebrates; aquatic or partially terrestrial in habit; body scaled or naked or partly covered with bony or horny plates; abdomen sometimes protected by closely packed scutes, scales, or rods ; skull completely roofed over or with a single vacuity ; pterygoid-palatine arch complete or wanting ; stapes always present ; two occipital condyles, sometimes cartilaginous ; skull bones pitted and grooved by the lateral-line canals, or smooth and lateral-line canals wanting; parasphenoid well developed ; palatine vacuities, large, small, or absent ; basioccipital partly or entirely cartilaginous; sclerotic plates present or absent ; mouth always terminal ; teeth sharply conical, smooth, or plicated, with walls sometimes extremely complicated by the infolding of the dentine and enamel. Vertebrae proccelous, opisthoccelous, amphiccelous, amphiplatyan, temnospondylous, stereospondylous, or cartilaginous; notochord often persistent ; column divisible into cervical, dorsal, and caudal series ; cervical series, so far as known, always short ; dorsal region long or short ; a single sacral or two ; caudal series short, very long, or absent. Pectoral girdle composed of an osseous scapula, cleithrum, clavicle, interclavicle, and coracoid with various relations ; sternum undeveloped ; pectoral girdle of membrane bones; in Triassic forms producing the effect of a plastron on account of the high development of the clavicles and inter- clavicle. Pelvis usually composed of an osseous ilium and ischium; pubis when osseous surrounded by large amounts of cartilage, usually cartilaginous, sometimes calcified. Limbs ambulatory, nata- tory, or wanting ; limb bones composed either entirely of perichondrium or of perichondrium and a small amount of endochondrium ; radius and ulna, and tibia and fibula free or fused. Digits 3 to 5, usually 4 for the hand and 5 for the foot. Terminal phalanges sometimes clawed. Carpus and tarsus osseous or cartilaginous, usually the latter. Ribs never attached to a sternal apparatus, single or double headed or intermediate, long and curved or short and straight. Articulation with vertebral column inter- or intra-central. Respiration both branchial and pulmonary; branchiae persistent and osseous in some forms. Development by metamorphosis either in the egg membrane, on the back of the mother, or in the water. No amnion or allantois. Heart with a single ventricle and 3 or 4 pairs of aortic arches; postcava always present in the recent forms. DEFINITION OF SUBCLASS EUAMPHIBIA, MOOD1E, 1909. COAL MEASURES TO RECENT. (World-wide distribution.) Moodie, Trans. Kans. Acad. Sci., 1909, p. 243. Moodie, Geol. Mag., n. s., Dec. v, vol. VI, p. 220, May, 1909. The present group was established for the reception of the Branchiosauria and their descendants, the Caudata, with the related forms, the Apoda. The Salientia are included provisionally, since there is no evidence of the origin or relationship of this group of animals to other Euamphibia save that they have attained the same stage of evolution. They are in no way closely related to any known group of Amphibia, recent or extinct, but they stand on the same plane of development as the Caudata and present similar structures, i.e., a single ventricle in the heart, external branchiae in the young, a glandular skin, perichondral bone, and a large parasphenoid. The origin of the Salientia is a puzzle and must remain so until further paleontological evidence is forthcoming. Wyman, Cope, and the writer have all remarked on the similarity of structure between the Salientia and the single known specimen of Pelion lyelli Wyman from the Linton, Ohio, Coal Measures. 49 50 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The subclass Euamphibia may be defined as follows: Aquatic or terrestrial Amphibia; development by metamorphosis; external branchiae present in the young; bones almost entirely perichondral ; carpus and tarsus never ossified ; osseous pubis absent; vertebrae usually amphiccelous with persistent notochord; ribs short and straight or flat and slightly curved, or absent; digits 4 in the hand and 5 in the foot; skull never grooved or pitted, nor cut by the lateral-line canals; lateral-line organs present in the skin ; sclerotic plates present or absent ; tail long and flattened or absent. Ribs in Triton walthi are secondarily long and curved. DEFINITION OF THE ORDER BRANCHIOSAURIA LYDEKKER, 1890. COAL MEASURES AND PERMIAN OF NORTH AMERICA AND EUROPE. Lydekker, Cat. Fossil Reptilia and Amphibia, pt. iv, p. 208, 1890. Extinct, salamander-like amphibians, with broad, obtusely rounded cranium; external branchiae present in young; sclerotic plates present; bones of the cranium not ornamented with deep pits and grooves nor cut by the lateral-line canals, though sometimes ornamented with slight scorbiculations ; notochord always persistent; vertebrae cartilaginous (in caudal region) or but partially ossified, the ossification being entirely perichondral ; a single sacral vertebra ; transverse process of vertebrae large in dorsal region; ribs always short, straight, and heavy, present throughout the length of the vertebral column and borne on the transverse processes centrally ; caudal region of moderate length with elongate fleshy tail ; usually 20 presacrals, of which 4 or 5 may be considered cervicals; limbs natatory and always present, well developed; elements of the appendicular skeleton composed entirely of peri- chondrium; carpus and tarsus cartilaginous; digits 4 in the hand and 5 in the foot; phalangeal formula for the hand usually 2-2-3-2 and for the foot 2-3-4-3-2 ; distal phalanges not clawed; abdomen covered with closely packed corneous scutes or scales; body naked or covered with minute horny scales; median and dorsal lateral lines present on the posterior part of body and on tail. CHAPTER IX. THE AMERICAN COAL MEASURES BRANCHIOSAURID/E. Definition of the Family Branchiosaurid^b Fritsch, 1879. Coal Measures and Permian of North America and Europe. Fkitsch, Fauna der Gaskohle, 13d. I, p. 69, fig. 30, 1879. Lydekker, Cat. Fossil Rcptilia and Amphibia, pt. iv, p. 210, 1890 (Protritonidae). Stegocephalic, salamander-like animals, with broad, anteriorly truncate skull. Teeth smooth with large pulp-cavity. The parasphenoid narrowed anteriorly, pos- teriorly expanded to a shield-shaped plate. Vertebrae with the notochord persistent and intravertebrally expanded. Pelvis well developed, the ilium and ischium osse- ous with large cartilaginous margins, the pubis unknown, possibly hyaline cartilage. Ribs short, straight, present on almost all vertebras. Skin with delicately ornamented scales. Eyes with sclerotic plates. Palatal elements toothless or with small tooth- like tubercles on pterygoids and palatines. Ventral armature on throat, chest, and abdomen, extending on to the limbs, consisting of small delicate scutellae arranged in a chevron pattern. The above definition is modified from Fritsch (Fauna der Gaskohle, Bd. I, p. 69, 1879). The North American species are: (?) Sparodus sp. indet. Dawson, Micrerpeton caudatum Moodie, Mazonerpeton longicaudatum Moodie, Mazonerpeton costatum Moodie, Eumicrer peton parvum Moodie. Genus MICRERPETON Moodie. Moodie, Jour. Geol , 17, p. 39, figs. 1 to 6, 1909. Type Micrerpeton caudatum Moodie. The genus Micrerpeton, of which the single species is described below, was the first evidence of the occurrence of the Branchiosauria in America. There have been three other genera referred to the Branchiosauria from North American deposits, but there is good evidence that none of them belong there. The genus Amphibamus was originally referred to the Xenorhachia by Cope (105, pp. 134-137) on account of the supposed cartilaginous condition of the vertebrae and the absence of ribs. Later he abandoned this order and placed the form in the Branchiosauria, where it was retained by Zittel (642). Recently Hay has shown (316), and I am able to corrob- orate his statement, that ribs are present in the species, and that they are long and curved, not at all like the short ribs of the true Branchiosauria. These long, curved ribs undoubtedly exclude Amphibamus from the Branchiosauria and indicate its close affinities with the Microsauria. The genus Pelion has also been referred to this order on purely negative evidence (642, p. 375). This genus is excluded from the Branchiosauria by the well-ossified condition of the limb bones, in which the endo- chondral ossification is seen to be well developed, a condition not found, so far, among the true Branchiosauria. The form of the head and the elongate hind limb would also tend to exclude the genus from the Branchiosauria. In the Branchiosauria 51 52 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. the fore limb is usually larger than the hind limb, the reverse of which is the case in Pelion. The genus Sparodus, as it occurs in North America, is uncertain. It is indicated by remains which are almost impossible of determination. The genus Micrerpeton may be distinguished from other known Branchiosauria by the large size and anterior position of the orbits, absence of a posterior table to the skull, the short, heavy limb bones, the slender ilium, and the expanded, elongate, and laterally compressed tail. The genus may be defined as follows : Small forms, the known representative attaining a length of less than 2 inches ; head broad and short ; sclerotic plates present ; interorbital space less than the least diameter of the orbit ; occiput concave; pineal foramen in the line which cuts the posterior edge of the orbits; teeth small, pleurodont denticles; presacral vertebrae 20 or 21, of which probably 5 are cervical; 1 sacral; ribs short, straight, and heavy, central; scapula ovoid ; limbs stumpy and heavy, fore limb exceeding the hind in size ; endochondral ossifications distinctly absent; tail long, expanded, and flattened, probably provided with a thin expanded membrane; body covered with minute, ovoid or rounded scales which are ornamented with concentric lines ; color markings vertical to the long axis of the body and abundantly present on the tail ; lateral-line organs repre- sented by the dorsal and median lateral lines on the tail, the sensory pits probably occurring in specialized darkened scales. Coal Measures of Mazon Creek shales near Morris, Grundy County, Illinois. Micrerpeton caudatum Moodie. Moodie, Jour. Geol., 17, p. 39, figs. 1-6, 1909. Type: Specimen No. 12,313, Walker Museum, University of Chicago. Horizon and locality : Mazon Creek shales, near Morris, Illinois. The species is represented by very complete remains (plate 2), which are pre- served on opposite halves of a nodule. The specimen was collected many years ago by Mr. W. F. E. Gurley at Mazon Creek, but it has never before been studied, although Dr. Newberry examined it and said in a note that Professor Cope should see it. Unfortunately Cope did not see it and it lay unknown for more than a quar- ter of a century. I am indebted to Dr. Stuart Weller for calling my attention to the specimen, as well as for the privilege of describing it. The specimen is exceptionally perfect (plate 25, fig. 4). Nearly all the skeletal elements are present, and the general contour of the body, the character of the dermal covering, the color-markings, the lateral-line system, and many other features of interest have been detected. Such completeness of preservation is very uncommon even among the remains obtained from this locality. In this case the entire form was preserved, but the collector, in cracking the nodule, lost the chips containing the feet, so that only portions of the limbs remain. It is thus impossible to deter- mine the phalangeal formula, but the feet were probably like those of Branchio- saurus amblystomus Credner, as given by Credner, to which species the present form is closely allied and indeed must be placed in the same family with Branchiosaurus, Pelosaurus, and Melanerpeton. The remains here described represent a small, salamander-like form, and they are among the earliest geological evidence of the group, which, without doubt, gave PLATE 2 Orbits blackened by pigment L-.. Pineal eye ■ ..Mandible Interclavicle Spines of vertebrae Centra of vertebje Tibia Femur , -' Impression of the fleshy and. membranous parts of the tail. Banded color markings on tail Impressions of cartilagi- nous vertebra _'— . Dorsal lateral line Median lateral line Drawing X 3. 5. of type specimen of Micrerpeton caudatum Moodie, from the Coal Measures of Mazon Creek, Illinois, showing skeletal elements, form of head and tail, the lateral-line sense organs, banded color- markings, and ventral scutellse. On the edges of the tail impression are indications that in life the tail had a thin fold of skin above and below the fleshy portion, much as in the larva; of . I ni/dys/oitia at the present day. THE AMERICAN COAL MEASURES BRANCHIOSAURID^E. 53 rise to the modern salamanders. The parts preserved in the specimen are: the complete outline of the head with most of the cranial elements clearly distinguish- able, as well as the black pigment of. the choroid; the entire vertebral column, in- cluding pits in the tail region, where the vertebrae were without doubt entirely car- tilaginous; parts of the pectoral girdle; the ilium; the left humerus; the ventral scutellation; the ribs of one side of the body and indications of ribs on the other; portions of both hind limbs; and a complete impression of the fleshy tail. On this impression of the tail are preserved small, horny scales, transverse color-markings, and the distinct impressions of the lateral-line system. The bones of Micrerpeton caudatum, as in so many of the fossils from this local- ity, have been replaced by a white, friable mineral which is probably kaolin. The animal is preserved on its back and it is thus illustrated from the ventral side. The entire length of the animal is only 49 mm., of which the tail occupies nearly half. The head has much the same shape as in the species of Branchiosaurus described and figured by Fritsch (251), Credner (181), and Theyenin (568). The eyes occupy relatively the same position as in that genus. The orbits are very large and broadly Fig. 13. — Restoration of Micrerpeton caudatum, a branchiosaur from the Coal Measures of Illinois. X 2. oval. Within the borders of the rim the stone is blackened as though by the black pigment of the iris, such as Cope has described in Amphibamus. Under a rather high power of magnification the cranial bones are seen to be represented by mere flakes of white mineral matter. The sutures separating the cranial elements are distinctly preserved on the main half of the nodule. The openings of the skull are five — the two orbits, the two minute nostrils, and the pineal foramen. A median suture separates the skull into halves and the pineal foramen lies slightly anterior to the posterior third of its length. The boundaries of the premaxillas are not distinct, but they are very small elements and form the inner border of the nostrils, which are clearly indicated by bosses of stone. The nasal element is nearly square and lies anterior to the frontal, which it borders broadly. The parietal is about the same size as the frontal and it apparently forms a portion of the inner border of the orbit, although this is not assured. The parietal is elongate and unites posteriorly with the postparietal. The postparietal, with the tabulare and the squamosal, form the posterior boundary of the skull, and they are hence not unlike the same elements in other Stegocephala. The prefrontal forms the anterior border of the orbit. The lacrimal has not been detected. The maxilla 54 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. is elongate and forms the antero-lateral border of the skull. The jugal forms an important element in the lateral border of the cranium and joins the quadratojugal posteriorly. The postfrontal is triangular and with the postorbital forms the pos- terior border of the orbit. Both of the elements are acuminate posteriorly, although the suture between them is indistinct, and they inclose between their posterior acumi- nations an anterior projection of the supratemporal. The squamosal has the usual relations and borders the supratemporal laterally. The latter element forms the quadrate angle of the cranium. The entire length of the vertebral column is preserved, although the nature and structure of its elements can not be determined. The impressions of a few of the vertebrae show that some of the centra were amphiccelous, but other than this nothing is definite. The cavities which the centra occupied were filled by the white mineral matter and the force of the blow which cracked the nodule destroyed the form of the mold. It is possible that where the mineral matter has filled the cavities the centra were osseous or partly cartilaginous, and where the cavities were unfilled the centra were entirely cartilaginous. The length of the vertebral column from the base of the skull to the last impression of a cartilaginous centrum is 33 mm. The number of centra between the sacral vertebra and the skull is 20 as they are preserved, but there may have been one more, the atlas. Fritsch has represented 21 in his restoration of Branchiosaurus salamandroides, and this is a further indica- tion of an affinity between the two genera, although Credner has represented 26 pre- sacral vertebras in Branchiosaurus ambly stomas. The presacral vertebrae are thus seen to vary within narrow limits, but the number of presacrals is near 20, and this may be taken as typical. It is interesting to notice that in modern forms of the salamanders the presacral vertebrae number about 20. There is but a single sacral centrum in Micrerpeton. The sacral rib has not been detected, but it is restored after the condition found in Branchiosaurus. The right femur partially covers the sacral vertebra, and its structure can not be determined. I count impressions of 17 caudal centra, of which at least 12 may have been partially ossified. In the cervi- cal region there are distinct impressions of transverse processes on at least 5 verte- brae, and this number is assigned to the neck, although it is by no means certain that this is the correct number. The neck was at least short, if we may judge from the position of the remains of the pectoral girdle. No cervical ribs are definitely determined. There is a short rib lying between the fifth and sixth vertebrae, but to which it belongs is uncertain. There are impressions of 10 ribs preserved on one side of the vertebral column and one on the other side. They are short, straight, and heavy, as are the same ele- ments in Branchiosaurus. This character alone is sufficient to place Micrerpeton among the Branchiosauria, since no such ribs are known in other groups of the extinct amphibians. The ribs preserved lie next the seventh to the seventeenth vertebrae on the left side, and there is one on the right side which may belong to either the fifth or sixth vertebra. They are central in their attachment, and in this they agree well with the mode of rib attachment in the modern salamanders. All of the ribs are single-headed and are composed, for the most part, of perichondral tissue. THE AMERICAN COAL MEASURES BRANCHIOSAURID/E. 55 The position of the ribs in the matrix inclined backwards, and, making a small angle with the vertebral column, is very suggestive of the condition in Branchiosaurus. The pectoral girdle is represented by three distinct elements of the left side, which are identified as scapula, clavicle, and coracoid, following the nomenclature given by Woodward (631), although Credner (186) would name them otherwise. The scapula is represented by an ovoid fragment lying next to the vertebral column. The clavicle was probably spatulate, as in Melanerpeton, but the inner end of the element is not visible. The coracoid is represented by its outer end only, and its inner pointed extremity is not visible. The interclavicle has not been detected. The humerus lies somewhat to one side of the pectoral girdle, as if there had been a large amount of epiphyseal cartilage. Its position may be due to post-mortem shifting, but there is little other evidence of any movement after deposition. The humerus is a large, heavy bone in comparison with the rest of the skeleton. It is expanded at each end, and its ends show concavities, proving that the bone is formed principally of perichondral tissue, as would be expected from such an early Branch- iosaurian. The endochondrium has not yet developed in this form, which is evi- dently adult. There is no other element of the arm present. There is but a single element of the pelvis preserved, a slender elongate rod which is undoubtedly the ilium, since it has the usual position for that element and is much too large for a sacral rib. It has much the same shape as the ilium in the modern Salamandra, and is not expanded as is the ilium of Branchiosaurus. This element, like the humerus, seems to have been but a hollow cylinder of bone and undoubtedly had cartilaginous ends, as in the ilium of the recent Salamandra. The two femora are preserved nearly entire, the right one lying upon and partly obscuring the sacral vertebra. The femur is much more slender than is the humerus, with slightly expanded ends, and, like the humerus, shows the concavities at the ends, indicative of the perichondral character of the tissue composing it. There are two elements of the leg preserved more or less entire. The larger bone may repre- sent the tibia and the smaller the fibula. They both present characters similar to those of the femur and humerus. They are simple rods of bone tapering at the distal end. The feet have been lost, though doubtless they were present at one time. The ventral surface of the body, as in other members of the Branchiosauria, was covered and protected by a series of small scutes arranged in the regular chevron pattern. The form of the scutes and their number can not be determined. The lines which represent them are, however, distinct. Some of the scutes are missing and some of them are obscured by lying over the vertebral column. They are all somewhat shifted to the left. The lines are very small and close together. I count 16 of them in a distance of 2 mm. In length the longest line preserved is a little more than 4 mm., measuring from the point of the chevron. The lines representing the scutes come to a point in a median ridge which is now represented by a line. The dermal scutes on the abdomen were probably the forerunners of the abdominal ribs of the reptiles (fig. 9). The impression of the tail contains some of the most interesting features in the entire specimen. Scattered over it and in places laid in mosaic are impressions of 56 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. small dermal scales, which may have covered the entire body. In form the scales are ovoid, being half as wide as long, and the markings on the scales partake of the nature of radiating lines, much after the pattern of the sculpturing of the cranial bones in the Microsauria. The scales are less than 0.5 mm. in diameter and their character can only be ascertained under high magnification. Near the middle of the tail there are preserved distinct transverse bands of dark color, which are more or less evident throughout the entire tail impression, but they are elsewhere not so distinct as in the central region. The lines are evidently due to rows of pigmented scales, and in all probability the animal's body was transversely striped. The most interesting and important single structure discovered on the specimen is the impression of the lateral-line system, which is clearly evident as two dark lines on the impression of the fleshy part of the tail. The sense-organs are repre- sented by two longitudinal rows of pigmented scales, one beginning at the tip of the tail, the other taking its origin from the median line somewhat further forward. I am indebted to Dr. Katashi Takahashi for calling my attention to the similarity of this arrangement to that found in the recent Necturus. The arrangement and dispo- sition of the lines containing the sense-organs is practically the same in the two forms. The median lateral line takes its origin from the extreme tip of the tail and is continued to the base, where the impression is broken. The dorsal lateral line has its origin rather abruptly from the median lateral line at a distance of 6 mm. from the tip of the tail. The sense-organs were undoubtedly located beneath specialized pigmented scales on the surface, and to this pigment is due the preservation of the lines. The fact that the arrangement of the sense-organs of Micrerpeton corresponds so exactly to the condition found in Necturus is of considerable interest. Necturus alone among the modern tailed Amphibia has the arrangement described for the lateral-line system of Micrerpeton. All other forms of the Caudata, as also the larval forms of the Salientia, have an arrangement of the lateral-line system which is per- fectly distinct from that found in Necturus, although the basis of the same arrange- ment is found in all. In Ambly stoma, for instance, the median lateral line is not present on the tail, and the dorsal line is incompletely developed. The close simi- larity of the arrangement of the systems of sense-organs in the two forms, Micrer- peton and Necturus, may be of genetic significance with regard to the latter form. The lateral-line sense-organs are of a very fundamental significance, and it is not at all improbable that the same arrangement of the lines has existed from the Carbo- niferous period or earlier. We know that such has been the case in a great many of the fishes. The ancestors of the modern Caudata must have originated somewhere in the basal Carboniferous or earlier periods, and, in the writer's opinion, the Branch- iosauria represent the ancestral group of the Caudata. This suggestion is by no means new, since Baur and others have held the same view. This topic has been discussed at length elsewhere (459) by the writer. The relations of the form Micrerpeton caitdatum are readily determined. The number of the presacral vertebrae, the form and position of the ribs, the shape of the skull, the arrangement of the cranial elements, the structure of the pectoral girdle THE AMERICAN COAL MEASURES BRANCHIOSAURID/E. 57 and the character of the ventral armature all clearly bespeak a close relationship with Branchiosaurus, Melanerpeton, Pelosaurus, and other European branchiosau- rian forms from the Upper Carboniferous and Lower Permian. The above-described species, with others given below, is the earliest geological evidence of the Branchiosauria, since the oldest European forms are from the Ste- phanian (Upper Carboniferous), which probably lies somewhat above the horizon of the Allegheny series of North America. The presence of the Branchiosauria in America is of considerable interest in the bearing it has on the distribution and migration of the Paleozoic animals. Knowledge of how the group came to occur in such widely separated localities in approximately contemporary geological strata is an unsolved problem of paleontology. It is possible that the piscian ancestors of the Amphibia migrated across or along the borders of the seas and began the amphibian phase of development independently in the two continents. That evolution should, in this case, have followed almost exactly parallel lines seems incredible. Measurements. ram. Length of entire animal 49 Length of rib 1.5 length of head in median line 6.5 Length of scapula 3 Width of head at posterior border 8- Maximum width of clavicle 2 Length of orbit 2.5 Length of humerus 2.5 Width of orbit 2 Length of ilium 1.5 Interorbital space 2 Length of femur 2 Length of the vertebral column 33 Length of tibia .... 1.5 Length of the vertebral centrum in dorsal series. . 0.5 Length of tail impression 21.5 Length of trunk from base of skull to sacrum. ... 22 Width of tail impression at base 4 Genus EUMICRERPETON Moodie. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 330, 1912. Type: Eumicrerpeton parvum Moodie. The genus is established on three well-preserved specimens representing nearly the entire anatomy. The generic characters are found in the very broad posterior table of the skull, with its short length, reduction of tympanic notch, and shortness of body. The body-length of Eumicrerpeton (plate 5, fig. 1) is less proportion- ately than that of other closely allied genera. Other generic characters are found in the sharp postero-lateral angle of the skull, and it is to be distinguished from Micrer- peton, especially, by the short, stumpy limb bones. The narrow, elongate eye, placed close to the edge of the skull, is a character not observed hitherto in the Branchiosauria. The genus is closely allied to Branchiosaurus of Europe. Eumicrerpeton parvum Moodie. Moodie, Proc. U. S. Nat. Mus., 40, p. 430, fig. 1, 1911. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, pp. 331-336, p'- 3. figs- 3 and 4; pi. 4; pi. 5, fig. 1; pi. 6, figs. 1 and 2, 1912. Moodie, Amer. Nat., 44, pp. 367-375, figs. 1-4, 1910. Moodie, Science, n. s., xxxi, No. 789, p. 233. Type: Specimen No. 803, Yale University Museum. Other specimens, No. 802, Yale University Museum, and No. 4400, U. S. National Museum. Horizon and locality: Mazon Creek shales, near Morris, Illinois. The impression of the outline of the entire body is preserved (plate 3, figs. 1 and 2) in three specimens, and in all are found molds and impressions of the alimentary 58 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. canal, which, in one specimen (471), are remarkably complete and instructive. The three specimens will be discussed separately, since they show different features. The impression of the larger animal (No. 803, Yale University Museum), which is probably an adult, presents the following elements: the entire skull, both humeri, impressions of posterior and anterior ventral armature, portions of the alimentary canal, one femur, portions of a fibula and tibia, and the entire impression of the tail, on which, as in Micrerpeton caudatum, there occur two definite dark lines, one begin- ning at the tip of the tail and running obliquely along the tail to where the impres- sion is broken at the anal region; the other beginning at a distance of 4.5 mm. from the tip and running almost parallel with the median line. These two lines undoubt- edly represent the lateral -line system. The skull is especially noted for its shortness and the great posterior width, as well as for the almost entire absence of the tympanic notch. The pineal foramen is located on a line with the posterior border of the orbits. The eyes themselves are narrow and acuminate at each end, with a pronounced convexity inwards and a flattening on the outer margin. They are located on the very border of the skull, but relatively more posterior than in Micrerpeton. No sclerotic plates are evident. The median suture can be indistinctly observed running the entire length of the skull. The sutures bounding the outside of the frontals and the squamosals are partially evident, but not satisfactorily preserved. The mandible is represented by a mold which in wax impression shows short, stumpy teeth. Posterior to the skull at a distance of a millimeter there are two sharp impres- sions which may represent the anterior edges of the interclavicle or they may be branchial elements. They are distinctly curved, however, and probably represent portions of the interclavicle. A wax impression does not show a discrete structure, but the boundaries of some larger element. No other remains of the pectoral girdle can be discerned. The humeri are short and relatively thick. Wax impressions show them to have had truncate or slightly concave ends, thus indicating the absence or slight development of the endochondrium. No other elements of the arm are preserved. The impression of an elongate femur and the heads of the tibia and fibula of the left side are preserved. The ventral armature is preserved in two small patches, and these show the chevron-shaped rods to have been very fine — much more delicate than in Micrerpeton. The body impression is very instructive and interesting, both in showing the form of the body and because in it are preserved the larger portions of the alimen- tary canal. The form of the body can best be discerned by reference to the figures (plate 3, figs. 1 and 2; plate 5, fig. 1). The portions of the alimentary canal preserved consist of the greater portion of the stomach, three coils or loops of the small intestine, the rectum, and a pit which undoubtedly represents the anal opening. The anus is found at a distance of 1 6 mm. from the tip of the tail and is somewhat removed from the body portion, as in mod- ern salamanders. On each side of the posterior end of the rectum there occur a pair of enlargements which probably represent the oviducts at their posterior ends (fig. 15, C) . MOODIE PLATE I 1. The larger specimen of Eumicrerpeton parvum Moodie. XI. 2. The smaller specimen of Eumicrerpeton parvum Moodie. X ]. 3. Type specimen of Erpetobrachium mazonensis Moodie. X 1. 4. Type specimen of Erierpeton branchialis Moodie. X 1. 5 and 6. Type specimen of Mazonerpelon longicauJatum Moodie. X 1. 7. A copy of Cope's drawing of the type specimen of Amphibamus grandiceps Cope. The original was destroyed by fire. X 3. THE AMERICAN COAL MEASURES BRANCHIOSAURID^E. 59 €^ ic * The tail impression is more acuminate than in Micrerpeton, but shows the same structures as in that form, i.e., the lateral lines which have already been mentioned. Micrerpeton was a more rapid swimmer than the present form on account of the greater development of the tail. The second specimen of the species (No. 802, Yale Museum) shows much the same character as the specimen already described, except that there are impressions of small, blunt teeth on the mandible. The two humeri and the femur of the left side are preserved and the interclavicle is represented by an identical impression, as in the first-described specimen. The tail impres- sion, though similar in form, does not exhibit so much of the structure of the lateral lines (fig. 15, B). The matter of especial interest in connection with this second specimen is the remarkably perfect preservation of the alimentary canal, which is entire, except for the very anterior end of the oesophagus. The posterior portion of the oesophagus, which measures 3.5 mm. , is clearly preserved. Its anterior end is thrown around posteriorly and indicates that this end was loosened after death and became dis- placed before fossilization. The length preserved may represent the entire oesophagus. The oesoph- agus is constricted before it enters the stomach, which shows the usual curvature found in modern salamanders. The stomach measures 6 mm. in length by 2 mm. in breadth, and consists of a single enlargement as in the modern Ambly stoma punctatum. It increases in size somewhat toward the pyloric end and then very gradually constricts to the pylorus. Three divisions of the small intes- tine can be seen. The most anterior one, corre- a. ThMmedmenatEimkrcrtttonpamm sponding to the duodenum, is segmented, as though S^hi^f xY^OrigSi the intestine had been filled with food before inter- in United states National Museum. a, anus; /, femur; ft, humerus; ic, lntrr- ment. The remainder of the intestine, correspond- clavicle ;"»«, intestine; m, mandible; or, ,. ., . . , . , . . , „ orbit; st, stomach; (, tibia and fibula. ing to the ileum, is looped in the iorm 01 two figures b. Type specimen of Amphibamus tiwra- 8 which are superimposed, with the upper portions £K». SJTffSL^S of the 8 at right angles to each other. The rec- see p- *32-) turn is clearly discernible, though its lower portion is somewhat obscured by having the lower part of the upper loop of the intestine lying over it. The anus lies at a distance of 1 .5 mm. posterior to the transverse line from the upper end of the femur, and is quite well back on the tail, as in modern salamanders. In this specimen also occur two oval bodies which may be identified as the lower ends of the oviducts, thus indicating, in all probability, that the animal was a female. A dissection of several species of modern caudates has resulted in the discovery that the adult condition of the alimentary canal of all the species dissected (.1 mhly- B 60 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. stoma pimctatum, Necturus maculosus, Diemyctylus torosus, D. viridescens, etc.) is much more complex than that exhibited by the specimen under discussion. A very near approach to the condition found in Eumicrerpeton parvum is found in an imma- ture branchiate individual of Diemyctylus torosus, 56 mm. in length, from a fresh- water pond on Mount Constitution, on Orcas Island, Puget Sound, Washington. The similarity of the intestinal structure is of considerable importance to our" understanding of the relationship existing between the Carboniferous Branchio- sauria and the modern Caudata, and only confirms other arguments, offered in another place (459), concerning their immediate relationship. The branchiosaurian affinities of the present species are almost too evident to need discussion. The entire structure is essentially similar to that of other genera of the order. The third specimen of this species (No. 4400 of the U. S. National Museum) is almost as perfectly preserved as were the other two specimens. The skull structure, the intermediate position of the pineal foramen, the epiotic notch, and the shape of the skull are essentially similar to the described specimens of the species. The present specimen is more developed than the other two and probably represents an adult. The alimentary canal is perfectly preserved. It is nearly half again as long as the smallest of the above-described specimens, and the skull is proportionately longer and wider. There is preserved also an impression of the anterior ends of both clavicles. The right humerus is imperfectly preserved, as is also the right femur and tibia; other than these the fossil is merely an impression. The skull is so similar to those described above that additional description is unnecessary. The pineal foramen is quite large and lies on a line which cuts the orbits into equal longitudinal parts. The interorbital space is about equal to the long diameter of the orbit. Traces of sclerotic plates are observed in the left orbit, but they are quite imperfect. The alimentary canal (fig. 14a) is unlike the previously described structures, in that the intestine is longer and more convoluted. It lies in five longitudinal folds and ends in an enlarged cloaca, near which there are impressions of two glands, or the pos- terior ends of the oviducts, as was suggested for the Yale specimens. The creatures undoubtedly fed on small plants and animals much as do the recent salamanders. Measurements of Eumicrerpeton parvum Moodie. No. 803 (222), Yale University Museum: mm. No. 4400, U. S. National Museum: mm. Length of animal 37.5 Length of entire animal 45 Length of skull 4.5 Length of skull 6 Posterior width of skull at table 6 Width of skull 9 Long diameter of eye 1.75 Transverse diameter of orbit 1 .50 Transverse diameter of eye 65 Long diameter of orbit 2.25 Length of left humerus 1.50 Interorbital space 2.50 Number of ventral armature rods in 1 mm .... 10 Diameter of pineal foramen 50 Length of femur 1.75 Length of body from back of skull to pelvis ... 22 Width across base of tail impression 3.5 Oreatest width of body 9 Length of tail from base to tip 17 Length of tail 16 No. 802 (471) Yale University Museum: Width of tail at base 5 Length of animal 30 Png&°r humerus 3 Length of skull ^4 'M8* °f *"»"*£ V v: 2'5° Posterior width of skull 5 Length of tibia (fibula?) 1.75 Length of oesophagus 3.5 Length of stomach 7 Length of stomach . 6 Width of stomach 3 Width of stomach 1 .33 Estimated length of intestine 18 Width across base of tail impression 2.5 Length of tail from base to tip 7 Length of intestine (estimated) 5.6 Width of intestine 1 1 and 2. Vertebrae of Spoil dylerpeton spinalum Moodie. X 1. 3. Tvpe specimen of Mazonerpelon rostatum Moodie. X 1. 4. Type skeleton of Cephalerpeton venliiarmnliim Moodie. X 0.9. 5 and 6. The halves of the nodule containing a practically complete skeleton of Ampliibamus gramtkeps Cope. X 1. Originals of Above figures in the Yale University Museum. THE AMERICAN COAL MEASURES BRANCHIOSAURIOE. 6l Genus MAZONERPETON Moodie. Moodie, Kans. Univ. Sci. Bull., vol. vi, No. 2, p. 336, 1912. Type: M. longicaudat&m Moodie. The genus is distinguished from other known branchiosaurian genera by the great length of the dorsal region, the elongate tail (plate 5, fig. 2), with its well- developed caudal ribs, the reduction of the tympanic notch, the broad nature of the scapula, the elongate interclavicle, and the slender ilium. The number of dorsal vertebras is identical with that of Branchiosanrus of Saxony. Mazonerpeton longicaudatum Moodie. Kans. Univ. Sci. Bull., vi, No. 2, p. 337, pi. 3, figs. i-2;pl. 7,fig.3;pl. 10. 1912.' Type: Specimen No. 795 (1234), Yale University Museum. Horizon and locality: Mazon Creek shales, near Morris, Illinois. (Plate 3, figs. 5 and 6.) The remains consist of the following elements: an incomplete skull; nearly the entire vertebral column, consisting of cervical, dorsal, sacral, and caudal vertebrae, 36 in number; several ribs preserved on each side of the vertebral column; a portion of the ventral armature; the scapulae; a clavicle; the interclavicle; both humeri; the radius and ulna of one side and the ulna of the other; portions of both hands; the ilium of the right side; both femora, and a partial impression of the left tibia. The skull is, unfortunately, very poorly preserved. Enough remains, however, to determine the essential characters. The skull bones, unlike any other American branchiosaurian, have an ornamentation consisting of sharp pits and elevations which in places have a quincuncial arrangement and in others take the form of defi- nite lines of pits or tubercles similar to the condition found in many of the Micro- sauria. The orbits are large and are situated back of the median transverse line of the skull. They are almost circular in form and contain 6 elongated sclerotic plates very closely arranged around the borders of the right orbit. The. plates are twice as long as wide. The interorbital width is 1 .25 times the transverse diameter of the orbit. Not many of the sutures of the skull are discernible. Portions of the frontals, the nasals, the prefrontals, the parietals, and the supratemporals can be identified. Their arrangement is shown in figure 14a. There is a decided posterior table to the skull, with truncate posterior border. The tympanic notch is shallow, with its outer border not so well protected as in Branchiosaurus. The cervical vertebrae arc incomplete, but their number was 4 or 5, as in M i< - rerpetott. The structure of the dorsal vertebrae is also uncertain, although the shape can be discerned. The vertebrae are short and thick, very unlike the long, cylin- drical vertebrae of Cephalerpeton. The heavy transverse process is quite evident on the best preserved vertebrae. This process recalls that described by Credner for the Saxony Branchiosauria. Several of the vertebrae show the articulation of the ribs with this process. The ribs of the caudal region recall very strongly those of Branch- iosaurus. They are quite heavy in the anterior caudal region and then diminish rather rapidly to the point where the tail is broken and lost. 62 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. They are quite different from The ventral armature is represented by a patch of chevron rods 21 mm. in length. The rods take a very peculiar form, being short crescentic bundles of fine rods, hair-like in appearance. In one of these bundles I count 5 smaller rods. The bundles are arranged in rows similar to the pattern so characteristic of the Car- boniferous Amphibia, as described elsewhere. The patch of ventral armature pre- served belongs to the abdominal region. A single row of the crescentic bundles measures 11 mm. Both scapulas are preserved in their entire form those of any other genus, being broadly crescentic with a posterior concavity and an anterior protuberance. The anterior surface of both scapulae is obscured. Vas- cular foramina occur near the base of both scapulas; there being three of them in the right scapula, arranged in the form of an isosceles triangle . The morphology of these foramina is uncertain. They have never been observed among the Carboniferous Amphibia, and, so far as I am aware, they are entirely unknown among higher vertebrates. The temnospondylous Amphibia of the Carboniferous and Permian possess, in the co-ossified scapula-coracoid, three fora- mina, very similar to the ones in the pres- ent form, but they are confined to the coracoidal region and, in the Branchio- sauria, as identified by Credner, the cora- coid is a free element, although I have never been sure of its identity among American forms. Williston has called these foramina the glenoid, the supraglenoid, and the supracoracoid foramina (Journal Geol- ogy, xvii, No. 7). They are not, however, to be correlated with the three foramina above mentioned, since in the Temnospon- dylia the foramina belong with the coracoid and not with the scapula. The condition of the Temnospondylia occurs in the bony fish Xiphactinus andax Leidy, and an analogous condition obtains in the reptiles, as in the Mosasaurs and Dinosaurs. Near the outer edge of the right scapula there is a large fragment preserved, which, I think, must be the misplaced clavicle. It is obscurely triangular or, more exactly, spatulate. The interclavicle is represented by fragments only, and seems to have had a narrow form. I I W f \<^ Si Fig. 14a. — Skeleton of Mazonerpeton longicaudatum Moodie. c, carpus; d, clavicle; cr, caudal ribs; cv, caudal vertebrae; h, humerus; /, femur; or, orbit; r, radius; sf>, sclerotic plates; sc, scapula; u, ulna; us, ventral scutelke. From Mazon Creek. Original in Yale University Museum. THE AMERICAN COAL MEASURES BRANCHIOSAURID/E. 63 The humeri recall those of Micrerpeton. They are somewhat elongate and appar- ently cylindrical in their normal condition, although somewhat flattened in the fos- sil. The shaft is considerably constricted at the middle and the ends are expanded, in which expansion the lower end exceeds. The ends are abruptly truncate, indi- cating a small amount of endochondral ossification or its entire absence. The mesopodial elements, unlike what is described for Cephalerpeton, are quite dissimilar in form, recalling the condition in Mesosanrus brasiliensis McGregor. The larger element is, apparently, the ulna. It has the lower end greatly expanded and the shaft is curved outward, resembling very much a reptilian ulna. The radius is much smaller than the ulna, lacks the lower expansion, and is shorter by i mm. The carpus is represented merely by a blank space, with evidences of impres- sions of cartilage in the sandstone. The hand of the right side contains 4 phalanges. There are 2 phalanges preserved in the first digit, including a sharp-pointed ter- minal phalanx, and the second digit has only the metacarpal. The third has the metacarpal and the first phalanx, which does not differ in form, but only in size, from the metacarpal. The fourth digit contains only the metacarpal. Of the left hand there are portions of 3 digits preserved, including 3 metacarpals and a phalanx, which in structure are not different from those of the right hand. The ilium of the left side is preserved, apparently entire. It is elongate and cylin- drical, its upper end adjoining the twenty-eighth vertebra. The head of the femur lies close to the lower end of the ilium, so that that element must have been suspended in the flesh, much as in modern salamanders. It could not have been of much use as a support for the body. The form of the femur is not unlike that described for the humerus, save that its lower end is smaller than the upper, while in the humerus the extremities are of equal diameter. A portion of the right femur is preserved, extending in an opposite direction to the left. Measurements of the Type. Length of entire specimen 64 Length of portion of skull preserved 6.5 Posterior width of same 7 Width across orbits 11 Long diameter of orbit 3 Transverse diameter of orbit 1.75 Interorbital width 4.75 Length of dorsal Vertebrae 48 Length of caudal scries 11 Length of anterior dorsal centrum 2 Length of anterior dorsal rib 4 Length of anterior caudal rib 1 .75 Length of scapula 5 Greatest width of scapula 4.25 Probable length of interclavicle 6 Width of same 3 Mazonerpeton costatum Moodie. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 341, pi. 2, fig. 3; pi. 8, fig. 4; pi. 9, fig. 2; pi. 10. 1912. The remains on which the present species is based are inclosed in a much frac- tured nodule. The parts of the animal which have been identified are: a part of the skull and left mandible, two clavicles, a humerus, impressions of several verte- brae, a portion of the dorsal region of the body with several ribs, two portions of the caudal region with several ribs and unidentified fragments. (Plate 4, fig. 3.) Length of clavicle 4.5 Width of same 1.5 Length of right humerus 6 Distal width of same 2 Length of ulna 3.25 Distal width of same 1 Length of radius 3 Width of carpal space 2 Length of metacarpal 1 .74 Length of first phalanx 1 .75 Length of distal phalanx of right hand 35 Number of chevron rods in I mm 4 Length of ilium 2.25 Length of femur 4 Proximal width of femur 1 .50 64 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. rb$Lcv The animal, from the shape and form of the ribs, is undoubtedly a branchio- saurian, since short, heavy, straight ribs have not yet been found to be associated with other than branchiosaurian structures. It is placed in the genus Mazonerpelon on account of the structure of the pectoral elements, the form of the humerus, and the length of the tail, all of which agree in structure with Mazonerpelon longicauda- tum. The animal attained, perhaps, a length of 4.5 inches, while that of M. longicau- datum was about 3 inches. The tail in the present species is very long and slender, more elongate than in any other known branchiosaurian. The part of the skull preserved is very unsatisfactory and, aside from the fact that it seems to represent the under side of the left half of the skull, little can be said. Three sutures can be observed, but what sutures they are is undetermined. The left mandible lies crushed on the edge of the skull and partially obscures what little there is of that structure. The slightly curved impression, from which the bone has been either broken or weathered, measures 13 mm. in length by 3 mm. in posterior diameter by 1 mm. in anterior diam- eter. These measurements show the element to have been slender and pointed anteriorly. Very little accurate information can be derived from the study of the vertebral column of the speci- men, nor can the dorsal vertebral formula be made out, since only a portion of the length of that region is preserved and only a few rather indefinite impressions can be dis- cerned. These impressions show Fig. 14J.— Skeleton of Mazonerpelon coslatum Moodie. X 1.5. ,, ... - . j 1 • 1 Original in Yale University Museum, dv, dorsal vertebra;; the Vertebrae tO be Short and higher ch, neutral spines; d, clavicle; cv, caudal vertebra:; /, femur: than in most Branchiosauria. *- humerus; m' mandible; '*• rib= **• « ». verteb™. The caudal series is represented by two sections, one of which is, apparently, from near the base of the tail, judging from the size of the caudal ribs preserved ; the other is from near the tip of the tail, and shows the constituents to have been long and slender. Ribs are apparently absent on this section. The position of the two caudal sections shows that when the animal died it was coiled up much like a snake, so that in the fractured nodule three sections of the body are visible. The tail was probably half as long again as the body. The ribs throughout the body are short, heavy, and straight, with, in the dorsal series, a lateral and a distal expansion which is taken as a distinctive specific char- acter. Judging from imperfect impressions in the dorsal series, the ribs were attached to a transverse process of the centrum, thus agreeing with other branchio- MOODIE PLATE 5 2. A reconstruction of the possible appearance in life of the Coal Measures branchiosaurian, Eumkrerpetoii parvum Moodie, a small, primitive salamander, less than 2 inches in length, based on three specimens from the Mazon Creek Shales. The lateral-line organs are represented as dark bands on the tail, the sense organs being, apparently, situated beneath specialized pigmented scales, to which is due the preservation of the lines. A restoration, natural size, of the branchiosaurian, Mazonerpeton , based on two specimens. The form of the animal is quite salamander-like. It is shown when about to feed on a specimen of Acanthotehon stimpsoni, which is said to be a brackish-water crustacean. The branchiosaur and crustacean may possibly have inhabited the same body of water. THE AMERICAN COAL MEASURES BRANCHIOSAURID.^. 65 saurians in this respect. The ribs show a progressive decrease in length from the cervical region to the point of their disappearance on the tail. The pectoral girdle is represented by two elements, one of which is certainly the right clavicle, and the other is possibly the left clavicle, though its form is some- what distorted by pressure. Both elements are in the form of an elongate spatula with the dorsal surface greatly concave and the inner end acuminate. The right humerus is imperfectly preserved, though the impression allows one to gain an exact idea of its form. It lies under the right clavicle. Its ends are truncate with a contracted shaft and expanded extremities; the bone was apparently hollow. in Fig. 15. A. Impression of Erierpeton branchialis Moodie. bb, basibranchial ; hyp, hypohyals; m, mandible; x/3, The reference by Cope of this genus to the Caudata is one of the most interesting facts connected with the Paleozoic Amphibia. He says: "The present genus is, then, to be referred to the neighborhood of Amphiuma and Protonopsis, but forming the type of another family" (123). He regards the branchial apparatus as being more fish-like than that of any of the modern genera. It is possible that Cocytinus gyrinoides was a larval branchiate and consequently aquatic form. It should be more fully compared with Erierpeton branchialis from the Mazon Creek shales when better known, as well as with Ilyphasma Icevis from the Linton locality. All three of these forms are included, provisionally, under the Cocytinidae. Cocytinus gyrinoides Cope. Cope, Proc. Am. Phil. Soc, xn, p. 177, 1871. Cope, Trans. Am. Phil. Soc, p. 278, 1874. Cope, Geol. Surv. Ohio, 11, pt. 11, pp. 364-365, pi. xxxix, fig. 4, 1875. Type: Specimen No. 8613 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. Two specimens of this interesting amphibian are known, one of them fairly com- plete (No. 2564, Am. Mus. Nat. Hist.). The type specimen consists of the inferior bones of the cranium in a fairly complete state of preservation, with the muzzle and its teeth; also 8 anterior vertebrae, with their short recurved ribs. The condition of the hyal elements in the type specimen is as follows: the haemal elements of the first branchial arch are partially concealed on both sides of the ceratohyal. An expanded truncate face for attachment to the axial element is visible on both sides, but the body of the bone is flat and presents the edge of the specimen. The first pleural element proceeds from just behind the axial- hyal; it is longer than the other pleural elements. A slender bone is visible extending from the space between the cerato- hyal and mandibular angle; it may, therefore, pertain to the sus- pensorium of the jaw as well as to that of the hyiod arch, or be squamosal as well as stylohyal. The second haemal bone is slender, but with an enlarged axial extremity; that of the right side is not so well preserved as to be safely determined. The third haemal elements are the smallest, and originate imme- diately in front of the occipital condyles and diverge outwards and backwards. They are little curved, subcylindric, and slightly expanded at the extremities. Of the pleural elements the first and second are little curved and the first is marked by a pit or foramen on the under side near the distal end, which is clearly visible on both sides of the specimen. The third and fourth pleurals are more curved and the outer ends slightly expanded and directed backwards. The obverse of the specimen (fig. 16) shows that the anterior axialhyal is wedge- shaped. The lateral basihyals are massive. The second hasmal branchihyal is FlG. 16. — Obverse of Cocy- tinus gyrinoides Cope, from the Coal Meas- ures of Ohio X 2. pmx, premaxillas; mx, maxilla; m, mandible; ah, axialhyal; h, basal branchihyal ; ch, cer- atohyal; bv, haemal branchihyal; b, bll, bill, bllll, pleural branchihyals. PI ATE 6 .2 *•* a H II 11 -* at 5 .if "C 80 .2 £ 01 u _ 3 a J= I< >.a » . v ■ u 2 3 »i 1> 3 -a ■ *-» * .a £ v 3 3 J= O - u / £ J) «-> .5 •« 5-3 I 9 s I B 5 a .<= if I M 5 O £ S x C £L 2 si! S 8 2 i> y g 5 -. «• w > z ^ z s Z ° S - H i s g ial ^ o * 5 •£ * i ORDER CAUDATA DUMERIL, l8o6. 69 dilated, fan-shaped distally, and supports two pleural elements. The muzzle pro- jects over the lower jaw and was rather broadly truncate. The premaxillary teeth are cylindric and 6 in number on each side. The maxillary bone is represented by a lamina at each lateral extremity of the premaxillary. The mandibular rami are very stout, as are also the ceratohyals. The vertebrae have possessed some apophy- ses, apparently keel-shaped diapophyses. The ribs are slightly curved. Measurements of the Type. mm. mm Length of specimen 32 Length of axialhyal 3 Length of skull 12 Length of postbranchial 4 Posterior width of skull 11 Width of vertebra 1 Length of premaxilla 5 Length of vertebra 3 Length of mandible 10 Length of rib 6 The other specimen of this species (fig. i6«) is interesting in having 40 consecu- tive vertebrae preserved, and 19 pairs of ribs attached in their natural relations to the skull and hyal elements. There are a few hyal elements preserved, but nothing is added to our previous knowledge. The ribs are quite as in the type specimen, as Fig. 16a. — Nearly complete specimen of Cocylinus gyrinoides Cope, from the Coal Measures of Linton, Ohio. Original in the American Museum of Nat- ural History. X 0.95. are also the vertebras. The animal was apparently a slender, eel-shaped amphibian comparing favorably with the modern Amphiuma in this respect. There are no indications of limbs or limb girdles. Measurements (No. 2564, American Museum of Natural History). mm. Length of entire specimen 113 Length of skull 15 Width of head posterior 15.5 Length of vertebra 2 Length of rib 4 Genus ERIERPETON Moodie. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 328, 1912. Type: Erierpeton branchialis Moodie. The generic characters are found, first of all, in the presence of hyobranchial arches which indicate its relationship to the formerly described Cocytinus gyrinoides Cope, from Ohio. The only other known extinct genera of Caudata which possess, or at least have preserved, the hyobranchial arches are the Jurassic Hylceobatrachus from Belgium and Lysorophus from the Permian of Texas. The present form is widely distinct from both of these genera in the shape of the mandible and the form and arrangement of the hyobranchial bars. The genus Erierpeton finds its closest ally in Cocytinus, in the family Cocytinidae, which possibly belongs in the order Caudata and the suborder Proteida of Cope. 70 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Erierpeton branchialis Moodie. Moodie, Kans. Univ. Sci. Bull., VI, No. 2, pp. 329-330, pi. I, fig. 3; pi. 2, fig. 1, 1912. Type: Specimen No. 801 (222) 5, Yale University, Museum. Horizon and locality: Mazon Creek shales, near Morris, Illinois. The amphibian remains designated by the above name consist of a distinct man- dible and some rather indefinite body impressions (plate 3, fig. 4). Three elongate impressions occur between the rami of the mandibles (fig. 15, A), which, I suppose, must represent hyoid bones belonging to the hyobranchial arches. The lateral ele- ments are paired and the median impression is straight and lies between the paired impressions of the hyoids. The paired portions probably represent the hypohyals or hypohyals plus the ceratohyals, and the unpaired portion of the first basibranchial, according to the nomenclature of Wiedersheim (Comparative Anatomy of Verte- brates, 1897, p. 86). If the impressions have been correctly interpreted the present specimen is of very great interest, since it is the first evidence we have of the hyo- branchial arches in the Amphibia of Mazon Creek, and the second in the Carbon- iferous of North America. Dawson doubtfully identified (216) some elements of the Joggins Amphibia as hyoids, but was uncertain as to their position. Cope described fully the well-developed hyobranchial apparatus of Cocytinus gyrinoides (123) from the Coal Measures of Ohio. Among other Paleozoic Amphibia Williston (614) has described branchial arches in the peculiar form Lysorophus tricarinalus Cope, from the Permian of Texas. The form of the impression of the mandible in the present specimen is unlike anything known to the writer among other Carboniferous or later Amphibia. The rami are long, slender, deep, slightly curved, and pointed anteriorly. The anterior symphysis was not a complete sutural union, but was occupied partly by cartilage or other connective tissue. There are no definite traces of the appendicular skeleton. The traces of the body (fig. 15, A) indicate an elongated, rather slender animal, but further than this noth- ing can be said in regard to its structure. The occurrence of a typically caudate form in the Carboniferous is unusual and complicates still further our understanding of the origin and relationships of the early Amphibia. Measurements of the Type. mm> mm. Length of entire impression 50 Width of basibranchial 0.75 Length of mandible along median line 10 Length of hypohyal 2.4 Width of mandibular ramus 9 Width of hypohyal \ c Length of basibranchial 2.5 Genus HYPHASMA Cope, 1875. Cope, Proc. Acad. Sci. Phil., p. 16, 1875. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 387, 1875. Type : Hyphasma lends Cope. ' ' Vertebrae osseous, the posterior dorsals, and probably the caudals, furnished with fan- like neural spines; limbs unknown-(?) wanting. Thoracic shields present. Ventral arma- ture, consisting of rhomboidal scuta, forming packed rows arranged in chevrons, directed backwards, on top of which are the usual rod-like scales arranged in packed chevrons, with the angle directed forward. ORDER CAUDATA DUMERIL, l8o6. 7 1 ' ' The general appearance of the type of this genus is that of a Ptyonius, but the ventral armature is different from anything observed in the known genera of this group. The larger external scuta are like those of the species of Colosteus (Sauropleura) , but their series have a different direction. The inner chevrons are those of many other genera" (123). Hyphasma laevis Cope. Cope, Proc. Acad. Sci. Phil., p. 16, 1875, Cope, Geol. Surv. Ohio., 11, pt. 11, p. 387, pi. 37, fig. 4, 1875. Type: Specimen No. 9023 (in counterpart), American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. "In the only known specimen the vertebrae have low and squarely truncate neural spines near the head, and some distance anterior to the tail they are quite conspicuous and delicately line-grooved. The body is slender and probably limbless. The thoracic scuta are large and close to the head; the median is produced at both ends, but chiefly anteriorly while the lateral are narrow; all are without sculpture The head is seen from below. The mandibular rami are not so slender as in most species of Ptyonius, but are rather stout. They are a little incurved distally, so that the form of the muzzle is somewhat narrowed, but not produced. The teeth are not visible. Ten rows of the outer layer of scuta in 0.005 m-" (I23)- The specimen is very indistinctly preserved and the characters given by Cope can not all be made out. It is puzzling to see just on what he bases his conclusion. It is possible that the specimen is a poorly preserved Ptyonius. The outlines of the vertebrae are so indistinct that I am uncertain about them. In certain lights there appear to be regular impressions which resemble the spines of the vertebrae of Ptyonius, but they are doubtful. The skull appears totally distinct from any known species of Ptyonius, but it is very imperfect. The condition of the pectoral elements is very uncertain and I can not be sure that what Cope described as tho- racic "scuta" are such. The interclavicle, however, is clearly preserved as a diamond- shaped structure. It is almost smooth, with a few faint radiating lines near the base. It measures 5 mm. in greatest breadth by 8 mm. in length. MEASUREMENTS OF THE TYPE. mm. mm. Length of specimen as preserved 64 Length of median thoracic scuta 10 Length of skull 15 Width of same 4 Greatest width of skull 8 Width of clavicle 2 Width of body 8 Length of mandibular ramus 12 Length of 7 cervical vertebrae 15 CHAPTER XI. DEFINITION OF THE ORDER SAL1ENTIA, LAURENT), 1768. (World-wide distribution.) Naked, tailless Amphibia of compact form, and with usually proccelous vertebrae. Cau- dal vertebrae coalesced into a slender elongate piece, the urostyle. Two elements of the tar- sus ossified and greatly elongated. Development by metamorphosis; gills never present in adult. Ilium greatly elongated. The order is suggested in the Coal Measures by a single species, known from a single poorly preserved specimen (plate 24, fig. 1 ) . The form Pelion lyelli Wyman was the first known of the Linton Amphibia, and its striking frog-like (123, 639) appear- ance was early noticed. There is no assurance that the species belongs with this order, but since a well-developed and highly specialized frog (480, 481) occurs in the Como Beds (405) of Wyoming, it is not impossible that we may have in the Pelion lyelli a suggestion (460), at least, of the ancestral structure. It is certain that the frogs have, in past ages, had a much greater length of vertebral column than they possess at pres- ent, as is witnessed by the coalescence of several vertebras to form the urostyle. It is suggested that the ancestral vertebral column is represented in Pelion. Family PELIONTIDjE Cope, 1875. Cope, Bull. U. S. Nat. Mus., No. 1, p. 11, 187.5. The present family includes but a single species, that of Pelion lyelli Wyman, first described in 1858 (640), from Linton, Ohio. The family characters are to be found in the broad and obtusely rounded cra- nium, in the frog-like scapular arch, the frog-like hind limb, and in the form of the palate, so far as these structures have been preserved. It has been suggested that the present form shows decided affinities with the frogs of to-day and it may possibly be looked upon as the actual ancestor of the liv- ing frogs. The length of the vertebral column would seem to militate against such a relationship, since it is well known that frogs have had a short vertebral column since the Jurassic (480, 481). But this is not a good argument, since the developing urostyles of modern tadpoles show metameric fenestrations in the developing bone which doubtless correspond to openings between the vertebras. The notochord of the tail is segmented, apparently through the influence of former vertebral structure. At any rate, the suggestion is an interesting one and, whether sustained or disproven, the present discussion is based on the probabilities of the case. Genus PELION Wyman, 1868. Wyman, Am. Jour. Sci. (2), xxv, p. 160, 1858 (Raniceps). Cope, Proc. Acad. Nat. Sci. Phil., 1868, 211 (Pelion, suggested in letter to Cope by Wyman). Type: Pelion lyelli Wyman. ' ' The only specimen of the species exhibits an inferior view of a portion of the skeleton ; and the obverse, on which the thoracic and abdominal armor could have been preserved, has not come under my observation. The specimen, however, does not exhibit any ribs, although the vertebrae are well preserved. As observed by Professor Wyman, the genus pre- sents some points of similarity to the Anura (Salientia) . The prolongation of the angles of the mandible is of this character, as well as the general form of the head. The bones of the 72 MOODIE PLATE 7 Uvlerpeton dawtoni Owen. Above: Mandible, teeth, rib, and bones of anterior extremity. Below: Bones of pelvis and posterior limb and bony scales. Nearly natural size. Erect tree, Coal formation, South Joggins, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original specimens in Peter Redpath Museum of McGill University. DEFINITION OF THE ORDER SALIENTIA, LAURENTI, 1 768. 73 forearm may be united as in the frogs, and the length and curvature of the femur are seen among these animals rather than the Salamanders. The form of the femur is different from that of Amphibamus grandiceps Cope, which also differs in the presence of dermal scales and ventral scutellae." (123.) Pelion lyelli Wyman. Wyman, Am. Jour. Sci. (2), xxv, p. 160, 1858. Cope, Proc. Acad. Nat. Sci. Phil., 1868, p. 211. Cope, GeoL Surv. Ohio, 11, pt. 11, p. 390, pi. xxvi, fig. 1, 1875. Moodie, Pop. Sci. Monthly, lxxii, p. 562, fig. 1, 1908. Type: Specimen No. 7909 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. (Plate 24, fig. 1.) This was the first species described from the Linton, Ohio, deposits. It was made known by Dr. Wyman in 1857 at the meeting of the American Association for the Advancement of Science for that year. The species was subsequently studied by Cope. He merely confirmed Wyman's observations. The following description is based on the descriptions of Cope and Wyman and on my own study of the type specimen. This is the most frog-like, in appearance at least, of all the Amphibia which have so far been discovered in the Carboniferous. The skull especially has a shape which is strikingly frog-like, and the long hind limbs lend further likeness to the tailless forms. Pelion may have been a jumping creature, if we may judge from its long hind legs. Wyman and Cope have both called attention to the frog-like appear- ance of the specimen, and this is apparent at the first glance. It is probable that the resemblance has some significance as to the ancestry of the Salientia, and it may indicate the first step in the origin of the tailless Amphibia. It is possible that the frogs began to be separated from the other Amphibia during the Carboniferous. The first frogs we know are from the Jurassic, where they are well-developed ranids. If Pelion be a frog ancestor, then the history of the group from the Coal Measures to Jurassic is an unknown story. The specimen is preserved on its back and it is thus impossible to tell as to the structure of the skull. Cope was of the opinion that the depressed areas on the sides of the elongate parasphenoid were the orbits, and if so the resemblance to the frogs is much more striking. In the frogs there is a strong process from the ptery- goid which projects inward to meet a corresponding process from the parasphenoid. This forms a heavy rod behind the palatine vacuity. There is a heavy rod repre- sented in the specimen and a part of it is certainly the external process of the para- sphenoid, but whether it is to be interpreted as in the frog is an open question. The outline of the cranium is partially obscured by the mandibles, but the anterior part is represented by a raised line, as shown in figure 17. In the anterior part of this space there are two ridges which may be tooth ridges. If they are teeth there is a great similarity to the premaxillary and vomerine teeth of Necturus, since the ridges are widely separated at the median line and approximated distally, as they are in Necturus. The mandible is preserved entire and its form is strikingly frog- like. Its posterior angles project over the quadrate area and seem to have had an upturned projection such as is found in the mandibles of the Crocodilia. There are impressions of 20 vertebras preserved, and they cover a little more than half of the presacral region. There may have been 28 to 30 presacrals. The 74 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. vertebrae, as preserved, are somewhat quadrate in outline and constricted at the mid- dle, as though they were of the typical microsaurian type. No ribs are preserved. There is an impression anterior to the right humerus which may represent a part of the pectoral girdle, but its form is so obscure that it can not be determined. The pelvic girdle is entirely wanting in the specimen. Remains of the fore and hind limbs are preserved. The arms are especially well pre- served and consist of a strong humerus, a separate radius and ulna, and phalanges, the carpus having undoubtedly been cartilaginous, since there are no traces of carpal bones. Wyman has figured a small ossicle (fig. 17) which might be interpreted as a carpal, but it is further removed from the carpal region than his figure shows and I would interpret it as a fragment of a phalange, since the first digit seems to be turned aside over the vertebral column. The right hand is but imperfectly preserved, but the left hand is nearly entire. There are evidences of 4 digits, possibly 5. The metacarpals are elongate and rather stout. The phalanges of the distal series have been lost, so the phalangeal formula can not be determined. On the whole, the hand has a very broad aspect and is not at all slender, as in the majority of the microsaurians from the Coal Measures of Ohio. It resembles in a great measure the broad hand of a toad and may thus be indicative of a terrestrial life. The humerus is well developed and has pronounced swellings, as though for the attachment of strong muscles. These indications would favor the view of the animal being a land dweller. The femur and a part of the tibia (?) of the right side are all there is preserved of the hind limb. These elements show the leg to have been quite long, though weaker than the fore limb. The femur has a large distal articular surface. The fibula is, apparently, absent, though it may simply be lost. The genus Pelion stands alone among the Carboniferous Amphibia. The form can not be placed in the order Branchiosauria on account of the well-developed limb bones and the large mandible. It may belong with the Microsauria. I have placed it under the Salientia in the hope of learning more about the early relatives of the tailless forms. There is no assurance at all that it is even ancestral to the Salientia, but the resemblances are striking. The following gives the measurements of the type specimens : Measurements of the Type Specimen.* Fig. 17. Peliim lyclli Wyman, an amphibian from the Coal Measures of Ohio, the sup- posed ancestral salientian. (After Wyman.) X 0.75. pmx, premaxilla; pv, palatine vacuity; m, mandible; sc, scapula-coracoid ; /;, hume- rus; r-u, radius and ulna;/, femur; /, tibia. Length of specimen, as preserved no Median length of the skull 24 Width across the mandibular angles 25 Greatest width of skull 30 Length of vertebral column from occiput to sacral region 80 Length of left humerus 19 Width of distal end of left humerus 5.5 Length of radius and ulna 1 1.5 Width of distal end of radius 2 Length of digit II, as preserved 16 Ixngth of digit in, as preserved 14 Length of femur 24 Width of distal end of femur 4 Length of tibia 18 *The type specimen was collected in 1857 by Dr. John V. Lauderdale, who presented it to Dr. J. S. Newberry. CHAPTER XII. SUBCLASS LEPOSPONDYL1A Z1TTEL, 1887. COAL MEASURES TO PERMIAN. (Europe and North America.) The group is here defined according to the English edition of Zittel's Text Book of Paleontology, 1902, p. 125. "Notochord persistent and enclosed in constricted bony cylinders, hour-glass-shaped in longitudinal section. Teeth simple, conical, hollow. " According to Zittel there are two families, the Microsauridae and the Ais- topodidae. The latter family is dealt with under Aistopoda (p. 76) and it is there shown that the group is in no wise a valid one. The former family is regarded as an order and is fully entitled to that rank. As defined here the subclass Lepospondylia contains but a single order, the Microsauria. Extinct, terrestrial, aquatic, or semi-aquatic amphibians; skull pitted and grooved by lateral-line canals and by sculpturing marks, or the skull may be smooth; teeth present on most of the palate bones; exoccipitals cartilaginous or calcified, never completely osseous ; sclerotic plates sometimes present ; skull of various shapes. Vertebra? with notochord largely persistent, hour-glass-shaped; neural spines low or high, or absent; ribs intercentral and single-headed, with an incipient tubercle in some forms; vertebral column differentiated into dorsal and caudal series; cervical series not clearly defined. Limb bones with well-ossified perichondrium, endochon- drium partly ossified; epiphyses absent; carpus and tarsus (tarsus osseous in two species) cartilaginous; phalanges clawed or not; digits 4 in hand and 5 in foot. Pubis sometimes calcified but never osseous. Definition of the Order Microsauria Dawson, 1863. Coal Measures and Permian. (Europe and North America.) Lizard-like, sometimes longicaudate, stegocephalous, lepospondylous, ambu- latory or legless amphibians; skull bones usually sculptured with pits and grooves; lateral-line canals well developed on skull bones; skull with horns from tabulare and supratemporals or without horns ; branchiae never persistent ; sclerotic plates present; orbits usually well forward. Vertebra; hour-glass-shaped; endochondral bone weakly developed throughout skeleton, especially in vertebras; notochord largely persistent; neural spines low and rudimentary or long, fan-shaped, and highly ornamented. Dorsal series of vertebral column variable; usually from 22 to 30; tail containing sometimes over 75 vertebras, or tail very short with 2 weakly developed vertebrae; caudal ribs present, in those forms with long caudal series the distal vertebras sometimes exhibiting 2 pleurocentra. Ribs long and curved, always intercentral in position, single-headed, with at times an incipient tubercle. Pectoral girdle composed of scapulas, clavicles, coracoids, and interclavicle. Pelvic girdle composed of osseous rod-like ilium, plate-like ischium; pubis carti- laginous, sometimes calcified. Limbs present or wanting or weakly developed, sometimes present in front and wanting behind. Radius and ulna and tibia and fibula free ; carpus and tarsus usually cartilaginous ; digits 4 in hand and 5 in foot, terminal phalanges sometimes clawed. Phalangeal formula for the hand 2-2-3-2, for the foot 2-2-3-4-3. Abdomen covered with dermal armature composed of osseous or corneous rods or scutes ; overlapping scales, fish-like in appearance, some- times present over the entire body ; body also covered with lizard-like scales or naked. 75 76 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The order Microsauria was established by Sir William Dawson in 1863 (208) as a family of "reptiles" for the reception of the genera Hylonomus, Hylerpeton, Smi- lerpeton, and Fritschia. Hylonomus lyelli is the type species of the order. Dawson (216, p. 635) says of the species Hylonomus lyelli Dawson: "It is the type of the genus Hylonomus and of the family Microsauria." The forms which have been referred to the genus Hylonomus, and hence to the order Microsauria, from the deposits of Europe are discussed under Hylonomus. The Microsauria have been regarded by the writer and others as being ancestral in a sense to some of the later reptiles (469) , but there seem to be insuperable obsta- cles in the way of a direct derivation of the reptiles from the Microsauria. One of these obstacles seems to be found in the structure of the hand. In all Microsauria, so far as is known, there is no evidence of more than 4 digits in the hand, while no true primitive reptile possessed less than 5. The carpus of all true reptiles is osseous, while that of the Microsauria is merely cartilaginous. It is possible that the Micro- sauria stand in some such ancestral relation to the later reptiles as the Crossoptery- gia (489ft) do to the Amphibia. The Microsauria had undergone adaptive modifica- tions as to structure and habit, so that they have paralleled many of the groups of reptiles, but their structure is quite different. The evidence, as far as we can see now, points to a close genetic relation between the reptiles and the Microsauria, but that this relation is ancestral I, for one, am not ready to say. The Group Aistopoda Miall, 1873, is untenable. The group Aistopoda was established in 1873 as section ix by the Committee of the British Association for the Advancement of Science, in their "Tabular View of the Classification of the Labyrinthodonts." L. C. Miall (449, 450) was the secre- tary of the committee, and the report was published in two parts. Two genera were at that time attributed to the Aistopoda, Ophiderpeton and Dolichosoma, both described by Huxley from the Coal Measures of Kilkenny, Ireland. Fritsch (251, pp. 107-126) in 1883 refers to the group as "Familie" and describes 4 genera and 9 species as belonging to the group. Zittel (642, p. 383) refers to the group as "Familie" and places 5 genera in it. Smith- Woodward (Vertebrate Pale- ontology, 1898, p. 129) refers the Aistopoda to a suborder. In Eastman's trans- lation of Zittel's Paleontology the group is called a family, " Aistopodidas. " Lydek- ker (393, p. 205) regards the group as a suborder. The writer (469) refers the Aistopoda to an order. The group Aistopoda has been adopted by practically all paleontologists and zoologists who have had occasion to refer to these animals. Lydekker (393) in 1890 defined the group as follows: "Body long and snake-like, without limbs, and apparently without pectoral or pelvic girdles. Vertebrae with elongated centra and aborted neural spines. Ribs slender, and barbed like those of fishes. Teeth smooth, without plications of the dentine. External gills probably persistent." If now we take up a consideration of each of the characters mentioned by Lydek- ker we find that the first one holds good for all examples of the group. The second character, "without limbs, " is not good. Species of QLstocephalus , Ptyonius, Molgo- phis all possess limbs; and doubtless Ophiderpeton will be found to possess limbs PLATE 8 Fritschia (iirtidentata Dawson. Above: Bones of skull and anterior extremity, and bony rods of belly. Below: Bones of pelvis and posterior extremity. Nearly natural size. Erect tree, Coal formation, South Joggins, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original specimen in the Peter Redpath Museum of McGill University. SUBCLASS LEPOSPONDYLIA ZITTEL, 1 887. 77 also, since it has a well-developed pectoral girdle. The limbs in all of these genera are small. The third character, "apparently without pectoral and pelvic girdles," is not at all a good character, since nearly every specimen of some species and almost all species show evidences of pectoral girdles and a few exhibit pelvic girdles. The fourth character, "vertebrae with elongated centra and aborted neural spines," is not a good distinguishing character, since Amphibamns , an undoubted microsaurian, possesses the same vertebral characters. "Ribs slender, and barbed like those of fishes " is a character which is common to several widely distinct genera. All Micro- sauria possess long, slender ribs, and the barbed condition is one which is possessed by only a few, Thyrsidium, Ophiderpeton, etc., the so-called "barb" being merely a highly exaggerated tuberculum. The teeth of nearly all Microsauria are smooth, so that the character "teeth smooth" is not a good one for a group definition. It has not been possible to examine any of the American specimens for the plication of the dentine, since the forms are so rare and the fossils very fragile. The last character, "external gills probably persistent, " is certainly not true for the American species, and the evidence for the European species is negative. Fritsch described and figured (Fauna der Gaskohle, Bd. I, 1883, p. 114, Tafeln 18 and 23) structures which he regarded as supporting structures for the external branchiae. He says in regard to these structures: " Bei dem Umstande, dass sie von der Kiemengegend aus sich buschelformig verbreiten und man ihren Contakt mit einer Art von Branchiae constatiren kann, zweifle ich nicht daran, dass diese Stabchen dem Kiemenapparate angehoren. Bedenklich ist nur ihre grosse Zahl und das Vorkommen bis zum i6ten Wirbel und ich erwog die Moglichkeit, dass diese Stabchen einem zarten Bauchpanzer angehoren konnten. Da aber weiter im Verlaufe der ganzen Wirbelsaule nichts Aehnliches vorkommt, so ist man gezwungen anzunehmen, dass Dolichosoma sehr grosse lange Kiemenbuschel besessen haben muss." John Samuel Budgctt (79, p. 162), in his discussion of the "Structure of the Larval Polyp terus, " refers to the above-described specimen of "aistopodous Stegocephali, " i.e., Dolichosoma longissimum Fritsch, and calls especial attention to the similarity of the external rod of segmented cartilage on the hyomandibular of Polypterus to this structure, to which Fritsch has assigned a branchiate nature in Dolichosoma. There is no doubt in the mind of the present writer, however, that the rod of carti- lage, referred by Fritsch to the gills, can be other than scutellate rods of the ventral armature, these rods belonging to the armature of the breast or throat. The evidence for this conclusion is furnished by Fritsch himself ( Fauna der Gaskohle , Bd . 1 , plate 1 8 , fig. 11), where all may read in the figure of the specimen the facts of the case. There is quite evidently no justification for Fritsch's conclusion of the branchiate nature of Dolichosoma. There is no evidence of any gill-like structure in the American snake- like amphibians of the Coal Measures. Reviewing, then, the characters of the group which have been assigned by vari- ous observers, it will be seen that there is but a single character which holds good: "body long and snake-like." This is totally insufficient for the retention of the group. I therefore propose to abolish the group entirely from zoological classifica- tion. It is not even a family. It will, however, be convenient to refer to the snake- like forms as "aistopodous." CHAPTER XIII. THE MICROSAURIAN FAMILY HYLONOMID/C, FROM THE COAL MEASURES OF NOVA SCOTIA. Family HYLONOMID^ Fritsch, 1883. Fritsch, Fauna der Gaskohle und der Kalksteine der Perm formations Bohmens, Bd. 1, p. 159, 1883. Lydekker, Cat. Fossil Reptilia and Amphibia, iv, p. 201, 1890. The following characterizations of the family are those given by Lydekker (393, p. 201) based on Fritsch (251): Body slender and lizard -like; skull narrow, with smooth or faintly sculptured bones ; neural spines of vertebras well developed, and long, slender ribs. Teeth smooth, or with grooved summits. The whole body covered with sculptured scutes. Internal gills may be developed. Fritsch (251, Bd. 1, p. 159) gives the following in his original description: " Stegocephali von Baue schlanker Eidechsen mit schlanken langen Rippen. Wirbel amphicoel mit stark entwickelten oberen Dornfortsatzen. Schadelknochen glatt oder schwach verziert. Schuppen gross, verziert, den ganzen Korper deckend. Zahne glatt oder mit verzierte Spitze. Kiemenbogen bei einigen angedeutet. Mittlere Kehlbrustplatte unbekannt. Coracoidea ahnlich wie bei Branchiosaurus schlank, winkelig gebogen." Fritsch includes the following genera in the family : Hylonomus, Hyloplesion, Smilerpeton, Seclcya, Orthocosta, and Ricnodon. The family includes the following species: Hylonomus lyelli Dawson, Nova Scotia. Hylonomus wildi Woodward, England. latidens Dawson, Nova Scotia. Smilerpeton aciedentatum Dawson, Nova Scotia. multidens Dawson, Nova Scotia. Hylerpcton dawsonii Owen, Nova Scotia. wymani Dawson, Nova Scotia. intermedium Dawson, Nova Scotia. iritschii Gein. and Deichm., Saxony. longidentatum Dawson, Nova Scotia. geinilzi Credner, Saxony. Fritschia curtidentata Dawson, Nova Scotia. (?) pictus Fritsch, Bohemia. Genus HYLONOMUS Dawson. Dawson, Quart. Jour. Geol. Soc. London, xvi, p. 274. figs. 14-18, i860 Dawson, Air-breathers of the Coal Period, p. 44, 18(13. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 634. Credner, Zcit. d. dcutsch. geol. Gesell., 1890 (ix Thcil, die Stegocephalen und Saurier). Type : Hylonomus lyelli Dawson. The genus Hylonomus is a very important one from a taxonomic viewpoint, since it was regarded by Dawson (216, p. 635) as the typical genus of the order Micro- sauria, the most abundant group in the Carboniferous. Unfortunately the species of the genus Hylonomus are known only from fragmentary remains. I have re- produced in plate 9 Dawson's figures of the remains of Hylonomus as published by him in 1891. Dawson (216) gave, in 1882, the following definition of the genus Hylonomus: "Form lizard-like, with the posterior limbs somewhat large in proportion to the anterior. Size, small. Mandibular and maxillary teeth numerous, small, conical, pointed. Palatal teeth minute. Abdominal scales oval. " 78 MOOOti PLATE 9 Hylonomus lyelii Dawson. 1, maxillse and skull bones; la, sternal bones; 2, mandible; 3, humerus, ribs, and vertebra; 4, posterior limb; 5, pelvis; 6, caudal vertebrae. Nearly natural size. Erect tree, Coal formation, South Joggins, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original in the British Museum. THE MICROSAURIAN FAMILY HYLONOMID^. 79 Credner (186), Fritsch (251, Bd. 1, p. 89, Taf. 12, figs. 1, 4, 15), and Woodward (629) have referred remains of Microsauria discovered in the Coal Measures or lower Permian deposits of Saxony, Bohemia, and Lancashire, England, to the genus Hylonomus. There is much uncertainty as to the validity of these references, due to the uncertain nature of the type of Hylonomus. There are 4 American species of the genus: Hylonomus lalidens Dawson, II. lyelli Dawson, II. multidens Dawson, and //. wymani Dawson. All the species are from the Coal Beds at the South Joggins, Nova Scotia. Hylonomus lyelli Dawson. Dawson, Quart. Jour. Geol. Soc. London, xvi, p. 274, figs. 14 to 18, i860. Dawson, Air-breathers of the Coal Period, p. 44, 1863. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 635, pi. 39, figs. 1 to 14 and 27. Dawson, Acadian Geology, 3d ed., 1880, p. 370. Type: Specimens Nos. R 443 to 445 in the British Museum (393, pt. iv, p. 223). Horizon and locality : Coal formation of the South Joggins, Nova Scotia. This species is by far the most abundant (plate 9) in the erect trees examined by Dawson. Its characters Dawson (216) defines as follows: ' ' General form lizard-like, with the hind limbs rather larger than the fore limbs. Length when mature, 5 to 6 inches. "Head somewhat elongate; bones of skull smooth or with microscopic striae, perfectly united, except at the parietal foramen. Occipital condyle double, and apparently bony. Teeth simple, conical, numerous, about forty in each mandible, and nearly equal, except that a few of the anterior ones are rather larger than the others. The teeth are anchylosed to the jaw in a furrow protected by an external bony plate. ' ' Vertebras with cylindrical bodies, slightly concave at the ends. When partly exfoliated they appear hour-glass-shaped, in consequence of the internal cartilage having the form of two cones attached by their apices. Zygapophyses conspicuous above ; neural arches united to the bodies of the vertebrae, and with broad neural spines. Dorsal vertebrae with strong lateral processes. Caudal vertebrae apparently simple and cylindrical. Number of verte- brae in neck and trunk about thirty. "Ribs long and curved, with capitulum and tuberculum, cartilaginous within. "Anterior limb slender, humerus with distinct keel; radius and ulna separate; toes four or five. "Posterior limb with well-developed femur; tibia and fibula shorter, separate; toes five, somewhat long and slender. "Pelvis large, composed of ilium and ischium." Interclavicle and numerous scutellse are present. Upper surface protected with imbricated horny scales. In front two rows of horny tubercles and plates, with epaulettes composed of bristle-like fibers projecting from the skin. The animal possibly fed on insects, as is indicated by the coprolitic matter asso- ciated with the remains of the species. The following measurements are given by Dawson for the largest individual discovered: Length of head about cm. 2 Length of rib cm. 1.3 Length of neck " " 1.3 Length of humerus " 1.4 Length of trunk " " 7 Length of femur " 1.8 Length of posterior limb to heel cm. 3 Length of tibia " 1.2 Length of mandible " 1.8 Length of body of vertebra mm. 2.5 Teeth, 5 in 1 mm. 80 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Hylonomus latidens Dawson. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. ii, p. 637, pi. 39, figs. 18-22. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, p. 74. Type: Specimen No. 3061-1, Peter Redpath Museum, McGill University. The British Museum (393, pt. iv, p. 224) also has a specimen, No. R 447. Horizon and locality: Coal formation at the South Joggins, Nova Scotia. Fragments of 3 specimens from 3 trees represent this species (plate 10). It seems to have been of stouter build than II. lyelli, with the limbs shorter in proportion. Its generic affinities are somewhat doubtful, as it presents in some respects characters intermediate between Hylonomus and Hylerpeton . Mandibular and maxillary teeth broadly conical, about 20 in each mandible— 3 in 1 mm. ; anterior mandibular teeth somewhat larger than the others, and bent or hooked. Vomer or palate with minute teeth. Thoracic plate large. Scales of abdo- men oval, but somewhat narrow, and tending to be oat-shaped. Length of mandible (imperfect) mm. 9 Length of tibia (?) mm. 5 Length of humerus mm. 7 Length of thoracic plate cm. I Length of vertebra mm. 2 Length of six caudal vertebra; mm. 8 Hylonomus multidens Dawson. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt.- 11, p. 637, pi. 39, figs. 23-26. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, p. 74. Type: Specimen No. 3061-2, Peter Redpath Museum, McGill University. Horizon and locality: Coal formation at the South Joggins, Nova Scotia. This animal is known only by portions of bones of the head and a few other frag- ments. The scattered bones of the extremities are inseparable from those of II. lyelli occurring with it. As compared with that species, the bones of this are smoother and more delicate. The teeth are more numerous and slender. The crushed distal end of a femur or humerus found near the skull indicates that the limbs were well developed. mm. Length of mandible 1 1 Length of skull 15 Length of femur 9 Teeth, 5 to 6 in 1 mm. Hylonomus wymani Dawson. Dawson, Quart. Jour. Geol. Soc. London, xvi, p. 277, figs. 27-29, i860. Dawson, Air-breathers of the Coal Period, p. 52, 1863. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 637, plate 39, figs. 15-17. Dawson, Acadian Geology, 3d ed., p. 378. Type: Specimen No. 3061, Peter Redpath Museum, McGill University. There is also specimen No. R 446 in the British Museum (393, pt. iv, p. 224). Horizon and locality: Coal formation at the South Joggins, Nova Scotia. As compared with the II. lyelli the present species is smaller in size, more elon- gated in form, had the teeth less numerous (about 22 in the mandible), and shorter and more obtuse in form. There are 6 to 7 in 1 mm. This species is much more rare than II. lyelli, but quantities of minute bones, probably belonging to it, occur in the coprolitic matter. In one specimen 38 verte- PLATE 10 Hvlonomits latidriis Dawson. Skull, portion of skeleton, foot, scapula, sternal bones, humerus, and rib, believed to belong to this species. Erect tree, Coal formation, Nova Scotia. Nearly natural size. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original in the Peter Redpath Museum of McGill University. THE MICROSAURIAN FAMILY HYLONOMIOE. 8l brae of this species were found partially associated, indicating a long, slender body. The body is covered with scales and ventral scutellae are present. Dawson ques- tions whether this species may not be the young of H. lyelli. mm. mm. Length of skull 8 Length of femur 6 Length of mandible 5 Length of humerus 5 Length of rib 5.5 Genus SMILERPETON Dawson. Dawson, Phil. Trans.^Roy. Soc, London, 1882, pt. 11, p. 634. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, p. 74. Type : Smilerpeton aciedentatum Dawson. The type species was originaHy referred to Hylonomus, but further study induced Dawson to refer it to a new genus. Dawson gives (216) the following character- istics of the genus: "Form somewhat elongated, and limbs short. Mandibular and maxillary teeth wedge- shaped, with cutting edges. Palatal teeth numerous, some of them large. Abdominal scales oval. A single species is known, S. aciedentatum, from the Coal Measures at the South Joggins, Nova Scotia." Smilerpeton aciedentatum Dawson. Dawson, Quart. Jour. Geol. Soc. London, xvi, p. 275, figs. 19 to 23, i860. Dawson. Air-breathers of the Coal Period, p. 65, 1863. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. n, p. 638, plate 40, figs. 28 to 45. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, p. 75. Dawson, Acadian Geology, 3d ed., p. 376. Type: Specimen No. 3061-3, Peter Redpath Museum, McGill University. The British Museum (393, pt. IV, p. 224) also has a specimen, No. R 433. The important characteristic (plate 12) is found in the form of the mandibular and maxillary teeth, which are of a peculiar wedge-shape, being broad and oval at the base and narrowed to a longitudinal edge at top. Thus, when viewed from the side they appear narrow and blunt, but when the jaw is broken across, and they are viewed from the rear or front, they appear broad and sharp-edged. The effect of this arrangement is that the jaw is armed with a closely placed series of chisels or wedges, giving an almost continuous edge. At the end of the mandible some of the teeth are longer and more conical. Another important character is that the palatal and vomerine bones seem to have bristled with teeth, mostly of very small size; but there are also some larger palatal teeth, of which some are sharply pointed and others blunt with furrowed points. The vertebras are of the same type as those of Hylonomus; but some which appear to be caudal have a pointed spine above, indicating perhaps a flattened tail. The ribs are short and stout. The body seems to have possessed an interclavicle and ventral scutellae. Above it was, apparently, clothed with small tubercles and horny scales, and to have had cuticular pendants like those of Dendrerpeton. 82 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. An additional species of this genus was apparently indicated by some fragmen- tary remains, but Dawson thought best not to describe them as such, since they might indicate only a young individual of the present species. Length of mandible Length of femur Length of humerus (?) Length of vertebra Length of rib There are 5 teeth in 2 mm. 1.5 cm. 1.5 cm. 1.3 cm. 3-5 mm- 1 cm. 1 l/tfN c; Fig. 18. — Skeletal elements of Smilcrpeton aciedentatum Dawson, from the Coal Measures of Nova Scotia. (After Dawson.) a, shaft of femur? X 2; 6, intermaxillary _and teeth, X 25; c, sections of teeth, X 25; d and e, palatal teeth, X 25;/, femur, X 2; g, rib, X 2; h, palate; i, caudal vertebrae; j, long palatal tooth, X 25; k, bony scale. Genus HYLERPETON Owen. Owen, Quart. Jour. Geol. Soc. London, xvm, p. 241, 1862. Dawson, Amcr. Jour. Sci. (3), xn, p. 443, 1876. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 634. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, p. 74. Dawson, Air-breathers of the Coal Period, p. 55, pi. vi, figs. 32-46, 1863. Body stout, with strong limbs. Mandibular and maxillary teeth strong, not numerous, grooved at apex. Palatal teeth numerous, and some of them large. Thoracic plate broad. Abdominal scales pointed or oat-shaped. Type: Hylerpeton dawsoni Owen. Hylerpeton dawsoni Owen. Owen, Quart. Jour. Geol. Soc, xvm, p. 241. Dawson, Air-breathers of the Coal Period, p. 55, 1863. Dawson, Acadian Geology, p. 380. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. n, p. 639, pi. 41, figs. 62-85. Dawson, Proc. and Trans. Roy. Soc. Canada, 1894, xn, p. 74. Type: Specimen No. 3061-4, Peter Redpath Museum, McGill University. There are also specimens, Nos. R 441 and 442, in the British Museum (393, pt. iv, p. 225). (Plate 7.) Horizon and locality: Coal formation at the South Joggins, Nova Scotia. Bones of skull slightly striated, but not sculptured as in Dcndrerpeton. Lower jaw with distinct ascending ramus or coronoid process, a feature not known in any other of the Nova Scotia fauna, but observed by Cope in Brachydectes. Teeth, 12 in each ramus of the mandible, bluntly conical, slightly striated at the apex. Pulp- cavities large and longitudinally striated at the sides, though the teeth are not MOODII PLATE 11 Hylerpeton longidentatunt 'Dawson. Mandible and other bones. Nearly natural size. Erect tree, Coal formation, South Joggins, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original specimen in the Peter Redpath Museum of McGill University. PLATE 12 SmiUrpetOH aciedentatum Dawson. Mandible, portions of skull, scales, and various bones. Nearly natural size. Erect tree. Coal formation, South Joggins, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original specimen in the Peter Redpath Museum of McGill University. THE MK'KOSAURIAN FAMILY HYLONOMID^E. 83 folded. Maxilla furnished with similar teeth, one of which near the front is larger than the others. Palatal teeth numerous, small, conical, with a few large teeth at the sides. Vertebras short, cylindrical, well -ossified, with well-developed zygapophyses and neural spines; ribs strong and much curved, with well-developed division of the proximal ends; pelvis imperfect, but apparently large, with broad ilium. Humerus half the length of the mandible ; radius half as long as humerus ; femur very large and stout, nearly as long as the mandible; leg bones and phalanges corre- spondingly stout. The thoracic plate (plate 7) is indicated only by some fragments. The abdom- inal scales are narrow and pointed (oat-shaped) , smooth externally and with a ridge at one side within. The following are the dimensions of the largest specimen: Length of mandible 4.4 cm. Length of radius 1.5 cm. Length of largest tooth 5 mm. Length of vertebra 6 mm. Length of femur 3.5 cm. Length of rib 3 cm. Length of tibia 2 cm. Length of scales 5 to 7 mm. Length of humerus 2 cm. Hylerpeton longidentatum Dawson. Dawson, Am. Jour. Sci. (3), xn, pp. 440-447, 1876. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 640, pi. 42, figs. 86 to 109. Dawson, Proc. and Trans. Roy. Soc. Canada, 1894, xn, p. 74. Type: Specimen No. 3061-6, Peter Redpath Museum, McGill University. There is also a specimen, No. R 440, in the British Museum (393, pt. iv, p. 225). (Plate 11.) Horizon and locality : Coal formation at the South Joggins, Nova Scotia. Head much elongated, with the bones minutely pitted, and with delicate microscopic stria;, but not sculptured. Mandibular and maxillary teeth long and acute, pointing backwards, with the apex of their inner sides finely striated; 20 or more in each ramus of the lower jaw ; palatal bones with several long, slender teeth and many minute teeth. The mandibles found are not complete, but there are indications that there was an ascending process as in H. dawsoni, but less developed. The narrowness of the dentary bone is caused in part by the lower posterior edge being bent inward and by the posterior end being broken off above. Vertebra; short and stout, and apparently well ossified. Ribs long, with double head and much curved. Humerus longer than femur, which is short and stout, if the bone taken for it is rightly determined. Abdominal scales narrow, oat-shaped ; thoracic plate large, broadly oval. Measurements of Hylerpeton longidentatum Dawson. Length of mandible 4 cm. Length of femur (?) 1.2 cm. Length of vertebra 5 mm. Length of tibia 8 mm. Length of rib ' 3 cm. Length of mandibular teeth 3 mm. Length of humerus i-5 cm. 6 to 7 teeth in 1 cm. Hylerpeton intermedium Dawson. Dawson, Proc. and Trans. Roy. .Soc. Canada, xn, p. 75, 1895. Type: Specimen No. 3061-5, Peter Redpath Museum, McGill University. Horizon and locality: Coal formation at the South Joggins, Nova Scotia. This species is known only by the mandibles and portions of the skull, which are rather shorter than those of adult individuals of the last species. The extremity of 84 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. the mandible and the cranial bones have the same slightly waved surface as in the other species. Mandibles 3 cm. long and the teeth, which are about 15 in each ramus of the lower jaw, are simple, with large pulp cavities, those of the maxillary bone slightly enlarging upwards, and intermediate in form between the long, slender teeth of H. longidentatum and the thick, obtuse teeth of H. dawsoni. Coal formations, South Joggins, Nova Scotia, in erect tree, discovered by P. W. McNaughton, 1893. Genus FRITSCHIA Dawson, 1882. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. 11, p. 634. Lydekker, Cat. Fossil Reptilia and Amphibia, pt. iv, p. 225, 1889. Type: Fritschia curtidentata Dawson. Body lizard-like ; limbs large and well-ossified, mandibular and maxillary teeth conical, grooved at the apex. Abdominal scales slender and rod-like. Fritschia curtidentata Dawson. Dawson, Phil. Trans. Roy. Soc. London, 1882, pt. n, p. 641, pi. 43, figs. 1 10-128. Dawson, Am. Jour. Sci. (3), xn, p. 444, 1876. Type: Specimen No. 3061-7, Peter Redpath Museum, McGill University. There is also specimen No. R 449, in the British Museum (393, pt. iv, p. 225). Horizon and locality : Coal formation at the South Joggins, Nova Scotia. Represented by 2 specimens (plate 8). Bones of the head very smooth, only a few microscopic punctures. Teeth conical, somewhat obtuse, striated at the inner side of the apices; there are about 30 in each ramus of the mandible, and about 27 in the maxillary bone. Teeth implanted in a furrow. Vertebrae short and well ossified, 3 in 1 cm. Ribs strong, curved, about 1 cm. in length. Limbs robust, the bones better ossified than in any other known species from Nova Scotia. Toes of foot probably 5, central ones long and slender. Interclavicle of moderate size and somewhat rounded. Ventral scutellas needle-like. Length of mandible (imperfect) 2.1 cm. Length of femur 2.4 cm. Length of maxilla 2 cm. Length of radius and ulna 1 cm. Length of rib 1 cm. Length of toe in foot 7 mm. Length of humerus 2 cm. 8 teeth in 5 mm. CHAPTER XIV. THE MICROSAURIAN FAMILY TUD1TANID/C, FROM THE COAL MEASURES OF OHIO AND PENNSYLVANIA. Family TUDITANIDiE Cope, 1875. Cope, Geol. Surv. Ohio, 11, pt. n, p. 357, 1875. Lizard-like microsaurians ; cranial elements strongly sculptured with pits or grooves or almost smooth, with weak punctulations. Orbits usually well forward; squamosal sometimes excluded from the parietal ; skull hornless ; teeth pleurodont, conical and sharp, smooth or slightly plicate; clavicle of a triangular shape, which is characteristic of all the species; vertebrae well developed and phyllospondylous, the osseous portion being merely a hollow cylinder, hour-glass-shaped ; ribs curved, long, attenuated and intercentral ; digits clawed; ventral armature absent in all but a single species and the association of the species is doubtful; tail moderate in length. Three genera with 13 species included in the family. These species are : Tuditanus punctulatus Cope, Linton, Ohio. brevirostris Cope, Linton, Ohio. longipes Cope, Linton, Ohio. minimus Moodie, Cannelton, Pennsylvania. walcotti Moodie, Linton, Ohio. Erpetosaurus radiatus Cope, Linton, Ohio. obtusus Cope, Linton, Ohio. tabulatus Cope, Linton, Ohio. minutus Moodie, Cannelton, Pennsylvania. sculptilis Moodie, Cannelton, Pennsylvania. acutirostris Moodie, Linton, Ohio. tuberculatus Moodie, Linton, Ohio. Odonterpeton triangularis Moodie, Linton, Ohio. The association of these species in the one family is provisional and will need revision on the acquisition of new and more complete material. Genus TUDITANUS Cope, 1874. Cope, Trans. Amer. Phil. Soc, xv, p. 271, 1874. Cope, Geol. Surv. Ohio, 11, pt. 11, pp. 391, 1875. Type: Tuditanus punctulatus Cope. The genus as here defined is a somewhat composite group and it is quite proba- ble that some of the species here included will have to be removed to another genus when the anatomy of the forms is better known. The species of the genus are all moderately small, the largest barely attaining a length of 8 inches. There are 5 species of Tuditanus thus far known. All of the species are charac- terized by the possession of a peculiar triangular-shaped clavicle with radiating grooves, and this has been taken as one of the distinctive characters of the genus, as well as of the family. The structure of the cranium where known is quite uniform 85 86 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. among the different species. The squamosal is quite large and the supratemporal is not always closely joined to the parietal. The species are: Tuditanus punctulatus Cope, Linton, Ohio. brevirostris Cope, Linton, Ohio. longipes Cope, Linton, Ohio. minimus Moodie, Cannelton, Pennsylvania. walcotti Moodie, Linton, Ohio. Tuditanus punctulatus Cope. Cope, Trans. Amer. Phil. Soc, xv, p. 271, 1874. Cope, Geol. Surv. Ohio, n, pt. 11, p. 392, pi. xxxiv, fig. 1, 1875. Type: Specimen No. no, American Museum of Natural History, where there is also specimen No. in. Horizon and locality: Linton, Ohio, Coal Measures. This species, together with the form, T. brevirostris, describedonp. 88, was used by Cope as the type of the genus Tuditanus. Cope subsequently associated some rep- tilian remains from the Linton mines with the type of T. punctulatus and changed the generic term to Isodectes, which was known by another species, /. megalops Cope, from the Permian of Texas. The remains associated by Cope with I. megalops undoubtedly represent a reptilian species and which has been described elsewhere under the name Eosauravus copei Williston. The species is of exceeding interest because it is the oldest known reptile and places the range of the Reptilia down towards the base of the Pennsylvanian. The species Tuditanus punctulatus Cope was founded on well-preserved remains of nearly the entire skeleton of a single individual. The bones are represented by shining carbonaceous matter, and since both of the slabs containing the impression were preserved, a great many characters have been determined. The head, I fore- limb, and 23 consecutive vertebras with ribs are well defined, but of the pelvis and hind limbs nothing is visible. The cranium (fig. 19) is very similar to that of T. minimus, from the Cannelton, Pennsylvania, slates. It is triangular in shape, with a narrowed obtuse muzzle. The orbit of the left side is well defined and lies well forward. It is oval in outline and its width is about two-thirds of its length. The nostrils are small and are located well toward the tip of the muzzle. The parietal foramen lies behind the median transverse line which divides the skull equally. The cranial elements are for the most part destroyed, but the outlines of a few can be determined. Those elements which are preserved are ornamented with a sculpturing of minute punctulations which, on the postfrontal, assumes a radiating arrangement. The ornamentation of the other elements consists of inosculating pits, but they seldom assume the form of ridges or grooves. The bones of the premaxillary region of the cranium are lacking. The first element which can be detected is the prefrontal, which occupies a position in front of the orbit. There seems to be space for a lacrimal, but its outline is not distinct. The frontal can be readily separated and is seen to be an elongate element occupying the median region THE MICROSAURIAN FAMILY TUDITANID.«. 87 of the skull between the orbits. The parietal is apparently the largest element of the cranial roof and the pineal foramen is located in the anterior fourth of the median suture separating the parietal elements. The form of the postparietal and the tabu- lare can not be determined, as the greater part of this region is lacking. The squa- mosal seems to be located well forward and is rather small, but has the usual rela- tion of this element. Only fragments of the other ele- ments remain and nothing can be said of their form. The mandibles of both sides are represented by depres- sions, and they are ornamented with longitudinal grooves and ridges. The teeth are not preserved, but there are evidences of the maxillary teeth. These are minute and sharply conical. Just posterior to the skull there is pre- served the impression of a short, round rod which is not definitely determined. It may be an element of the hyoid apparatus, although it is rather stout for such. It does not have the relations indicated by Cope in his figure (123, pi. xxxiv, fig. 1). There are three elements of the pectoral girdle pre- served. These undoubtedly represent the interclavicle and the clavicles. The interclavicle is rhomboid in shape and is attenuated posteriorly. The attenuation is ab- ruptly truncate posteriorly and it is thus of quite a dif- ferent character from the acutely pointed interclavicle of T. minimus. The clavicle has a somewhat semicir- cular form, but is not attenuated at either end. It seems to be uniformly broad. The forearm of the right side is preserved in part. The humerus is seen to be a heavy, somewhat expanded element lying displaced with relation to the pectoral girdle. It is greatly expanded at the ends. The ulna presents characters similar to the humerus and only dif- fers from it in being shorter and less stout. The radius is not preserved. The carpus is unossified and its posi- tion is occupied by a blank space. The digits are repre- sented by 4 metacarpals, and this may have constituted the entire number of the fingers. The phalangeal bones preserved are a little scattered. They are elongate with expanded ends. Evidences of 23 consecutive osseous vertebrae are preserved. Their character can not be determined, al- though Cope (123) describes them as amphiccelous. This may be inferred to be the case, but I am unable to verify his observation. In form the vertebras are subquad- rate. The neural spines are not evident. The osseous ribs articulate, apparently, between the bodies of the vertebrae. Cope figured them as intercentral. There are 22 Fig. 19. — Drawing of skull and skele- tal elements of Tuditanus punctu- latus Cope from the Coal Meas- ures of Linton, Ohio. X 1.5. fr, frontal; ic, interclavicle; d, clav- icle; h, humerus; ph, phalanges; par, parietal; pp, postparietal; lab, tabulare, supratemporal and squamosal; u, ulna; r, radius. 88 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. or 23 pairs preserved. They are single-headed and the extremities are attenuated. No traces of ventral scutella are present. The entire length of the animal probably did not exceed 5 or 6 inches. Its form was quite lizard -like and it was probably of an ambulatory type, though it may have spent a part of its time in the old lagoon in which its remains were finally buried. No traces of external gills have been detected in this or any other Linton species. Measurements of the Type of Tuditanus punctulatus Cope. ram. mm. Length of entire specimen 94 Width of humerus at proximal end 2.5 Median length of skull 22 Length of ulna 6 Width of skull at occiput 19 Width of ulna at proximal end 2 Depth of mandibular ramus 4 Length of phalanx (metacarpal?) 2 Length of the 23 vertebra? 61 Expanse of longest ribs 16 Length of interclavicle 6.5 Length of rib 10.5 Width of interclavicle 4.5 Width of rib 5 Width of the three pectoral plates 10 Length of a vertebra 2.5 Length of humerus 8 Tuditanus brevirostris Cope. Cope, Trans. Amer. Phil. Soc, xv, p. 272, 1874. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 393, pi. xxvi, figs. 3, 4, 1875. Moodie, Bull. Amer. Museum Natl. Hist., xxvi, art. xxv, pi. lxiv, fig. 4, 1909. Type: Specimen No. 8609 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. This species was associated by Cope with the type T. punctulatus in the descrip- tion of the genus. Represented by a portion of the skeleton of one individual, the skull is preserved on one block, with a considerable part of the anterior ribs, pec- toral girdle, and vertebral column, although this last is not clearly represented, but as in so many of the coal specimens the bones are covered with a thin layer of car- bonaceous matter which makes it impossible to definitely determine the form. The cranium is large in proportion to the size of the body. The skull is in the form of a wide oval and is wider than it is long. The elements of the skull were orna- mented with a coarse sculpturing which partakes of the nature of incomplete radia- tions on the squamosal region. The different elements of the cranium can not be distinguished, although I think the outlines of the parietal are indicated. The position of the nostrils is well forward and they are slightly elongate transversely. The pineal foramen can not be determined. The orbits are oval in shape and their width is about equal to two-thirds of their length. The interorbital space is greater than the length of the orbit. Cope's figure of this specimen is not accurate, since he has the orbits drawn too far to the side. They are located near the central line of the skull and resemble in some respect those of the preceding species. Cope has described teeth in the maxillary region, but I am unable to detect them. There are portions of two pectoral elements which may represent a clavicle and a portion of the interclavicle. The clavicle has much the same shape and practically the same ornamentations as has the clavicle in Tuditanus minimus. The clavicle preserved shows a somewhat triangular shape and is slightly acuminate at the anterior end, as preserved, and obtuse at the posterior end. The nature of the interclavicle can not be determined. THE MICROSAURIAN FAMILY TUDITANID^. 89 The vertebral column is represented by a line which Cope suggests (123) may be the chorda dorsalis (notochord). Osseous vertebrae were probably present, but their nature is obscured by the carbonaceous matter covering them. The ribs as preserved are long and curved. They are slender and attenuated at the distal ends. They were probably single-headed, but whether their articulation was intercentral or not can not be determined. The other specimens which are referred to this species show nothing of impor- tance in the way of structure. They consist for the most part of fragments which may or may not represent the species. The species differs from the type of the genus (T. punctulatus) in the possession of a broadly rounded muzzle. This character will also separate it from other spe- cies of the genus. The sculpturing of the bones of the cranium is coarser in the pres- ent species than in the type. The form of the clavicle is different in the two species. The above-described species seems to be more closely allied to the form described as Tuditanus walcotti than to other species of the genus. I have been unable to detect the presence of limbs, although Cope says they are present. Measurements of the Type of Tuditanus brevirostris Cope. mm. mm. Median length of skull 15 Interorbital space 4.5 Width of skull at posterior border 18 Length of clavicle 7 Width of skull across orbits 1 1 .5 Width of clavicle 3 Length of orbit 3 Length of longest rib preserved 9 Width of orbit 2 Length of entire specimen 54 The material consists of the type specimen with counterpart and two fragments which probably are to be associated with this species. Collected by Doctor J. S. Newberry. Tuditanus longipes Cope. Cope, Trans. Amer. Phil. Soc, xv, p. 210, 1874 (Sauropleura). Cope, Geol. Surv. Ohio, 11, pt. II, 398, pi. xxvi, fig. 2, 1875. Type: Specimen No. 1099 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The cranium of this species is quite unknown. The only genus with which the specimen can be compared in the structure of the skeleton is Tuditanus. From the other species of the genus the present form differs in the presence of ventral chevron rods and the elongate character of the limbs, as well as in the possession of large iliac bones, which, in the only other species in which the ilia are known, are small and slender. It seems best to locate the species in this genus for the present, although it may eventually have to be removed to another group. Very little is known of the main portion of the skeleton of the species of Tuditanus, other than in T. longipes, so an exact comparison is impossible. From all the species of Tuditanus thus far known the present species differs in the elongate character of the limbs and in the presence of ventral scutellation. There are three other species of Tuditanus in which the limbs are known. These are: T. walcotti Moodie, T. minimus Moodie, and T. punctulatus Cope. In these three species the limbs are short and weakly developed. From the otherspecies of Tuditanus the present species maybe separated by its size principally, since nothing of the bodies of the other species is known. 90 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The body of the present species is elongate and slender, with a long neck and prob- ably a long tail. Ribs, as preserved, are 19 to 22, though there may possibly have been more. They are moderately curved backwards, have intercentral articulation, are attenuated at the distal extremity, and are single-headed. The anterior ribs are stouter, with a widened upper portion and attenuated distal part. The posterior ribs are more slender. There are evidences of 28 vertebrae present. All regions of the vertebral column are present and the dorsal region is preserved entire. The cervical series is repre- sented by the posterior vertebrae only. These are very indistinctly preserved. The dorsal vertebrae are elongate and were prob- ably amphiccelous, although this has not been definitely determined. They are ex- panded at each end, thus ending in a slightly raised rim. The single-headed ribs articulate between the vertebrae. The exact number of the dorsal series can not be ascertained, although this may have been 25. The spines of the vertebrae are not determinable, since the animal is preserved on its back. The caudal vertebrae are represented by two patches of the remains of what was once probably the entire series. Cope ascribes 70 mm. to the tail, but I do not find that much. The specimen may have been mutilated since he studied it. The caudals are slender and, like the dorsals, are expanded at the ex- tremities. The scapular arch is not preserved, but the pelvic arch is represented by the two iliac bones in good state of preservation. These are short, flat bones expanded at the anterior extremity, as preserved. They lie turned a little to each side of the vertebral column and fig. 2o.— Cope's drawing of TudUanus longipes, partially Obscure the femora. The iliac bones from the Linton, Ohio, Coal Measures. XI. are quite characteristic of this form, since similar-shaped elements have not been observed in any of the other Carboniferous forms from the same deposit. The greater part of the forelimb is preserved, although much of the hand is missing. The humerus is an unusually elongate bone and lies somewhat across the vertebral column. It is crushed flat and the ends are partly destroyed. It shows evidences of expansion at the ends, although not a great deal. It is much longer than the radius and ulna, which are of about equal length. The ulna is larger than the radius and has expanded ends, with the upper end more expanded than the lower and both ends slightly truncate. The radius is a simple rod of bone and is but slightly expanded. The carpus was evidently cartilaginous, since there is no evi- THE MICROSAURIAN FAMILY TUDITANID/E. 91 dence of osseous material in its place. There is but one phalangeal bone preserved, and, since this is displaced with reference to the ulna and radius, its position can not be determined. It may have been a metacarpal. It is short and expanded at the ends. The hind limbs are represented by the two femora and the upper portion of the tibia. The femur is almost as elongate as the humerus and is more slender. It is not so much expanded as the humerus. Its ends appear to have been cartilaginous and do not represent the well-formed articular surfaces preserved in the T. minimus. The upper part of the tibia is preserved, and appears to have been truncate. If this species belongs with Tuditanus it is of interest in that the ventral chev- rons are present. The species is particularly characterized by the elongate limbs. Measurements of the Type of Tuditanus longipes Cope. mm. mm. Length of vertebral column between pelvis and end Length of radius and ulna 12 of humerus 70 Length of a vertebra 3 Length of vertebral column anterior to humerus 18 Length of ilium 7 Length of caudals present 42 Length of femur, estimated 18 Length of humerus 19 7 chevrons in 4 mm. Width of humerus 2 Tuditanus minimus Moodie. Moodie, Jour. Geol., xvn, No. [, p. 56, fig. 10, 1909. Moodie, Proc. U. S. Nat. Mus., 37, p. 23, pi. 8, fig. 2, 1909. Type: Specimen No. 4555, U. S. National Museum. Horizon and locality : Cannelton slates of Pennsylvania (Upper Freeport) . The species is represented by a nearly complete skeleton preserved on a slab of slate from the Cannelton shales of Pennsylvania. The obverse slab has been lost, which is very unfortunate, since there is no doubt that the entire skeleton was origi- nally present. The species is placed in the genus Tuditanus on account of the close resemblance to the type form T. punctulatus Cope, although it is much smaller than that species. The type specimen of the species did not attain a length of more than 3.5 inches. Its form is very lizard-like, but its structure is typically amphibian. The form of the skull is especially similar to that of the type species T. punctulatus, which it resembles in the narrow posterior truncation of the skull, as well as in the anterior position of the orbits. The skull is in the form of a narrow oval, sharply narrowed posteriorly and trun- cate. The orbits are located well forward and their posterior border lies in front of the line dividing the skull transversely into equal parts. The interorbital space is greater than the diameter of the orbit. Impressions of teeth are preserved on the premaxilla? and maxillae; there are 8 of them in a distance of 3 mm. The teeth appear to be mere blunt denticles and were possibly pleurodont. The elements of the cranium are very poorly preserved. It has been impossible to determine all of the sutures. The bones of the premaxillary region have been destroyed, but the arrangement of them was probably not far different from that which obtains in other members of the genus. The posterior boundaries of the nasals 92 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. are preserved and prove this element to have had an obtuse posterior border. The sutures bounding the frontals are clear and show that they were small and that they formed a part of the inner boundary of the orbits. The parietal is recognized as a large element, apparently the largest in the skull. Together the parietals form a wide oval inclosing, on the median suture, the circular pineal foramen. The parie- tals are sculptured with coarse radiating grooves and ridges, much after the manner of Erpetosaurus radiatus Cope. The pittings present on that form are, however, absent in T. minimus. The sutures bounding the postparietal are tolerably well defined and these show that element to have been rather large and quadrate, with the usual relations. The tabulare is distinct, triangular, and small. It is produced into an angle on the posterior border strongly recalling a similar condition in T. punctulatus. The boundaries of the prefrontals and the upper borders of the max- illae are not clearly ascertained. The lacrimal has not been detected. The post- frontal and postorbital form the posterior boundary of the orbit, although all of the limits of the latter element have not been definitely determined. The position of the supratemporal is well assured, although its entire boundaries are not determined. It has the usual relations and joins the parietal broadly. The jugal is broad and widens posteriorly to join the squamosal, which, as usual, forms the quadrate angle of the skull. The sutures bounding the quadratojugal and the posterior end of the maxilla are not determined. There are but two fragmentary vertebras preserved and an estimate based on the length of these remains gives about 30 presacral vertebrae. The structure of the vertebrae preserved can not be ascertained, but the neural spines appear to have been low and stout. There are six elements of the pectoral girdle preserved. These are: the six clavicles, the interclavicle, the coracoid of one side, and the two scapulas. The interclavicle is rhomboid in form and acuminate posteriorly. It is sculptured with radiating grooves and ridges. It is quite different from the same element in T. punctulatus, in that the base is acuminate, not truncate. The clavicle presents much the same shape as does that element in Erpetosaurus tabulatus. It is ornamented by a sculpturing of radiating lines which take their origin from the lower external angle as the bone lies in the matrix. The clavicle is somewhat trian- gular in shape and lies close to the skull, but this close approximation of the pecto- ral elements to the cranium is due probably to post-mortem shifting, since the scapulae are shifted backward. There can be little doubt, however, that the pectoral arch was close to the cranium. There is an oval fragment preserved on the left of the specimen which I take to be a portion of the coracoid. The scapula is preserved entire on the left side and is represented by fragments on the right side. It is almost semicircular in form and narrows externally until it is somewhat fan-shaped. There appears to be an ornamentation of lines on the surface of the bone. These lines follow the contour of the anterior border. The arm is preserved nearly complete on the left side, and the right side shows the humerus and the forearm. The humeri are unusual in having well-developed THE MICROSAURIAN FAMILY TUDITANID^E. 93 articular ends as though the endochondral tissue was well developed. The hume- rus is expanded at the ends and it is larger at the upper than at the lower end. The ulna is expanded at the proximal extremity, but is more attenuated at the distal portion. It is shorter than the humerus by about one-third of its own length. The radius is a mere slender rod of bone and presents well-developed articular ends. It is slightly shorter than the ulna. The carpus is unossified and its position is repre- sented by a blank space. There are phalanges of 4 digits preserved and they are 4 in number. The phalangeal elements, like the other bones of the extremity, have the articular surfaces prominent, with the terminal phalanx claw-like. There are no ribs nor traces of them preserved, and a conjecture as to their character can not be hazarded, since they are known in but two other species, in which they are slender and curved. There is no evidence of a ventral scutellation, and so far as is at present known this structure is absent from all of the species of the genus, or at least it is weakly developed. The ilium is all that is preserved of the pelvis. The bone itself has disappeared and has left a depression which shows this element to have been an elongate rod very similar to that described for Micrerpeton. The sacral vertebra seems to be indicated by a depression between the iliac depressions. One hind limb is preserved nearly entire and the greater part of the other is also preserved, although some of the phalanges are disturbed. The femur is slender and more elongate than the humerus. It has well-formed, rounded, articular ends. The tibia presents unusual characters in that its ends are truncate, as though the car- tilage composing its articular surfaces was not so highly calcified as in the other limb bones. It is somewhat expanded at the ends and is throughout its length broader than the femur. The fibula, like the tibia, is a slender rod of bone, although it is somewhat shorter than the tibia. The tarsus is unossified and its position is occupied by a blank space. Portions of both feet are preserved, but only one digit in the right foot is complete. The metatarsals are elongate and slightly expanded at the ends. There are 4 phalanges present in the complete digit, which possibly represents the fourth. The first digit is wanting, with the only terminal phalanx preserved claw-like. Measurements of the Type of Tuditanus minimus Moodie. mm. mm. Median length of skul! 15 Width of scapula, maximum 2.5 Width of skull at posterior border 16 Length of coracoid (?) 2 Length of orbit 3.5 Length of humerus 4 Width of orbit 2 Length of radius and ulna 3 Interorbital width 2.5 Length of metacarpal I Length of clavicle 6 Length of ilium 2.5 Width of clavicle, maximum 3.5 Length of femur 4.5 Length of interclavicle, estimated 5 Length of tibia and fibula 3 Width of interclavicle 3.5 Length of foot ; 3.5 Length of scapula 3.5 Length of metatarsal 75 Tuditanus walcotti Moodie. Moodie, Proc. U. S. Nat. Mus., xxxvn, p. 16, pi. 6, figs. 1, 2; pi. 7; 1909. Type: Specimen No. 4474, U. S. National Museum. Horizon and locality: Linton, Ohio, Coal Measures. 94 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. A small species of Microsauria is preserved as a smooth impression on a block of soft coal from Linton, Ohio. Nearly the entire form of the body is discernible. The specimen is especially interesting and valuable as exhibiting for the first time among the Linton forms the shape of the body of the small microsaurians of the Tuditanus type. It differs so markedly in the form of the skull from other species of the genus that it is regarded as a distinct form, and the name Tuditanus walcotti was proposed for it as an expression of the writer's indebtedness to the Secretary of the Smithsonian Institution for the use of the material among which the present form was included. The specimen includes, besides the body impression, the complete skull, a right clavicle, with portions of the left, a left humerus, 12 cervical and dorsal vertebrae, 10 pairs of ribs somewhat disturbed as to position, and a portion of the mandible. There are no traces of ventral scutellae nor body scales in the smooth impression of the carbonized skin. One would expect to find impressions of the ventral scutse in this specimen if they were present. Cope remarked on the apparent absence of scutellae from members of the genus Tuditanus as they were known to him, and no contrary evidence has since been brought to light. Until such evidence is forthcoming the absence of scutes will be taken as one of the generic characters of the genus Tuditanus. Under a magnification of 50 diameters the carbonized skin shows as folds and wrin- kles, like muscle fibers, in some places ; in others no traces of the muscular structure can be detected. The wrinkles may be impressions of the internal musculature of the body -wall of the abdomen. It is especially well preserved in the pelvic and pygal regions. Sections of the coal were made, but nothing definite could be determined as to the character of the impressions, as they were too poorly preserved and the coal was too soft to bear much handling. The specimen is preserved on the belly, with the dorsum of the skull uppermost. It has been practically impossible to determine the arrangement of any of the cranial elements except the frontals, parietals, and postparietals, which have the relations indicated in figure 21, A. A median suture is clearly evident, with the pineal foramen well back in this suture. The bones of the skull are marked with faint radiating lines. It is in the form of the skull and the position of the orbits that the specific char- acters are found, as follows: the backward position of the eyes and the oval, pointed shape of the skull. The species is closely related to Tuditanus minimus Fig. 21. — A. Outline drawing of type of Tuditanus walcotti Moodie, from the Coal Measures of Linton, Ohio, showing impres- sion of body and muscle at M. X 25. cl, clavicle; fr, frontal;/, femur; h, humerus; nos, nostril; or, orbit; par, parietal; rb, rib; pp, postparietal ; v, vertebra; pfo, pineal foramen. B. Left leg of second specimen of Tuditanus walcotti. X 3. THE MICROSAURIAN FAMILY TUDITANID/E. 95 Moodie, from the Cannelton slates of Pennsylvania, and serves further to connect the forms from the Ohio and Pennsylvania localities. It differs from the last-named species in the position and form of the orbits, these structures being more oval in the present form and placed further back. The shape of the skull differs also in the almost entire absence of the posterior table. The median points of the orbits occupy the line which bisects the skull, and the interorbital width is less than the width of the orbit. The mandible is heavy and appears to have borne sharp, pleurodont teeth. The vertebral column is represented by little more than a mold of the form of the vertebrae, so that little can be said of its character. The individual vertebrae are short and hour-glass-shaped. The ribs are borne intercentrally, as in all the micro- saurians which have been studied from the Linton deposits. The ribs are rather long and somewhat heavy, slightly curved and expanded at the proximal end, as though an incipient bicipital condition were present. The right clavicle, which is preserved as an impression, is entire. Its impres- sion shows this element to have been ornamented on its ventral surface with radi- ating grooves and ridges which started at the lower angle of the bone. The element is distinctly triangular, which is characteristic of the genus Tuditanus, so far as known. The fragment of the left clavicle adds nothing to our knowledge of the element. The left humerus recalls in a striking way that of Tuditanus longipes Cope, and it was once entertained as a possibility that the present form might be a member of that species, since the skull is lacking in T. longipes. Sufficient specific differences were found, however, in the ribs, which, in T. longipes, are very long, slightly curved, and delicate, but which, in the present form, are comparatively heavy. Other char- acters sufficiently diagnostic are found in the form assumed by the vertebrae in the two forms. Measurements of the Type of Tuditanus walcotti Moodie. mm. mm. Length of specimen 70 Length of vertebral column, as preserved 50 Length of skull 20 Length of a vertebra 1.75 Posterior width of skull 14 Width of a vertebra 50 Width of skull, anterior to orbits 10 Width of body impression 15 Length of orbit 4 Length of humerus 6 Width of orbit 2 Median width of humerus 50 Interorbital width 3 Width at end of humerus 2 Length of clavicle 9 Length of rib 8 Greatest width of clavicle 4 Width of rib 25 The above-described specimen was collected by Mr. R. D. Lacoe, of Pittston, Pennsylvania, from Linton, Ohio. A second individual (No. 4481, U. S. National Museum) of this species is indi- cated by a rather poorly preserved specimen on a slab of soft coal from the Linton mines. The following portions of the animal have been detected and will be dis- cussed: partial impression of the skull, with a fragment of a minute jaw, in which are minute teeth ; right clavicle ; part of the impression of the body ; nearly entire left hind limb ; impressions of about a dozen vertebrae, very indistinct. The impression of the skull is distinct only in a favorable light, and even then the boundaries of the cranium are a little uncertain. For this reason no representation g6 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. of the form will be attempted. The sculpturing on the parietals is, however, dis- tinct enough to show relationship with the previously described specimen, and the form of the body impression, the absence of abdominal scutes, the shape of the clavi- cle and its sculpture, and the proportions of the hind limbs all agree with the charac- ters which have been assigned to the genus Tuditanus. The fragment of the jaw is interesting as giving the first information as to the character of the mandible in the genus Tuditanus. It is very slender and of uniform width as far as preserved. The teeth are short, blunt cones, apparently pleurodont. The clavicle is of the typical Tuditanus form, with the sculpturing lines radiating out from the angle. The impression of the body adds nothing to that already described for the type specimen. The nearly entire hind limb is of great interest as adding another example of the phalangeal formula. The foot is almost perfectly preserved, and the formula was probably 2-2-3-3-2. The endochondrium of the limb bones is not highly developed. About a dozen vertebrae are represented by molds in the soft coal, but nothing of their structure can be determined. The sharp, reptile-like claws in which the toes end (fig. 2 1 , B) recall those of Eosau- ravus and of Tuditanus minimus Moodie. It is another link in the chain of the suggested relationship between the microsaurians and the early reptiles. Measurements of the Second Specimen of Tuditanus walcotti Moodie. mm. mm. Length of entire body impression 75 Length of clavicle 8 Width across belly, maximum 16 Width of clavicle, maximum 4 Length of skull ?I7 Length of hind limb 22 Posterior width of skull ?I4 Length of femur 8 Length of fragment-of jaw 4 Length of tibia (?) 6 Width of fragment of jaw 1.5 Length of metatarsal 2 Length of tooth in jaw .25 Length of first digit 6 Genus ERPETOSAURUS Moodie, 1909. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, p. 348, fig. 1, 1909. Moodie, Proc. U. S. Nat. Mus., 37, p. 21, 1909. Type: Erpetosaurus radiatus Cope. Skull stout, elements sculptured with radiating grooves, ridges, and pits; orbits large and usually placed far forward ; occiput sometimes with posterior table ; skull more or less rounded; lateral-line canals consisting of supraorbital, suborbital, jugal, and temporal canals, the last two uniting to form a circular canal in one species; clavicle triangular, sculptured like the skull. Our knowledge of the genus is confined to the skull. The genus was established for certain members of the genus Tuditanus and other forms which have been re- cently described. The species of the genus are: E. radiatus Cope type, E. tabulatus Cope, E. tuberculatus Moodie, E. obtusus Cope, E. minutus Moodie, E. acutirostris Moodie, E. sculptilis Moodie. All of the species are from the Linton, Ohio, Coal Measures, with the exception of E. sculptilis and E. minutus, which are from the Cannelton, Pennsylvania, slates. The position of the genus as to family is a little uncertain, since family charac- ters are not yet well understood among the Carboniferous forms on account of the lack of information as to the structure of the animals. If we take the absence of THE MICROSAURIAN FAMILY TUDITANID^E. 97 the ventral scutellae as a family character, the genus will be in the family Tuditan- idae, but the evidence on this point is negative. For the present we may place the genus only provisionally in the family Tuditanidae. The arrangement will undoubt- edly require revision later. Erpetosaurus radiatus Cope, 1874. Cope, Trans. Amer. Phil. Soc, xv, p. 273, 1874. Cope, Geol. Surv. Ohio, 11, pt. 11, pp. 394-395, pi. xxvii, fig. 1; pi. xxxiv, fig. 3; text-fig. 10, 1875. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, p. 348, pi. lxii, fig. 1, 1909. Type : Specimen No. 8600 G, American Museum of Natural History. Locality and horizon: Linton, Ohio, Coal Measures. (Plate 25, fig. 1.) Cope originally described this species from a portion of a skull. He (123) char- acterized the form as follows: "The marked character of this form is seen in the very anterior position of the orbits and the contraction of the muzzle. The orbits are large and are separated by a little more than their own diameter; their posterior border is in front of a line measuring the anterior third of the length to the supraoccipital crest, and nearly to the line marking the fourth of the length to the quadrate region. The posterior outline of the skull is deeply concave, the quadrate angle projecting beyond the occipital condyles." The base of the specimen is broken and there is no place for the occipital condyles. Unless the specimen has been mutilated since Cope studied it, the occipital condyles are not present. The restoration of the skull given in figure 22, B varies but little from that given by Cope in 1875. The elements are practically as he represented them. The premaxilla? are small and lie in the usual relations to the other elements. Minute conical teeth are present as impressions. They are quite similar to the teeth found in other Microsauria. The nasals are nearly square and form the inner boun- dary of the somewhat oval nostril, which is represented by a depression in the coal. The frontal is elongate. It is about twice as long as wide. It forms a portion of the inner border of the orbit, the remainder being made up by the prefrontals and the postfrontal. The parietals are the largest elements of the skull, but they do not greatly exceed the jugals. Together the parietals form a somewhat obtuse oval in the median region of the skull and they contain between them, in their posterior third, the small circular pineal foramen. The postparietal forms the posterior boundary of the skull. The prefrontal forms the anterior border of the orbit and is triangular in shape. The lacrimal is not identified. The maxilla is an elongate element the boundaries of which are uncertain, though probably somewhat as given. The postfrontal and the postorbital form the posterior boundary of the orbit, inclosing between them the anterior projection of the squamosal. The squamosal is an elongate element and is acuminate at each end. The tabulare is a large element lying lateral to the postparietal. The jugal is a very elongate element, apparently acuminate anteriorly. The quadratojugal is small and elongate. The supratemporal is definitely bounded and its limits are as indicated (fig. 22 B), being a large element which forms the quadrate angle. There are two other specimens of this species in the collections and a fragment of a fourth which it is difficult to make out. Cope identified and figured one of these 98 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. as E. radiatus (Geol. Surv. Ohio, II, pt. II, pi. 34, fig. 3), but the identification is doubtful and the figure shows structures which I am unable to identify in the speci- men. Nothing of importance is to be learned from the other two specimens, except that they show a diversity of size. They consist of incomplete skulls, concerning which Cope (123) remarks: "There are no mucous canals. The sculpture consists of strong ridges radiating and inosculating. Radiation is more uninterrupted on both jugal, supratemporal, and anterior part of the tabulare; on the first they originate in front of the middle exteriorly; on the supratemporal near the anterior part. The inosculation is honeycomb-like on the parietal, postfrontal, and posterior parts of the tabulare." Measurement of the Type Specimen. mm. mm. Length of the skull along median line (estimated) ... 60 Length of orbit 9 Length from muzzle to quadrate angle 71 Width of orbit 8 Width at posterior border 69 Interorbital width 8 Width at orbits 40 Length of nostrils 2 Measurements of Another Specimen of the Species. (No. 8598 G, American Museum of Natural History.) mm. mm. Median length of skull 56 Length of orbit 7 Length to quadrate angle 61 Width of orbit 6 Width at posterior border 50 Interorbital width 4.5 The specimens of this species were collected by Dr. J. S. Newberry. Erpetosaurus obtusus Cope, 1868. Cope, Proc. Phil. Acad. Nat. Sci., 1868, p. 213. Cope, Trans. Amer. Phil. Soc, xiv, p. 12, fig. I, 1869. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 396, fig. 11, 1875. Cope, Proc. Amer. Phil. Soc, 1885, xxn, p. 407 (Pal. Bull. 40). Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, p. 350, pi. lx, fig. 2. Type: Specimen No. 8601 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The species, Tuditanus obtusus, was first described by Professor Cope as Den- drerpeton obtusum, but he subsequently referred the species to the genus Tuditanus. It was removed by the writer to the genus Erpetosaurus in 1909. The species is known from two partially preserved crania. The skull elements seem to have disap- peared and left only the impressions. The sutures are, for the most part, clearly represented, but the skull shows no sculpturing. On the posterior third of one cranium there is a small space which seems to be slightly sculptured, as Cope indi- cated in his drawing. The general form of the skull is that of a broad oval, truncate posteriorly. The orbits lie in the anterior third of the skull, and the pineal foramen in the posterior third. Cope compared the skull to that of Huxley's Erpetocephalus, to which it has some resemblance. The nostrils are elongate and are situated at an obtuse angle with relation to the main axis of the skull. The premaxilla is small and forms the inner border of the nostril. There seem to be impressions of small teeth, but no large ones are evident. The nasals are sepa- rated by a zigzag suture, and are nearly square. They have the usual relations. The frontals form a portion of the inner boundary of the orbits and unite behind THE MICROSAURIAN FAMILY TUDITANID/E. 99 with the parietals, the anterior extensions of which they inclose. The parietals are large elements and together form a broad oval, truncate posteriorly. They inclose between them the pineal foramen in the median suture. It lies in the posterior fourth of the parietals. The postparietals are elongate transversely, and have the usual relations. The prefrontal is somewhat triangular and forms the anterior boundary of the orbit. The lacrimal has not been detected. The maxilla is elon- FlG. 22. A. ( hillinc of skull and cranial elements of Erpetosaurus minulus Moodic, from the Cannelton slates of Pennsylvania. Original in U. S. National Museum. X 2. fr, frontal; mx, maxilla; par, parietal; po, postorbital; pp, post- parietal; pr, postfrontal; sq, squamosal; spt, supratemporal; tab, tabulare. B. Outline of skull and cranial elements of Erpetosaurus radialus Cope, from the Coal Measures of Linton, Ohio; partially restored. X 0.75. Original in American Museum of Natural History, fr, frontal;./, jugal; mx, maxilla; «, nasal; or, orbit; par, parietal; pf, postfrontal; po, postorbital; pr, prefrontal; pp, postparietal ; pmx, premaxilla; qj, quadra to jugal; sq, squamosal; spt, supratemporal; tab, tabulare. C. Palate of Erpetosaurus {tabulatus?) , from the Coal Measures of Linton, Ohio. X 1. American Museum of Natural History, ex, exoccipital; m, mandible; mx, maxilla; pal, palatine; pt, pterygoid; prv, prevomer; pd, para- sphenoid ; *, anterior and posterior palatine vacuities. D. Outline of skull and cranial elements of Erpetosaurus acutiroslris Moodie, from the Coal Measures of Linton, Ohio. Original in American Museum of Natural History. X I. ex, exoccipital ;fr, frontal; j, jugal; mx, maxilla; n, nasal; or, orbit; po, postorbital; par, parietal; pp, postparietal; pf, postfrontal; sq, squamosal; spt, supra- temporal; tab, tabulare. B, ( Hitline of larger part of skeleton of Odonterpeton triangularis Moodie, from the Coal Measures of Linton, Ohio. Original in U. S. National Museum. X 4. h, humerus; il, ilium?; v, vertebrae, all of which are not represented. F. Right mandible of Erpetosaurus tabulatus Cope, from the Linton, Ohio, Coal Measures. Original in American Museum of Natural History. X 2. art, articular; cor, coronoid; d, dentary; ang, angular; sa, surangular. G. Skull elements and lateral-line canals of Erpetosaurus tabulatus Cope, from the Coal Measures of Linton, Ohio. X 1.5. fr, frontal; j, jugal;//, jugal lateral-line canal; mx, maxilla;n, nasal; or, orbit; par, parietal; po, postorbital ; pf, lacrimal or prefrontal ; so, suborbital lateral -line canal; pp, postparietal; pmx, premaxilla ; qj, quadratojugal ; spo, supraorbital lateral-line canal; sq, squamosal; spt, supratemporal; tm, temporal lateral-line canal; tab, tabu- lare; so, supraoccipital cross-commissure of lateral-line canal. IOO THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. gate, and with the quadratojugal forms the exterior border of the cranium. No teeth are observed on the maxilla. The postfrontal and the postorbital form the posterior border of the orbit, and between them inclose the anterior extension of the squamosal. The supratemporal is pointed anteriorly and has the usual relations of that element, joining the postorbital, the postfrontal, the parietal, the tabulare, and the squamosal. The tabulare is larger than the postparietal and is acuminate, the point being inclosed by the squamosal and the supratemporal. The jugal widens fan-shaped posteriorly. It forms a portion of the border of the orbit. The supratemporal, as usual, forms the quadrate angle of the skull. In front of it lies the elongate quadratojugal. Dendrerpeton is not well enough known for an exact comparison with Erpeto- saurus obtusus, but Cope separated the latter from Dendrerpeton on the position of the orbits and the broadly rounded muzzle. This species differs from the other species of the genus in the form of the cranium, as well as in the characters which separate it from Dendrerpeton. Measurements of the Type of Erpetosaurus obtusus Cope. mm. mm. Median length of the skull 48 Width of orbit 7 Width of skull at posterior border 56 Interorbital space 15 Width across orbits 37 Diameter of nostril 1.5 Length of orbit 12.5 Diameter of pineal foramen 1.75 Two other specimens, Nos. 8602 G and 8608 G of the American Museum, are associated in this species. Erpetosaurus tabulatus Cope. Cope, Proc. Amer. Phil. Soc, xvi, p. 577 (Tudilanus tabulatus). Moodie, Jour. Geol., 17, p. 52, figs. 8, 9, 1909. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, pp. 347, 351, pi. 59, fig. 2; pi. 62, fig. 2, 1909. Type: Specimen in the Zoological Collection of Columbia University. Horizon and locality: Linton, Ohio, Coal Measures. (Plate 25, fig. 2.) The species is known from a single well-preserved skull and its obverse in the collection of Columbia University in New York City. I am indebted to Dr. Bash- ford Dean for the privilege of studying this interesting form. It is from the Linton deposits of Ohio. The remains include a nearly complete cranium and a complete clavicle of the right side. The species agrees in all essential respects with the char- acters of the genus Erpetosaurus, presenting a broad, flat head and a triangular clavicle. The cranium is wider than long, the muzzle broadly rounded. The orbits are wide ovals, and their posterior borders fall little behind the transverse line dividing the skull equally. The interorbital width equals the long diameter of the orbit. The posterior outline of the cranium is truncate in a straight transverse line between the prominent tabulare angles. The composition of the cranium is different from that of any other species of Erpetosaurus in the large size of the tabulare and the fact that the supratemporal is excluded from the parietal by the extension of the postorbitals and the tabulare. This may be a generic character and entitles the species to be placed in a new genus, but it will be retained here until more of the anatomy of the PLATE 13 Dendrerpeton mveni Dawson. Above: Skull, mandible, and bones of anterior limbs. Below: Pos- terior limb, pelvis, and bony scales. Nearly natural size. Erect tree, South Joggins, Coal formation, Nova Scotia. Photograph by Dawson, published through the courtesy of Dr. Arthur Willey. Original specimen in the Peter Redpath Museum of McGill University. THE MICROSAURIAN FAMILY TUDITANID^E. IOI species is known. The elements of the anterior part of the skull are not preserved, but they are indicated by the broken lines in the drawing (fig. 22, G) . The nostrils are, however, clearly indicated as bosses of shale. There is a mere fragment of the nasal preserved posterior to the crack indicated by the transverse line in the drawing. The frontal is elongate as in other species of the genus and forms the inner border of the orbit. The parietal, as usual, is one of the larger bones of the skull roof and the pineal foramen is inclosed in the median suture by the two parietal elements. The pineal opening lies in the posterior half of the parietal. The postparietal is almost square, being slightly elongate transversely, uniting laterally with the tabulare, with which it forms the truncate table of the skull. The suture separating the tabulare from the supratemporal is clearly distinct. Although such a position for the supra- temporal is unusual it is not unique, since the same character has been observed in Diceratosaurus Icevis Moodie, described elsewhere (p. 120) in this paper. The post- frontal is rather small and it, together with the postorbital, forms the posterior boundary of the orbit. The postorbital is truncate posteriorly and joins the tabu- lare broadly. The supratemporal lies posterior to the postorbital and jugal and borders the quadratojugal, which is an unusual condition, but what significance the condition has remains to be determined. Posterior to the supratemporal lies the squamosal, which forms the quadrate angle of the skull. The quadratojugal is a small element and forms part of the lateral boundary of the skull. The jugal is a large element and forms the entire lateral border of the orbit. There are no teeth preserved on the fragment of the maxilla, but there are some impressions farther forward which resemble the pleurodont denticles of the modern Amphibia. The sculpture of the surface of the cranial bones consists of parallel ridges which are separated by grooves equal to them in width. The ridges radiate inward on the squamosals and frontals and outward on the supratemporals. They are somewhat interrupted on the other skull elements. The right clavicle is ornamented with a sculpture of similar radiating grooves and ridges. Cope described an atlas in connection with this skull, but I do not find it. The slender impressions to the right of the clavicle may possibly represent ribs. They are gently curved and truncate at the inner end. A nearly complete system of lateral -line canals has been detected on this skull. The canals preserved are : the temporal, the jugal, the infraorbital, the occipital cross- commissure, and the supraorbital. These terms were used for the first time for the Amphibia by the writer (458) in a discussion of the organs and their significance in the correlation of the skull elements. The occipital cross-commissure in the present skull is represented by a row of elongate pits, such as Andrews (8) has described for Ceraterpeton galvani Huxley from the Coal Measures of England. The cross- commissure is contained within the tabulare. The jugal and temporal canals form a complete ring, much as the same canals do in Trematosaurus. The supratemporal in Erpetosaurus tabulatus Cope is excluded from the parietal by the extension of the tabulare and the postorbital, and it is to be noticed that the temporal canal has a changed position to correspond with the changed condition of the squamosal. This is of considerable interest in connection with the correlation of the supratemporal 102 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. in fishes and amphibians. This subject has been fully treated in another place (458) and it will only be necessary to state here that on the basis of the lateral-line canals and their arrangement in fishes and the Amphibia the true correlation of the supra- temporal elements in amphibians and fishes has been made. The temporal canal in the present specimen has, apparently, an indication of a connection with the supraorbital canal, but of this I am not sure. The jugal canal occurs on the supra- temporal and quadratojugal, and it joins the infraorbital on the jugal. The infra- orbital is indicated by a short portion a few millimeters long under the orbit and the remainder, i.e., its connection with the jugal canal, is restored (fig. 22, G). There is, nothing unusual to be observed in that portion of the infraorbital canal which is preserved. The supraorbital canal is indicated by a curved, broad, shallow groove on the inner side of each orbit. As stated above, there seems to be a connection between this canal and the temporal, but I am not sure. The primitive conditions shown in the lateral-line canals in Erpetosaurus tabulatus Cope are the presence of the occipital cross-commissure and the ring-like formation of the temporal and jugal canals, which is too clearly indicated to be overlooked. Measurements of the Type of Erpetosaurus tabulatus Cope. mm. mm. Median length of skull 29 Diameter of nostril — I Width of skull, posterior border, estimated 37 Diameter of pineal foramen — 1 Width between tabulare angles 18 Length of right clavicle 13 Length of orbit 8 Width of right clavicle S-5 Width of orbit 6.5 MANDIBLE PROVISIONALLY ASSOCIATED WITH ERPETOSAURUS TABULATUS COPE. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, pp. 351-352, pi. lix, fig. 2; pi. lxiv, fig. 3, 1909. This specimen is preserved almost completely on a slab of soft coal. It is im- possible to determine positively to what microsaurian species the mandible be- longs, but it may, for the present, be associated with Erpetosaurus tabulatus Cope on account of its size and the character of its sculpture. This is the first and almost the only known example of a mandible of an American microsaurian. The form of the jaw is perfectly preserved, although the condition of the articular surface can not be determined. The proportions of the mandible , as may be j udged from the table of measurements , are rather stout and the teeth are strong and numerous. There are evidences of 19 teeth preserved. The sutures separating the articular (art) , angular (ang) , surangular (5a) , coronoid (cor) , and the dentary (d) (fig. 22, F) are clear for at least the greater part of their length and they may be easily restored for the remainder of their course. The surangular is thus seen to rival the dentary in size and on it occurs the peculiar sculpturing which approximates so closely that on the skull of Erpetosaurus tabu- latus Cope. The presence of the long anterior tooth is strikingly characteristic of many Microsauria. It is well developed in Sauropleura longidentata Moodie and Sauropleura (Anisodexis) enchodus Cope. It is also present in well-developed form in the later labyrinthodonts. The teeth are all, with the exception of the fourth from the anterior end, rather short, curved, and sharply pointed, with an indication THE MICROSAURIAN FAMILY TUDITANIOE. 103 of longitudinal fluting. The arrangement of the mandibular elements recalls in a striking way the mandible of Eryops megacephalus Cope as figured by Branson (49). Measurements of the Mandible Provisionally Associated with Erpetosaurus tabulatus Cope. (No. 8542 G, American Museum of Natural History.) mm. mm. Length of mandible 32 Length of one of the posterior teeth 1.25 Pi 1 terior width across surangular 6 Width across base of same tooth 50 Width of dentaiy 3 Length of long anterior tooth 2 Width of jaw at tip 1.5 Width of same tooth at base 75 Another specimen of this same species is 8550 G, of the American Museum. Linton, Ohio, Coal Measures. PALATE OF ERPETOSAURUS TABULATUS COPE. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, pp. 352-354, pi. lxi, fig. 2, 1909. The specimen is half a cranium with its impression. It is referred to Erpeto- saurus on the basis of the sculpturing of the mandible and the posterior table of the skull. On the surangular there is seen the rugosity which is common to other mem- bers of the genus. The characters presented are those of Erpetosaurus tabulatus Cope, but the reference is rather uncertain. The specimen is unique among the known American material in showing the structure of the palate. Jaekel (347) has figured and described the palate of Diceratosa urus puuctolineatus Cope from the Linton beds. The present palate differs from that species only in the enlarged ectopterygoid and the smaller palatine. The parasphenoid, in the present form, does not differ from that element in other Paleozoic Amphibia. Its form is slender, arising from an enlarged base and sepa- rating the pterygoids by its own width. The exoccipitals are probably represented in the specimen and they have been indicated in the drawing (fig. 22 C). They are rather large and extend some distance under the base of the skull to unite anteri- orly with the pterygoids, a very unusual arrangement. The pterygoids are elongate elements and are bounded anteriorly by the vomer and laterally by the ectoptery- goid. The vomer shows no evidence of being toothed, although it may have been so anteriorly. The same may also be said for the palatines. The relations of the ecto- pterygoids are rather unusual for the Amphibia, especially in the posterior exten- sion of the element. The bone lies all along the side of the pterygoid and anteriorly projects forward between the pterygoid and the palatine. In this unusual posterior projection the ectopterygoid has almost obliterated the infratemporal foramen, which possibly may be still represented by the triangular space between the bases of the pterygoid and the ectopterygoid. The anterior palatine foramen (internal nares) lies between the anterior ends of the palatine and the vomer, its usual rela- tions in the labyrinthodonts. The foramen may be recognized as the rounded depression slightly anterior to the palatine. The mandible is rather heavy and is coarsely sculptured with radiating grooves and ridges. The character of the teeth can not be determined, save to say that they were present. The posterior end of the mandible projects somewhat beyond the quadrate angle of the skull. 104 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The interest in the present specimen is heightened by the light it throws on the characters for the separation of the Amphibia and Reptilia. The wide separation of the pterygoids by the parasphenoid is an amphibian character of undoubted value. The reduction of the parasphenoid in this specimen is noteworthy. Measurements of the Skull of Erpetosaurus tadulatus Cope. (No. 8607 G, American Museum of Natural History.) mm. mm. Length of skull in median line 45 Width of pterygoid 5 Estimated posterior width of skull 50 Length of ectopterygoid 17 Estimated width of parasphenoid 6 Posterior width of mandible 12 Erpetosaurus minutus Moodie. Moodie, Proc. U. S. Nat. Mus., 37, pp. 21-23, pi. 8, fig. 1, 1909. Type: Specimen No. 4545, U. S. National Museum. Horizon and locality: Cannelton slates, Pennsylvania (Upper Freeport). (Plate 20 D.) The specimen on which the species is based is composed of the greater portion of a small skull preserved in the hard shale from Cannelton, Pennsylvania, and was collected by Mr. R. D. Lacoe, of Pittston, Pennsylvania. The characters of the specimen had not previously been determined, since the museum label and number had partially obscured the snout of the skull. The skull is very small, but has the form assumed by other members of the genus. The specimen may belong to a young individual, but even though it does, it is, nevertheless, quite distinct from the other species of Erpetosaurus. At first sight the specimen looks like a broken scute of some larger form. Close inspection, however, revealed the two impressions repre- senting the orbits, and a Zeiss binocular revealed the characters. The large size and anterior position of the orbits, the character of the sculpturing, the presence of the posterior table of the skull, as in Erpetosaurus (Tuditanus) tabulatus Cope, are the characters on which a specific diagnosis is possible. The specific characters which distinguish this form from E. tabulatus Cope, its nearest ally, are the slight development of the posterior table, the more delicate form of the sculpturing, the more posterior position of the orbits, and the varying shape assumed by the parie- tals in the two species. Any one of these characters would be valid as a specific character. The pineal eye is indistinct, but is observed to lie in the broken tract in the median line of the skull, in the middle of the portion posterior to the orbits. The interorbital space is equal to the width of the orbit. The orbits themselves are slightly oval and not round, as in the case of E. tabulatus Cope. The skull elements are sculptured with radiating grooves and ridges, and on the postparietals and tabulare the grooves take the form of pits in a row, which undoubt- edly represent the occipital cross-commissure of the lateral-line system first observed in a microsaurian by Andrews (8) in the skull of Ceraterpeton galvani Huxley. The supraorbital canal is represented by a slight elongate depression observable over each orbit and extending, in one case, for about 5 mm. The presence of the circular arrangement of the lateral-line canals in the jugal region is suggested by a depres- sion on the posterior edge of the squamosal. THE MICROSAURIAN FAMILY TUDITANIDiE. IO5 The portion of the skull anterior to the orbits is wanting, curiously enough, just as in Erpetosaurus tabula t us Cope. In the remainder of the skull, the post-parietals, the tabulare, the parietals, the supratemporal, and a portion of the right frontal can be detected, although the boundaries of but three can be accurately denned. The depression bounding the anterior outline of the skull is taken to be the impress of the mandible, in which case this structure would be of some depth, as in the case of the mandible associated with E. tabulates Cope, described below. This specimen is of interest in respect to the presence of the lateral-line canals, its small size, and its generic identity with forms from Ohio. There is still another form known from the Cannelton slates, described below as Erpetosaurus (Tuditanus) sculpt His Moodie. Measurements of the Type of Erpetosaurus minutus Moodie. mm. mm. Length of skull 18 Length of orbit 4.5 Posterior width of skull 17 Width of orbit 3.5 Width of skull across orbits 14 Interorbital width 3.5 Erpetosaurus sculptilis Moodie. Moodie, Jour. Geol., 17, No. 1, p. 6i, figs, n, 12, 1909 {Tuditanus sculptilis). Moodie, Bull. Am. Mus. Nat. Hist., xxvi, p. 347, 1909 (Erpetosaurus sculptilis). Moodie, Proc. U. S. Nat. Mus., 37, p. 22, 1909 (describes pectoral girdle). Type: Specimen No. 12,315, University of Chicago, Walker Museum. Horizon and locality: Cannelton slates, Pennsylvania (Upper Freeport). (Plate 18.) There is preserved in the collections of Walker Museum of the University of Chicago a small amphibian skull pressed flat on a slab of slate from Cannelton, Pennsylvania. It formed part of the Hall collection acquired by the University of Chicago in 1908. The specimen presents only a portion of the skull and fragmentary pectoral plates. The skull is wider than long and the muzzle is broadly rounded. The orbits are narrow ovals and their posterior border falls on the transverse line dividing the skull equally. The interorbital width is slightly greater than the width of the orbits and about equal to their length. The posterior outline of the skull is somewhat truncate, as in E. tabulates Cope and other species of the genus. The distal extremities of the quadrates do not project as far backward as do the supraoccipitals. The skull roof is formed of the regular elements, except that a quadrate seems to be indicated by a scale of bone on the posterior angle. The nostrils are oval and the pineal opening is small. The premaxillae are apparently relatively large elements, though their bounda- ries are not definite. The nasal is of an oblong shape and borders the frontal ante- riorly. The frontal forms the whole of the interior border of the orbit and borders the parietal broadly behind. The parietal is a large element and the pineal foramen is inclosed in the median suture about midway of the parietals. The postparietal is wider than long and with the tabulare forms the greater part of the posterior border of the skull. The prefrontal (plate 18, fig. 1) apparently forms the entire anterior border of the orbit and sends an acuminate projection to the side of it. The IOG THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. maxilla is excluded from the orbit and is an elongate element with sharp conical teeth, of which there are 4 preserved. These measure about 1 mm. in length. The jugal lies along the lateral border of the orbit and it is acuminate both anteriorly and posteriorly. It borders the supratemporal broadly. The postfrontal forms the greater part of the posterior boundary of the orbit. It is triangular and acuminate behind, and is bordered broadly by the parietal and supratemporal. The supratem- poral is also triangular and it borders the parietal broadly. The squamosal is evi- dently the largest element in the skull and on its posterior corner there is a flake of bone which may represent the quadrate, though this is by no means certain. The quadrate has not been detected in any of the Carboniferous Microsauria so far studied. The tabulare is an elongate element in the transverse line of the skull. Its entire boundary is uncertain, though part of the sutures are present. The quadratojugal is elongate and lies posterior to the maxilla and with that element forms the lateral boundary of the skull. The canals of the lateral-line system have not been detected on the skull. The sculpturing of the cranial elements consists of grooves and ridges which radiate from a center. They are more prominent on the parietals than elsewhere, although the other skull elements present a strong sculpturing. There are also preserved on the slab of slate, about 10 mm. posterior to the skull, fragments of the pectoral plates, probably representing the clavicles and the inter- clavicle. They are so badly fractured that their form can not be determined. No limbs or vertebrae have been observed. Measurements of the Type of Erpetosaurus sculptilis Moodie. mm. mm. Length of skull in median line 20 Interorbital space 4 Width of skull at posterior border (estimated) 24 Diameter of nostril — 1 Diameter of orbit 3 Pineal foramen, diameter 50 Length of orbit 4 Pectoral Girdle Provisionally Associated with Erpetosaurus sculptilis Moodie. Moodie, Proc. U. S. Nat. Mus., 37, p. 22, 1909. The present specimen is preserved on a block of slate from Cannelton, Pennsyl- vania. It is associated with the previously described Erpetosaurus sculptilis Moodie on account of its size, the geological and geographical distribution, and the charac- ter of the sculpture. It may pertain to an unknown species. Other remains besides the 3 elements of the pectoral girdle are preserved on the block of slate, but they are, for the most part, too imperfectly preserved for recognition. Some of them are phalanges, and I believe I detect a scapula in the rounded curved plate lying near the right clavicle. The 3 pectoral elements, the interclavicle and the 2 clavicles, are preserved intact, with the ventral surface uppermost. The specimen is particularly important in that it furnishes further evidence of the simplicity of the microsaurian pectoral girdle, which Jaekel regarded (347) as being extremely complex, in one species at least, Diceratosaurus punctolineatus Cope. The 3 elements are broken, but either the elements or their impressions are pres- ent, so that identification is possible. The elements are sculptured with radiat- 1 and 2. Photograph of the specimen of Amphibamus grandiceps Cope, from the Mazon Creek shales. X 1.5. Original in possession of Mr. L. E. Daniels, Rolling Prairie, Indiana. 3. Specimen of Sauropleiira (Coloslnis) saitrllata Newberry, from the Linton Coal Measures. The species was the first known of the Ohio Coal Measures Amphibia; at first ascribed by Newberry to the fishes, but later correctly identified by Cope. X 1. Original in American Museum of Natural History. 4. The type of Diceratosaurus (CeraterpetoiA punctolineafus Cope, from the Linton Coal Measu res. X 1. Original in American Museum of Natural History. THE MICROSAURIAN FAMILY TUDITANID^E. 107 ing grooves and ridges, as in so many of the Microsauria. The interclavicle is spatulate and bears a general resemblance to the same element of Metoposaurus fraasi Lucas from the Triassic (383) of Arizona. The clavicles are triangular, with rounded angles and the hypothenuse on the interior border. Measurements of Pectoral Girdle of Specimen No. 4539, U. S. National Museum. mm. mm. Width across the entire girdle 17 Length of clavicle 11 Length of interclavicle 15 Width of clavicle, maximum 6 Width of interclavicle 10 Erpetosaurus acutirostris Moodie. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, pp. 349-351, pi. lxi, fig. 1, 1909. Type: Specimen No. 8598 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The present species adds another form to the diversity of structure presented by the Carboniferous Microsauria. It is closely allied to Erpetosaurus (Tuditanus) obtu- sus Cope, from the same beds (Linton, Ohio), but differs from it especially in the position and shape of the orbits and the acute form of the skull. Other characters which amount almost to generic significance are found in the posterior prolongation of the frontal and in the triangular form of the skull. Only the skull of the animal is preserved. The character which is common to all members of the genus Erpeto- saurus, the cranial rugosity, is present in this species on the squamosal and supra- temporal. This character alone would not, however, suffice to separate the form generically, but the general morphology and arrangement of the cranial elements is such that reference to any other genus save Erpetosaurus would not be possible. The skull of E. acutirostris takes the form of a rounded triangle. Its base is some 50 mm. in extent, and this width gradually narrows to 31 mm. across the orbits and still more towards the snout. The form of the skull is not widely different from that of the type species, E. radiatus Cope, but the differences are sufficiently apparent. Nearly all the elements of the cranium can be detected (fig. 22, D). The bony portion of the cranium has nearly all been lost, leaving only the impression; and the matrix in which the skull was embedded has been forced up into the sutures between the cranial elements, thus forming ragged ridges where the bones of the skull joined. The position of the nostrils can not be determined accurately. The orbits are placed well forward, a character common to several species of the genus. The inter- orbital space is equal to the long diameter of the eye. The orbits are sep- arated by narrow prolongations of the postfrontals and by the anterior por- tion of the frontals. The frontals are remarkable in their great back- ward extension. In E. obtusus the frontals are nearly confined to the inter- orbital space. The parietals, which, on the median suture, inclose the parietal foramen, lie well posterior, and the parietals and the tabulare are small. A portion of the sculpturing of these elements has been preserved and 108 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. it is seen to be made up of pits and elevations much as we find in the skull of Saurerpeton latithorax Cope. The left squamosal also shows sculpturing, which here tends to take the form of grooves and ridges, and also of pits and elevations. It is quite probable that the anterior portion of the skull was ornamented with grooves and ridges and undoubtedly the lateral-line canals were well developed. The post- frontals and the postorbitals are both large and elongated. The postorbital is especially large. The supratemporal apparently separates the tabulare and the squamosal in their posterior extremities. The squamosal projects posteriorly to the tabulare and apparently even goes beyond the limits of the exoccipital. The out- lines of the jugal are fairly definite, as are also the limits of the maxilla. Measurements of the Type of the Skull of Erpetosaurus acutirostris Moodie. mm. mm. Length of skull 50 Width across orbits 31 Interorbital space 9 Posterior width of skull 50 Width of orbit 7 Diameter of pineal foramen I Length of orbit 10 Length from tip of snout to posterior angle of skull. . 65 An additional specimen shows that the skull, of which the anterior half is pre- served, is practically of the same size as the type and shows much the same characters, though more extensively. The sculpture of the squamosal region is not confined to that portion of the skull, but extends throughout the cranial elements, apparently including the premaxillae. The sutures, which are clearly distinct, are of the same type as has been described for the type specimen. Perhaps one of the most interesting characters discovered on the present specimen is that of the greater part of the left supraorbital lateral- line canal, which is exhibited as a rather deep and broad canal running in a slight curve across the lower edges of the postorbital and the parietals and partly cutting the jugal. On the left of the fossil, as it is preserved, there is an indistinct impression of the clavicle, with the sculpture in radiating lines from the apex as a center, such as have been found in other species of the Tuditanidae. Measurements of Additional Specimen. (No. 8607, American Museum of Natural History.) mm. mm. Length of skull as preserved 40 Length of clavicle 18 Anterior width across orbital region 22 Greatest width of clavicle 9 Greatest width of the skull 33 Still a third specimen of this species is possibly represented by a nearly complete skull of a small individual which exposes the mandible and the ventral portion of the skull. The remains are crushed flat, though not at all distorted. It is quite pos- sible that the present specimen represents a distinct species, but only a small portion of the dorsum is present and the shape of the cranium is indistinct, so it is retained in this species. The portion of the skull shows the sculpture to be quite similar to that of Erpetosaurus acutirostris, so far as the species is known; and the skull appar- ently tapers to a point anteriorly. It may be a dwarfed or immature form. The sculpturing of the jaw is such as we would expect of this species. THE MICROSAURIAN FAMILY TUDITANIDiE. 109 The specimen is about half the size of the type. The palate of the skull is well preserved and is extremely interesting. The sutures separating the various palatal elements are not distinct. The parasphenoid is especially large and the exoccipitals arc partly ossified, if we may judge by the projecting condyles. Anteriorly the parasphenoid contracts and then expands and on each side of the expanded part lie fragments of the palatines. To the right of the posterior end of the parasphenoid lies a portion of the dorsal element showing the cranial sculpture. The left mandible is somewhat displaced to the right of the skull, and crushed and weathered to such an extent that the sutures are entirely obliterated. There are 3 teeth, with indications of others. They are typically pleurodont and sharp and slender. The mandible tapers somewhat anteriorly and at the tip bears an elongate enlarged tooth. Measurements of the Third Specimen of Erpetosaurus acutirostris Moodie in the American Museum of Natural History. mm. mm. Length of skull, as preserved 25 Length of mandible 25 Posterior width of skull 15 Anterior width of mandible 2 Anterior width of skull 10 Length of large tooth I Width across occipital condyles 4 Erpetosaurus tuberculatus Moodie. Moodie, Bull. Amer. Mus. Nat. Hist., xxvi, pp. 348-349, pi. lviii, 1909. Type: Specimen Nos. 8693 G and 8610 G, American Museum of Natural His- tory. Horizon and locality: Linton, Ohio, Coal Measures. This species is based on a fragmentary cranium (plate 26, fig. 1) consisting of the posterior part of the right side of the skull. Its association in the genus is solely on the character of the sculpturing of the cranial elements. It is most closely related, in the characters preserved, to the form described by Cope as Tuditanus radiatus, from which it differs especially in the character of the sculpture and in the position of the orbits, as well as the arrangement and size of the various cranial elements, so far as these elements can be detected in the present specimen. In Erpetosaurus radi- atus the skull is sculptured by radiating grooves and ridges which did not arise from a definite center. In E. tuberculatus this center of radiation is marked by an eleva- tion or tubercle on each cranial element exposed, from which the grooves and ridges radiate outward. These tubercles have an elevation of 4 mm. above the cranial element proper. The orbit is located near the median line of the skull, so far as can be determined. In E. radiatus Cope the orbits are located well forward. In that species also the postparietal is smaller than in the present species and the squamosal is longer and more slender. (Plate 25, fig. 1.) The fragment of a skull on which the above comparison has been made consists of the right postparietal, a portion of the tabulare, the parietal, the frontal, and a portion of the squamosal. The other elements are not clear. The elements in the median line are elongate, as in Erpetosaurus radiatus. The pineal foramen is located well back on the median line and lies posterior to two-thirds of the length of the HO THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. parietals. The sutures separating the frontal and parietal elements from each other in the median line are of the zigzag form so characteristic of the labyrinthodonts. Measurements of the Type of Erpetosaurus tuberculatus Moodie. mm. mm. Length of portion of skull preserved 52 Maximum width of parietal n Width across tabulare (estimated) 60 Length of frontal 22 Length of parietal 15 Width of frontal 12 Genus ODONTERPETON Moodie. Moodie, Proc. U. S. Nat. Mus., 37, p. 19, 1909. Type: Odonterpeton triangularis Moodie. The generic characters may be found in the triangular shape of the skull, the large size of the teeth, the shape of the vertebrae, the small size of the orbits, and their anterior position as shown in the type specimen (fig. 22, E). The name of the genus is derived from the remarkable size of the teeth as compared with the size of the skull. Odonterpeton triangularis Moodie. Moodie, Proc. U. S. Nat. Mus., 37, p. 19, pi. 6, fig. 3, 1909. Type: Specimen No. 4465, U. S. National Museum. Horizon and locality: Linton, Ohio, Coal Measures. The specimen described under the above name is a representative of the small- est species of the Microsauria so far described from North America. Orthocosta microscopica Fritsch, from the Carboniferous of Bohemia, is a rival of the present form in size, but the form described by Fritsch is an entirely different animal and was formerly included among the so-called Aistopoda, which are regarded by the writer as merely specialized microsaurians. The present form shows clear affinities with the Microsauria. As may be seen by referring to the list of measurements, the skull of this form measures only 6.5 mm. in length. The form may possibly be larval, though I do not think so, if I may judge from the well-developed condition of the skull bones and the complete ossification of the vertebras. The sides of the skull are equal and the occiput is a straight table, so that the skull forms almost an exact equilateral triangle. The orbits are very small and are placed well forward. The interorbital space is four times that of the diameter of the orbit, a very unusual character and in itself worthy of ranking as a generic character. The median suture of the skull is zigzag and incloses the minute parietal foramen near the posterior end of the skull. The relations of the elements of the skull, with the exceptions of those of the frontals and parietals, can not be determined with accuracy, although there are here and there indications of sutures. The characters of the cranial elements, so far as they can be determined, are those of the family Tuditanidae, and the form may, for the present, be regarded as a member of that group. The teeth are very long, slender, and sharp, and are placed close together. There is no indication of fluting on the teeth. They are slightly curved inward. There are 13 vertebrae present. The centra are hour-glass shaped, and are apparently phyllospondylous, with the notochord largely persistent. The vertebral THE MICROSAURIAN FAMILY TUIHTANID^. Ill centra are unusually long and slender, with the ends rounded. The humerus of the right side is preserved. It is a long, slender bone with expanded extremities. There is no evidence of abdominal armature nor of ribs (fig. 22, E). The discovery of this form in the Linton deposits is of considerable interest in that it indicates a wide range in size and character of the fauna of the time. The forms now known from the Linton beds range from Odonterpeton, which possibly had a total length of 2 inches in life, to the form designated Macrerpeton huxleyi Cope, with a skull of at least 8 inches in length and whose body may have attained a length of some feet. The large rib described below undoubtedly indicates a large form of the ancient Amphibia from Linton, as do also the vertebrae described by Marsh in 1863 from Nova Scotia. Measurements of the Type. mm. mm. Length of animal, as preserved 18 Length of tooth 0.25 Length of skull 6.5 Length of vertebra 1.45 Posterior width of skull 5.5 Width of vertebra 35 Length of side of skull 6.5 Length of humerus 2.25 Diameter of orbit 65 Distal width of humerus 35 Intcrorbital width 2 CHAPTER XV. THE MICROSAURIAN FAMILY STEGOPID^E, FROM THE COAL MEASURES OF OHIO. Family STEGOPIDjE Moodie, 1909. Moodie, Jour. Geol., xvii, No. I, 79, 1909. The chief family characters are the large lacrimal, unknown in other species of Coal Measures Amphibia, the central position of the orbits, the general form of the skull, and the peculiar, short, divaricate horns from the squamosal. If an inter- temporal element is present in the skull, which is suggested as a possibility, the family is further distinct. The type species is Stegops divaricata Cope from the Linton, Ohio, Coal Measures. The group seems to be distinct and has no immediate allies, being confined to the American Coal Measures. Genus STEGOPS Moodie, 1909. Moodie, Jour. Geol., xvii, 79, 1909. Type: Stegops divaricata Cope. This genus has been erected for the reception of the peculiar form described by Cope as Ceraterpeton divaricatum, but there are good reasons why the form can not be retained in the genus. The position of the orbits in Stegops (plate 25, fig. 3) is dif- ferent from Ceraterpeton and the muzzle is rounded, not truncate as in the latter form. The horns are of a different type and there is no indication of the tabulare protuberance which is present in Ceraterpeton. The sculpturing of the cranial elements is also distinctive in the present form, consisting of radiating grooves and ridges; the cranium of Ceraterpeton appears to be but slightly sculptured. There is no lateral projection from the border of the skull in Stegops, as there is in the other genus. The structure of the skull of Ceraterpeton is practically unknown, except in a very general way, although Andrews (8) was able to make out some of the elements and to trace the lateral-line canals. A structural comparison is thus impossible, but on the basis of form alone there are good generic distinctions. The present genus is apparently distinct from other genera in the presence of an inter- temporal, but additional material will be required before a satisfactory determina- tion is possible. The genus Diceratosaurus of Jaekel (347) is distinct in the arrange- ment of the elements of the cranium, the general form of the skull, and in the two known species of Diceratosaurus the orbits are located well anteriorly, but in Stegops they are in the median transverse line of the cranium. The genus Stegops is distinct from Eoserpeton in the smaller size of the prosquamosal, in the broadly rounded muzzle, in the larger and more posteriorly placed orbits, and in the pres- ence of an intertemporal bone, or at least in the elongate character of the post- orbital if the intertemporal is not present. The species on which the genus Eoserpe- ton is based was first described by Cope as Ceraterpeton tenuicorne. The form is THE MICROSAURIAN FAMILY STEGOPIM;. 113 quite distinct, in spite of Jaekel's protestations to the contrary. The genus Stegops stands alone among the Carboniferous Amphibia of North America, so far as I am aware, in the possession of a well-defined lacrimal of the labyrinthodont type. Stegops divaricata Cope. Cope, Proc. Am. Phil. Soc., xxn, p. 406, 1885 (Keraterpeton divaricatum) . Moodie, Jour. Geol., xvn, No. 1, p. 79, fig. 22, 1909 {Stegops). The Type: Specimen No. 2559 G, American Museum of Natural History obverse of this is No. 12,311, Walker Museum, University of Chicago. Horizon and locality: Linton, Ohio, Coal Measures. The skull on which this species is based consists of the impressions on two slabs of coal, one belonging to the Newberry Collection of the American Museum of Natural History, No. 2559 G, and the other to the Gurley Collection of the Uni- versity of Chicago, No. 12,311. The slab be- longing to the University of Chicago contains the better-preserved remains, so that the descrip- tion is based largely on that portion (plate 25, fig. 3). Nearly all of the elements of the skull are determined with considerable certainty and many important characters in the morphology of the Microsauria are thus brought out. The skull is oval, elongate, truncate behind, and the quadrate angles project into sharp horns. The orbits are elongate ovals and their center lies in the median line which divides the skull transversely. The nostrils are elongate and have an oblique position. The pineal fora- men lies in the posterior third of the skull. Teeth are preserved on both maxilla? and pre- maxillae. They are simply sharp pleurodont denticles, and seem to have been fairly abun- dant. The bones have been completely carbon- ized and nothing of the original texture is preserved, although the details of the structure are beautifully preserved. (Plate 25, fig. 3.) The skull is somewhat triangular in its general form. The premaxilla lies on the anterior border of the cranium, and forms the median border of the nostril. The suture which separates the maxilla and the premaxilla is not evident, and it may not be correctly defined in the figure (fig. 23). The nasal is a very large element and is elongate. It unites with the premaxilla, the lacrimal, the pre- frontal, and the parietal. It is acuminate behind and the point is inclosed by the prefrontal and the parietal. The frontal is quite narrow and elongate, and does not border the orbit; its posterior boundary is not accurately represented. The radiations on the surface indicate the extent of the element. The parietals are remarkable in being smaller than the frontal and nasal. The pineal foramen is Fig. 23. — Skull elements of Stegops divaricata Cope, ft, frontal ; it, intertemporal ; j, jugal ; la, lacrimal; mx, maxilla; n, nasal; par, parietal; pf, prefrontal; po, postorbital; pof, postfrontal; pmx, premaxilla; sq, squamosal; spt, supratemporal ; qj, quadratojugal; tab, tabulare; pp, postparietal. X 1.3. 114 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. inclosed in the median suture in the anterior third of the parietals. The parietal has the usual relations of that element. The postparietal lies posterior to the parie- tal, but a portion of its bounding sutures are destroyed. The prefrontal forms the antero-interior border of the orbit and borders the postfrontal posteriorly, in an unusual manner. The lacrimal is a large element and is clearly separable from the other cranial elements. It, with the prefrontal, forms the anterior border of the orbit. The maxilla is very elongate and forms the larger part of the lateral border of the skull. Sharp-pointed teeth are present on it and they may have been pleurodont. The lateral border of the orbit probably received a portion of the maxilla. The postfrontal and the postorbital form the greater part of the poste- rior boundary of the orbit. The postorbital seems to be divided by a median suture which would indicate an intertemporal bone, but this is not certain. The appear- ance may be due to a fracture. The supratemporal is a large element bordering the parietal and lies in front of the tabulare. The squamosal is elongate to form the posterior horn -like extension of the skull. The tabulare is transversely elon- gate and has the usual relations. The jugal widens to a fan-shape backwards, and helps to form the lateral border of the orbit. Its lateral and anterior boundaries are not assured. The quadratojugal seems to lie as indicated, although the ante- rior part of the suture is not distinct. It is apparently an elongate element and with the maxilla forms the lateral border of the skull. The base of the skull as restored (fig. 23) is irregular and may have had a slightly different form. The genus Stegops is exceptional in the elongate character of the cranial ele- ments of the single species known. In this respect it recalls the species Dicer ato- saurus Icevis described below. The large size of the nasals, frontals, and lacrimals and the small size of the parietals are, so far as I am aware, unparalleled among the other Coal Measures Amphibia of North America. Measurements of the Type. mm. Median length of the skull 56 Interorbital space 16 Width across tips of horns 46 Length of the nostril 2 Width at base of horns 40 Diameter of the pineal foramen 1 Width across orbits 44 Length of the teeth 1.5 Diameter of the orbit 8 Length of the horn from base 7.5 Length of the orbit 15 Width of horn at base 4 MOODIE 1. Dorsum of skull of Diceratosaurns punctolineatus (Cope), from the Coal Measures of Linton, Ohio. Original in the Museum at Berlin University. X 2. After Jaekel. /o=postorbital; /r=frontal; ^o/=postfrontal; ;'=jugal; /a=laerimal; »/j-=maxilla; /».s="perisquamosal;" pp= post-parietal; ^«/:r=premaxilla. 2. Ventral surface of the skull of Diceratosaurns punctolineatus (Cope), from the Coal Measures of Linton, Ohio. X 2. After Jaekel. «=anterior palatine vacuity; *r=:exoccipital; /=jugal; »tr=maxilla;/r=transverse;/><7/=palatine;^=pterygoid;^r=prevomer;/.s=''perisquamosal;'' ^z'=posterior or suborbital palatine vacuity; /A=parasphenoid. 3. Pectoral girdle of Diceratosaurns punctolineatus (Cope), from the Coal Measures of Linton, Ohio. Original in the paleontological Museum at Berlin. X 2. After Jaekel. f//'=inner side of clavicle; ;'r=interclavicle; <7=clavicle (.lower side); fte=cleithrum. 4. (e ) Cervical or anterior dorsal vertebra of Diceratosaurns punctolineatus (Cope), from the Linton, Ohio, Coal Measures. rj<=c:uidal vertebra shown with its cap of dermal bone; dv—<\oxs&\ vertebra with ribs; .r-dcrmal plate <>n neural spine; A=humeius; «=ulna; r— radius. CHAPTER XVI. THE MICROSAURIAN FAMILY UROCORDYLID^E, FROM THE COAL MEASURES OF OHIO. Family UROCORDYLID^E Lydekker, 1890. Le of Eoserfe- ton tenuicorne. mm. Median length of skull 26 Width across squamosal enlargement.. . 30 Width across base of horns 25 Length from muzzle to tip of horn 40 Length of horn 10 Width of horn at base 4 Length of orbit 4 Width of orbit 3 Interorbital space 6 Nos. 4472 and 4473, U. S. National Museum, from Linton, Ohio, Coal Measures. mm. Median length of skull 16 Maximum width of skull 20 Length of horn from base 7 Length of orbit 3 Interorbital width 3.5 Length of vertebral column to sacrum . . 33 Length of femur 5.5 Length of tibia and fibula 3 Length of second digit (incomplete) 7 Length of metatarsal 1 .5 Length of clavicle 6 Width of clavicle 3.5 6 tff I C 1 / w H Fig. 25. — Restoration of skeleton of Eoserpeton lenuicorne Cope. X 1 . 5. Skull: pmx, premaxilla; n, nasal; fr, frontal; par, parietal; pf, prefrontal; nut, maxilla; pof, postfrontal; po, post- orbital; pp, postparietal; j, jugal; qj, quadratojugal; spt, supratemporal ; sq, squamosal; tab, tabulare. Skeleton: ic, interclavicle; cl, clavicle; sc, scapula; h, humerus; r, radius; u, ulna; c, carpus; sr, sacral rib (un- certain?); il, ilium; /, femur; fb, fibula; /, tibia; ts, tarsus. tals form a large oval space, so characteristic of many of the Carboniferous Micro- sauria, in the anterior third of which occurs the median parietal foramen. The postparietal is almost square, and forms part of the posterior boundary of the skull. The tabulare has the usual position and relations. The prefrontal is ill defined. The postfrontal is small and forms a slender rod on the postero-inner boundary of the orbit. The postorbital is small and its bounding suture with the postfrontal is THE MICROSAURIAN FAMILY UROCORDYLIOE. 125 indefinite. The jugal is only partially represented in the specimen, and that part forms the outer boundary of the orbit. The maxillary sutures are not defined. There are no evidences of teeth, since the skull is compressed dorso-ventrally. The quadratojugal is, apparently, a larger element than usual, with the usual rela- tions. The supratemporal lies in great part in front of the squamosal, but still has the normal relations of that element. The squamosal is the characteristic feature of the skull. It is very tumid at its base and projects into a long, slender, acumi- nate horn, the tumid portion being ornamented by radiating striae. Another specimen of this species presents the greater part of the skeleton. How- ever, very little can be added to our knowledge of the skull structure. It is barely possible that the second specimen may be distinct from the type. The horns are curved inward, but otherwise there is little or no difference. One of the most inter- esting and important features of the complete specimen is the unusual preservation of a leg, with impressions of 15 or more vertebras, and traces of curved ribs which are intercentral in position. The femur is slender and expanded at the ends, with the articular surfaces well formed. The tibia and fibula are mere rods of bone, although the tibia has slightly expanded extremities. There is no osseous tarsus. There are 5 digits in the foot; the second one is entire and contains 4 phalanges; the other digits are incomplete. The foot is remarkably long and slender, and is fully as long as the tarsal space plus the tibia, with the terminal phalanx clawed. There are impressions of 2 oval and elongate clavicles in the pectoral region. The outer end is not expanded as is usual, and the surface is ornamented with grooves and ridges which radiate from a common center. The entire remains measure scarcely 3 inches in length and it is to be doubted if the creature attained a length of more than 4 inches. It is probably a young form, but there are no evidences of external gills. The chevron armature is but poorly preserved, but so far as can be determined it is not different from that of other Microsauria, such as Amphibamus. CHAPTER XVII. THE M1CROSAUR1AN FAMILY AMPH1BAMID/E, FROM THE COAL MEASURES OF MAZON CREEK, ILLINOIS. Family AMPHIBAMID^ new family. Small, lizard-like, terrestrial or semi-aquatic, megacephalic microsaurians, known from 3 species. The family characters are the huge size of the head as compared to the body, the short, stumpy body with about 25 short dorsal verte- bras, a very short tail, phalanges clawed, pubis of calcified cartilage, sclerotic plates in the orbit to the number of 29 or 30 in each, ventral armature well developed. Teeth anisodont, sharp, conical, non-striate. Two genera are associated in this family: Amphibamus grandiceps Cope, known from three nearly complete skeletons; A. thoracatus Moodie, known from a single incomplete skeleton; Cephalerpeton ventriarmatum Moodie, anterior portion of body and skull. The species are all from the Mazon Creek shales and the family seems unrepresented elsewhere. It may be necessary to compare the Amphibamidae with the Hylonomidas when the latter group is better known, but in the light of our present knowledge the two families are distinct. The genera may be distinguished as follows: I. Size small, less than 3 inches in total length, skull with deeply incised tympanic notches (ear-slits) . .Amphibamus II. Size relatively large, body-length 6 inches or more, teeth distinctly anisodont, skull with nearly even posterior table, limbs very long, ventral armature highly developed Cephalerpeton Genus AMPHIBAMUS Cope, 1865. Cope, Proc. Phila. Acad. Nat. Sci., 1865, pp. 134-137. Geol. Surv. Ills., 11, pp. 135-141, pi. xxxii, 1 text-fig. Hay, Proc. Am. Phil. Soc, xxxix, p. 120, 1900. Moodie, Jour. Geol., xvn, p. 81, fig. 24, 1909. Type: Amphibamus grandiceps Cope. The publication of the type species of this genus began the researches of Pro- fessor Cope on the extinct Amphibia of North America, which he continued for so many years with such excellent results (105-177). The description was based on a single specimen (plate 3, fig. 7) belonging to Mr. Joseph Evans, of Morris, Illinois, who loaned it to Dr. Worthen for the Illinois Geological State Survey (107), in order that it might be described. The type has been destroyed by fire; so I am informed by Mr. L. E. Daniels, of Rolling Prairie, Indiana. There are two other known speci- mens of the species. One is in the collection of Mr. Daniels and the other No. 794, of Yale University Museum. This genus may be clearly separated from all the other microsaurians by char- acters which are peculiar to the form. Among these may be mentioned the posses- sion of sclerotic plates in the eyes; the large size of the orbits in comparison with the dimensions of the skull; the short, broad form of the body; the very short tail; the possession of a calcified cartilaginous pubis; clawed phalanges; presacrals 22. The character which places the genus distinctly in the Microsauria is the possession of long, slender, curved ribs, first detected on Mr. Daniels's specimen (plate 14, figs. 126 MOODIE PLATE 17 Type of Saurerpeton latithorax Cope. X 1.5. Original in U. S. National Museum. THE MICROSAURIAN FAMILY AMPHIBAMIOE. 127 1,2), by Dr. Hay (316). Its stegocephalian characters are evident in every partic- ular of its anatomy — the roofed skull, the arrangement of the cranial elements, the presence of a well-developed ventral armature, and the digital formula (4 for the hand and 5 for the foot). The genus A mph ibamus was regarded by Cope as a representative of a new order of vertebrates which he called (105) Xenorachia. He later ( 1 23) abandoned this, how- ever. Fritsch (251), Zittel (642), and others regarded Amphibamus as a branchio- saurian. The exact position of the form was uncertain until 1900, when Dr. Hay (316) described the long, curved ribs and suggested its place among the Microsauria. He, however (Cat. Foss. Vert., p. 410), made the mistake of including the branchio- saurian family Protritonidae, under Microsauria, thus confusing the subject further. The genus (462) has not the slightest relationship with the Branchiosauria. Amphibamus grandiceps Cope. Cope, Proc. Phila. Acad. Nat. Sci., pp. 134-137, 1865; Geol. Surv. Ills., 11, pp. 135-141, pi. xxxii, and 1 wood- cut, 1866. Hay, Proc. Am. Phil. Soc, xxxix, p. 120, 1900. Moodie, Jour. Geol., xvn, No. 1, p. 82, fig. 24, 1909. Moodie, Kan. Univ. Sci. Bull., vi, No. 2, pp. 343-349, pi. 1, figs. 1 and 2; pi. 5, fig. 3; pi. 7, fig. 1; pi. n, 12, 13, 1912. Type: Specimen has been destroyed. There is an excellent specimen (plate 4, figs. 5, 6), No. 794 (1234), in Yale University Museum, and another nearly as good in the possession of Mr. L. E. Daniels, of Rolling Prairie, Indiana. Horizon and locality: Mazon Creek shales, near Morris, Illinois. The form of the skull of Amphibamus grandiceps Cope is not unlike that of Tuditanus minimus Moodie (462) from the Linton, Ohio, beds, but it is less acumi- nate than in that form. The large size of the orbits is especially striking. The shape of the skull is triangular, with concavities in the posterior table which correspond to the ear-slits so characteristic of Metoposaurus (242) from the Keuper of Germany. The narrowed posterior table of the skull is truncate, as in several other genera of Microsauria, notably Tuditanus and Saurerpeton. In structure the skull differs but little from many of the other Carboniferous forms, but the arrangement of the elements of the skull is more regular than in other genera. The premaxillaries are very small elements in the anterior tip of the skull. They border the nares. The skull is rather peculiar among the Microsauria in the pos- session of a distinct lacrimal. I have detected this element in the cranium of Stegops divaricata Cope. As here defined the lacrimal is triangular, with its pos- terior border formed exclusively by the prefrontal. Its other relations are the nor- mal ones. The nasal is elongate, with the usual relations of that element. The frontal is slightly longer and broader than the nasal. It apparently forms a por- tion of the inner border of the orbit. The parietal foramen lies in the anterior fourth of the parietal, a rather unusual position for this structure. The parietals, as in so many of the Microsauria, together form the largest element of the skull and are roughly a triangular area in the postero-median portion of the skull. The post- parietal and the tabulare are clearly distinguishable and they have the usual rela- tions for those elements. The maxillary, jugal, and quadratojugal together form the greater part of the maxillary border. The postero-lateral angle of the skull is, 128 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. as usual, formed by the squamosal. The orbit is bounded posteriorly by the post- orbital and the postfrontal, which include in the angle between them the quad- rangular squamosal. The orbit is especially remarkable for its size as compared with the dimen- sions of the skull, being without a parallel among other known Microsauria. Around the border of the orbit in the specimen Cope studied (105) there were found 14 quadrangular plates which he called "superciliary plates." Hay (316) was inclined to regard them as sclerotic plates. In the Yale Museum specimen (plate 4, figs. 5, 6) there are 2a of these plates, and there seems to be no doubt that they are sclerotic ele- ments. In the restoration (fig. 26) 29 sclerotic plates are given, but there is no assurance that this number is the exact one. They may also have been slightly larger, but not as large as in Branchio- saurus. The vertebral column is pre- served nearly entire in the Daniels specimen and quite entire (478) in the Yale specimen. Cope, in his study of the type (105, 107), thought there could be no more than 13 presacrals, but the speci- men was poorly preserved and in- decisive on this point. Dr. Hay (316) was inclined to the opinion that there were less than 20. The Yale specimen shows 22 centra, which are elongate, hour-glass- shaped bodies, with the neural spine a long, low crest running the entire length of the centrum, Fig. 26. — Restoration of body outline and skeleton of .4 mphibamus grandiceps Cope, from Mazon Creek, Illinois, shales. Restora- tion is based on complete specimens of the species and on Cope's drawing. Form of body is indicated in one specimen, that in possession of Mr. Daniels. X 1.5. Skull: pmx, premaxilla; n, nasal ; fr, frontal; par, parietal; la, lac- rimal; pf, pref rental; pof, postfrontal; po, postorbital; pp, post- parietal; spl, supratemporal; mx, maxilla;j, jugal; qj, quadrato- jugal; sq, squamosal; tab, tabulare. Skeleton: ic, interclavicle; cl, clavicle; sc, scapula; h, humerus; r-u, radius, ulna; c, carpus; pu, pubis; il, ilium; /, femur; /, tibia; fb, fibula; Is, tarsus; x, ischium. with a median elevation, so that in lateral view the spine would be triangular in form. The body of the centrum is expanded laterally into a diapophysis which extends anteriorly. The posterior vertebrae, at least, had the notochord largely persistent. THE MICROSAURIAN FAMILY AMPHIliAMID^E. 129 t The osseous part of the vertebra seems to have been but a thin shell, and the struc- ture of the zygapophyses can not be determined. That they were dorsal in position is, however, evident from several vertebrae. The points of these structures project laterally. The tail is short and the caudal vertebrae weakly developed. There are distinct impressions of at least 12 pairs of ribs in the Daniels specimen. They are long, slender, and curved, and there is no definite assurance that there were as many ribs as are indicated (fig. 26) in the restoration (462). The ribs are inter- central (469) and probably occu- pied the full length of the verte- *"^ ~r bral column. There may have been as many as. indicated in the restoration. One of the most interesting features of the Yale specimen is the preservation of a small patch of skin, evidently from the back, lying to one side near the head, measuring 5 mm. in length by 3 mm. in width. The fragment shows the skin to be of tubercu- lated scales, 4 of which occupy the length of 1 mm. The scales are somewhat hexagonal, almost rounded, and were relatively quite thick. They lie in a close mosaic (fig. 27). The Yale specimen has, very well preserved, a portion of the ventral scutelke, of the throat, chest, and belly. The arrange- ment of the plates on the throat and chest is almost exactly the reverse of what Credner has described (190) for Bran- ch iosa urus amblystomus Cred. On the throat, in the present form, the chevron points anteriorly, and it is the anterior prolongation of the belly scutes with the postero- lateral projection of the gular scutes which form the chest and arm scutellation. The belly chevrons point anteriorly, as in Branchiosaurus, the rods formed by the scutes being straight and not curved as in Bra nch iosa urus . The entire ventral armature pre- served is displaced to the left of the animal and only the anterior portion is preserved. The pectoral girdle is only partially known. The scapula is crescent-shaped. The other elements are indicated only by fragments and nothing is known of their form. The arm elements are nearly all known. The humerus is slender and expanded at the ends, with its articular surfaces well developed. The separate radius and ulna are of approximately the same size and length. The carpus is unossified. The com- // ^J A ts^- % Fig. 27. — Skeleton of Amphibamus graiidiceps Cope. X i-4- carpus; cl, clavicle; cr, caudal rib; cv, caudal vertebra;/, femur; h, humerus; il, ilium; s, skin; or, orbit; r, radius; «, ulna; sc, scapula; sp, sclerotic plates; /, tibia and fibula; Is, tarsus; vs, ventral scutelte. Specimen No. 794, Yale University Museum. I30 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. plete phalangeal formula for the hand of Amphibamus is unknown. The third digit seems to have 4 elements. The formula 2-2-3-2 has been suggested (462). The pelvis is very satisfactorily known. The ilium is a long, slender, straight rod, with expanded ends. The ischium is shown on both sides of the vertebral column in the Yale specimen. Its form is almost identical with that of Paleohatteria longicaudata Credner, from the Rothliegenden of Saxony. The ischia are apparently approximate in the median line, though this character is somewhat obscured by the impression of the caudal vertebrae. Their relation with the ilium, other than that they were posterior to it, is uncertain. The pubis is, apparently, calcified cartilage. It is a squarish plate, somewhat corrugated, lying anterior to the ilium in the Dan- iels specimen. The elements of the pelvis were undoubtedly hung loosely in the flesh, as in modern salamanders, since there is no indication of articular surfaces. The hind limb is well known, the type having a nearly complete leg with the foot. The Daniels and the Yale specimens supplement and substantiate the type. The femur is longer than the humerus, but more slender, with its articular surfaces Fig. 28. — Restoration of probable appearance of Amphibamus grandiceps Cope on the basis of the material described herewith. X 1.5. about as well developed as in the humerus. The element is a simple rod of bone without muscular crests of any kind. The tibia and fibula are, likewise, slender separate rods of bone. The tarsus is unossified. The phalangeal formula is 2-2-3- 4-3, and is fairly definite. In the type specimen the matrix in the orbit was blackened as if by the pigmen- tum nigrum of the choroid. The same has been noticed in other specimens. Pro- fessor Cope thought this indicated that the animal was nocturnal. There are many characters in Amphibamus which seem to approximate the rep- tilian type of structure. Among these may be mentioned the character of the artic- ular surfaces of the limb bones, the intercentral position of the ribs, the incipient double-headedness and the curvature of the ribs, the presence of a cartilaginous calcified pubis, the length of the limbs, and the clawed character of the phalanges. Amphibamus was a low, flat, short, and undoubtedly a creeping, crawling ani- mal, possibly spending a portion of its time in the water; but it could not have been a swimmer. It was one of nature's first attempts at constructing a land vertebrate. THE MICROSAURIAN FAMILY AMPHIBAMID.E. 131 Measurements of Amphibamus ukandiceps Cope. Collection of Mr. L. E. Daniels, of Rolling Prairie, Indiana: mm. Entire length of specimen 62 Posterior width of head 15 Length of head 15 Posterior height of skull 3 Length of orbit 5 Width of orbit 3.5 Interorbital width 4 Width of skull in front of orbits n Width of skull just back of orbits 16 Length of presacral region of the vertebral column 30 Length of tail 13 Length of fore limb 13.5 Length of humerus 4 Length of radius and ulna 3 Length of right hand as preserved 3.5 Length of rib along curve 5.5 Length of hind limb 17 Length of ilium 4 Length of vertebral centrum 1 .75 Length of portion of scapula (?) preserved. 4.5 Length of foot 6.5 Width of impression of body midway 16 No. 794 (1234), Yale University Museum: mm. Length of skeleton 67 Length of skull 15 Posterior width of skull 15 Depth of tympanic notch 4 Width of tympanic notch 6 Long diameter of the orbit 7 Transverse diameter of the orbit 5.5 Interorbital width 4.5 Diameter of pineal foramen .75 Length of cervical series of vertebra? 9 Length of dorsal series 35 Length of caudal series 13 Length of a centrum of the dorsal series .... 1 .5 Length of dorsal rib 3.5 Length of arm 20 Length of humerus 7 Length of radius and ulna 4 Width of carpal space 3 Length of third digit 5 Length of leg 25 Length of ilium 3 Length of femur 9 Length of tibia and fibula 5 Length of carpal space 4 Length of 1st digit 3 Length of 2d digit 4.5 Length of 4th digit 7 3 ventral scutellae in 1 mm. Amphibamus thoracatus Moodie. Moodie, Proc. U. S. Nat. Mus., 40, pp. 431-433, fig. 2, 191 1. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, pp. 347-349, pi. 5, fig. 2, 1912. Type: Specimen No. 4306, U. S. National Museum. Horizon and locality: Mazon Creek shales, near Morris, Illinois. The type is a part of the collection of Mr. R. D. Lacoe, in the U. S. National Museum. The fossil is very poorly preserved, but the remains are to be seen on both halves of the nodule, so that considerable can be made out as to its structure. The chief diagnostic characters which will at once distinguish the species are the elongate arm, large interclavicle, shape of the vertebra, and triangular skull. The portions of the animal which are preserved are the impression of the skull with one orbit, the right humerus and radius with portions of others, and traces of ventral scutellae. These remains are so intermingled with the remains of plants that it has been quite difficult to distinguish bone impression from plants. This, however, has been done by whitening the fossils with ammonium chloride, when the texture of the fossils serves to distinguish the one from the other. Parts of the plants have been converted into galena and kaolin, as have also parts of the bones, so the task has been rendered doubly difficult. There can be no doubt, however, that the observations recorded below are correct. The position of the arm in relation to the pectoral girdle and the position of the girdle in relation to the skull impression first called attention to the possible presence of a fossil amphibian. There is little to be said of the skull. It is merely an impression in the nodule. It is triangular in form, with the snout an acute angle. The angle is, however, exag- gerated by the compression to which the fossil has been subjected. The right side of the skull lies over a portion of some plant. The animal is preserved on its back, so that this gives a good opportunity for the study of the pectoral girdle, which is par- 132 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. tially preserved. The interclavicle is very large and from it the species has been given its specific name {thoracatus — armed with a breast plate) . It is an exaggerated T, with the stem very short with its anterior margin curved, and ending in a rather sharp, elongate point. The interclavicle recalls, in a measure, the same element of the Branchiosauria, although it is much more expanded anteriorly and has a shorter spine. In these respects it resembles more nearly a reptilian interclavicle (fig. 14 B) . The clavicle is of the simple triangular shape so characteristic of the Microsauria. It is somewhat displaced backward and its inner margin is slightly obscured. The humerus is elongate, apparently cylindrical, and with expanded ends, resembling very closely the humerus of Amphibamus grandiceps, although its proportions are much greater than in that species. Its length is almost equal to the length of the skull, while in A. grandiceps the length of the humerus is only half that of the skull. The radius (ulna?) resembles in its general proportions those of the humerus. It is a more elongate, slender, lighter bone. The impression of the other bone of the fore- arm is obscured. A portion of a single vertebral centrum from the posterior part of the dorsal series is preserved. It is apparently amphiccelous ; its width is nearly half greater than its length. Measurements of the Type of Amphibamus thoracatus Moodie. (No. 4306, U. S. National Museum.) mm. mm. Length of entire specimen, as preserved 60 Greatest transverse diameter 3 Length of skull impression 18 Length of humerus 10 Greatest width of same 15.5 Greatest diameter ( >f same 4 Long diameter of right orbit . 4 Least diameter of same 1.5 Transverse diameter of same 3 Length of radius (ulna?) 11 Transverse width of interclavicle 14 Length of vertebral centrum 2 Long diameter of same 7(?) Width of same 3 Long diameter of clavicle 9 Genus CEPHALERPETON Moodie. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 349, 1912. Type: Cephalerpeton ventriarmatum Moodie. This genus is founded on remains of a nearly entire individual of a relatively large microsaurian from the Mazon Creek shales. The genus is most immediately related to the Amphibamida?, of which two species are already known, Amphibamus grandiceps Cope and A. thoracatus Moodie. The present genus differs from these species in many respects, notably in size. The skull in Cephalerpeton is nearly as long as half the entire body of Amphibamus grandiceps Cope, inclusive of the tail. Other structural differences are the anisodont teeth, the large size and the more median position of the orbits, and the absence of the posterior tympanic notch in Cephalerpeton. The form of the skull recalls that of Melanerpeton and Pclo- saurus (190) of Europe, but those genera are branchiosaurian, while the present form, from the structure of the vertebrae and the long, curved ribs, is an un- doubted microsaurian. Nothing like it occurs in any of the amphibian faunas thus far made known. It is most nearly approached by a member of the genus Erpetosaurus, but from this genus the present form is readily distinguished by the smooth skull bones, the absence of a posterior table to the skull, and the presence of a highly developed ventral armature. The interorbital width is less than the transverse diameter of the orbit. PLATE 18 &A 4. Type specimen of Erpetosanrtis sailptilis Moodie, from the Cannelton Shales of Pennsyl- vania. Original in the University of Chicago, Walker Museum. Skeletal elements of Eryops sp. indet., from the Pittsburgh Red Shale at Pitcairn, Pennsylvania. <7=nearly complete vertebra; b and r=ribs; hio, Coal Measures. Originalin American Museum of Natural History. X 1. Skull of Sauroptcnra scntcllata Newberry. From the Coal Measures of Ohio. X 1. Tooth of Mastodonsaurus sp. indet. of the Carboniferous of Kansas. Original in University of Kansas Museum. X 1. Tooth of Mastodonsaurus °ivan/eus Jaeger, from the Triassic of Germany. Introduced for comparison with the tooth from the Kansas Carboniferous. XI. THE MICROSAURIAN FAMILY NYRANIIDjE. 1 37 Micrerpeton caudatum Moodie (462, 478) from the Mazon Creek, Illinois, beds, and Sir William Dawson (208) described scales accompanying several forms from the Joggins deposits of western Nova Scotia. Measurements of the Types of Ichthyerpeton squamosum Moodie. Length of animal as estimated from two impressions 3 ft. Length of longest impression 21 in. Length of specimen containing tail impression 9 in. Width of tail impression: Maximum 50 mm. Minimum 6 mm. Width of a single scale 1 mm. Distance from base of tail to tip 125 mm. Length of specimen as preserved 225 mm. Width of chevron rod space 30 mm. Length of rib 25 mm. 8 chevrons in a distance of 3 mm. Genus CERCARIOMORPHUS Cope. Cope, Proc. Amer. Phil. Soc, 1885, p. 405. Type: Cercariomorphus parvisquamis Cope. The type specimen of this genus is supplemented by a portion of the body of another specimen which adds a little to our knowledge of the animal's form, but nothing as to structure. Cope's original description is as follows : "Represented by a fusiform body which terminates in a long, slender, cylindrical tail, and which is covered with small subquadrate scales quincuncially arranged. No fins or limbs are preserved, and the form of the head can not be made out. Probably a portion of the skull is preserved. There are some scattered bodies in the body portion, which look like deeply concave vertebrae with the zygapophyses of batrachians. There are some linear impressions at one point, which resemble the bristle-like rods on many Stegocephali. They are so few as to be of little importance. The scales are like those of fishes. There are traces of segmentation in the axis of the long tail. "The position of this curious form is quite uncertain. It is quite different from any- thing observed hitherto in the American Coal Measures." Cercariomorphus parvisquamis Cope. Cope, Proc. Amer. Phil. Soc, 1885, p. 405. Moodie, Science, n.s., xli, No. 1056, p. 463, 1915. Type : Specimen No. 2560, Newberry Collection, American Museum of Natural History. Horizon and locality: Discovered by Samuel Huston at the Linton, Ohio, Coal Mines. (Plate 21, figs. 3, 4; 24, figs. 2, 3.) The scales (plate 24, fig. 2) in their present condition are entirely smooth. At a distance of 20 mm. from the base of the tail they are in 20 longitudinal series. At that point the transverse diameter of the body is 140 mm. The outline contracts rather abruptly to the tail, of which 66 mm. are preserved. The surface of the tail is obscured by a thin layer of carbonaceous matter not sufficiently thick to obscure scales, which are evident at distances of 16 mm., 43 mm., and 52 mm. from the tip. The scales on the tail are smaller than those on the body and are without markings of any kind. The anterior half of the body is depressed and distorted, but the remainder is well preserved and shows a fairly good outline of an apparently limbless body. I38 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. An additional specimen (No. 8683 G, of the Newberry Collection, American Museum of Natural History) reveals no new facts as to structure, but serves to show that the body of the animal was long and slender (plate 21, fig. 4). The por- tion studied comes undoubtedly from the middle of the body. No limb elements are preserved. The scales are somewhat larger, especially toward the sides of the body, than in the type. The fragment measures 70 mm. in length by 18 mm. and 26 mm. in width. One of the largest scales measures 1 mm. in diameter. Measurements of Type. mm. Length of entire remains 180 Greatest width 22 Greatest width of undisturbed portion 15 Length of an individual scale .75 CHAPTER XIX. THE AISTOPODOUS MICROSAURIAN FAMILY PTYONIID/E, FROM THE COAL MEASURES OF OHIO. Family PTYONIID^ Cope, 1875. Cope, Gcol. Surv. Ohio, 11, pt. 11, p. 357, 1875. Elongate, slender, weak-limbed, aquatic microsaurians. Neural and haemal spines of vertebra elongated, expanded and sculptured. Ventral armature weakly developed or absent. Skull lanceolate, with long, slender teeth. Three genera are assigned to this family: Ptyonius, (Estocephalus, and Thyr- sidium. The forms are very closely related, and when additional material is secured the three genera may be found to be identical. The species included in this family are: Ptyonius pectinatus Cope, P. vinchellianus Cope, P. marshii Cope, P. nummifer Cope, P. serrula Cope, (Estocephalus remex Cope, 0. rectidens Cope, Thyrsidium fas- ciculare Cope. The species are all exclusively from the Linton, Ohio, Coal Measures, and most of them are known from abundant material. Genus PTYONIUS Cope, 1875. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 373, 1875. Cope designated no species as the type, but we may regard Ptyonius pectinatus as typical. Form elongate, with long tail and lanceolate cranium. Limbs weak, a posterior pair only discovered. Three clavicular elements; abdomen protected by packed osseous rods, which are arranged en chevron, the angle directed forward. Neural and haemal spines of caudal vertebrae expanded and fan-like. Ribs well developed. The various species vary in length from 3 to 10 inches. They are the most abundant amphibian in the Linton beds. The present genus resembles Lepterpeton Huxley (334), of the Kilkenny, Ireland, Coal Measures. But that genus possesses divided abdominal rods, or "oat-shaped scales," and the form of the cranium and propor- tions of the body are different. The genus is closely related to, possibly identical with, (Eslocep)ialus, but addi- tional material will be required to settle this point. Cope (123) gives the following key for the separation of the 5 species: x. Abdominal rods coarser, not more than 10 in 5 mm. Median pectoral shield discoid, radiate ridged; muzzle short P. nummifer Median pectoral shield oval, pitted and ridged P. marshii xx. Abdominal rods hair-like, 15 or more in 5 mm. Median pectoral shield with radii from the center, the principal forming a cross; form wider. . P. vinchellianus Middle pectoral with pits at the center and few or no radii; form narrow P. pectinatus Middle pectoral shield narrow, closely reticulate medially, and radiate towards the circumference; size half that of last P- serrula Ptyonius pectinatus Cope. Cope, Proc. Acad. Nat. Sci., 1868, p. 216. Cope, Trans. Am. Phil. Soc, xiv, p. 20, 1869. Cope, Trans. Am. Phil. .Soc, xv, p. 266, 1874. Cope, Geol. Surv. Ohio, 11, pt. n, p. 377, pi. xxvii, fig. 7; xxviii, figs. 2, 3, 6; pi. xxix, fig. 2; pi. xxx, fig. 2; pi. xxxv, figs. 1-3; pi. xli, fig. 1, 1875. Moodie, Proc. U. S. Nat. Mus., 37, p. 24, pi. 8, fig. 3, 1909. Schwarz, Beitrage zur Paleontologie und Geologic Osterreich-Ungarns und des Orients, Bd. xxi, p. 83, figs. 23, 24, 26, 1908. 139 140 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Type : It is impossible to determine which one of the specimens is the type. There are numerous representatives of the species, as follows: Nos. 140, 1096 G, 8345 G, 8555 G, 1089 G, 2, 132, 133, no number, 1094 G, 8545 G, 8677 G, 1159 G, 105, no number, 1091 G, ya, 1092 G, 1093 G, 1095 G, 153, and others unnumbered in the American Museum of Natural History; in the U. S. National Museum are the fol- lowing: Nos. 4458, 4463, 4464, 4514. (Plate 20, fig. 2.) Horizon and locality: Linton, Ohio, Coal Measures. The most abundant species of the Linton Coal Measures. There are over three dozen specimens preserved in the Newberry collection. The species is a clearly marked one, as a rule, though there is great variation in the size of the body and the form of the vertebra?. Though there are several apparently complete skulls pre- served in the collection, it is impossible to make out the morphology of the ele- ments on account of the amount of crushing to which the skulls have been subjected. The head is lancet-shaped, and the muzzle very elongate, slender, and acute at the extremity. The head is in fact a miniature of an ichthyosaur cranium. (Plate 20, fig. 2.) The orbits are large and posterior to the median line. The anterior por- tion of the skull is narrow, poste- riorly truncate, and the mandibular angle is projecting. The posterior portion of the mandible is sculp- tured. Possibly the entire cranium was also, and this has been lost; in fact, this sculpturing is indicated in one or two specimens. The teeth are conical and sharp, longitudi- nally striate, and anisodont. There seemstobe evidence of palatine or pterygoid teeth, though this needs confirmation. The pectoral plates are well preserved, with the interclavicle a narrow oval, with anterior and posterior prolongations. In one speci- men it is sculptured. The clavicles are narrow and slightly sculptured. The abdom- inal scutellae are bristle-like. The vertebrae are short, with expanded neural and haemal spines. The expanded condition of the neural spines begins over the thoracic region, where they are low. They become well developed in the posterior dorsal region. The caudal fan-shaped spines are larger. The dilated portions form equilateral triangles which stand on moderately short pedicels. They are weakly ridged, and each ridge is prolonged into a narrow acute tooth beyond the margin, 1 1 of which may be counted on one of the best -preserved spines. The longitudinal striae are terminated near the pedicel by two others which cross obliquely from each side, and, meeting, present the appear- ance of the margin of a cup sculptured in relief, from which the striae arise. Pedicels smooth. The spines are in contact at their angles, thus forming a continuous line. In a typical specimen there are 6 in half an inch, in another 7, and in a third 8. The ribs are well-developed and slender. Fig. 30. — Restoration of Ptyonius. X I. THE AISTOPODOUS MICKOSAURIAN FAMILY PTYONIID/E. 141 No traces of fore limbs have been detected in the numerous specimens, but ele- ments of hind limbs are preserved. In one of these the femur is a small bone, contracted at the middle. The form of the body is snake-like. There were probably from 75 to 100 vertebrae in a single animal. The form may be well compared to the modern Amphiuma so far as appearances are concerned; structurally they are widely separate. This species is one which is peculiarly char- acteristic of the Linton fauna. Measurements of Ptvonius i-ectinatus Cope. Nos. 107 and 10Q4 C, American Museum. Measurements of a small jaw, No. 8555 C, American Museum. mm. mm. Length of specimen 137 Length of jaw 15 Length of skull 26 Greatest width 1.5 Posterior width of skull 8 Length of tooth I Intcrorbital width 3 Measurements of specimen No. 4458, U. S. National 1 Mameter of orbit 1.5 Museum. Vertical expanse of vertebra 6 Length of specimen '. 65 \\ ldth of neural fan 2 Length of skull 22 Diameter of pedicel I Width of skull 6 Ptyonius vinchellianus Cope. Cope, Proc. Am. Phil. Soc, p. 177, 1871. Cope, Geol. Surv. Ohio, 11, pt. n, p. 376, pi. xxviii, fig. 1, 1875. Type : Specimen in the American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The species is represented by the opposite halves of a single specimen, which includes only the cranium and anterior half of the body. The fan-shaped neural spines commence but a short distance behind the line of the pectoral shields. They are low, with a few coarse ridges, the margin being entire. The abdominal rods are delicate and hair-like. The interclavicle is oval, with a few radiating crests, which originate at the center; in the areas behind there are a few scattered tubercles. The clavicles are ridged near the margin. The cranium is lanceolate in form, and the bones of the dorsum are marked with a few raised points and ridges. The species is about the size of Ptyonius pectinuttis Cope, and differs, apparently, from that species in the rather insignificant charac- ter of a narrower interclavicle and in the ornamentation of the same. Dedicated to Professor Alexander Winchcll, of the University of Michigan. Measurements of Ptyonius vinchellianus Cope. mm. Length of cranium 20 Width of same 8 Length of interclavicle 4-2 Ptyonius marshii Cope. Cope, Trans. Amer. Phil. Soc., xiv, p. 24, 1869 (Colostcus marshii). COPE, Geol. Surv. Ohio, 11, pt. 11, p. 375, pi. xxvii, fig. 6; pi. xxviii, fig. 3, 1875. Cope, Proc. Am. Phil. Soc, xn, p. 177, 1871. Type: Specimen No. 1157 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The head is elongate lanceolate. The upper surface of the frontal bones is punctate-rugose in relief, with short radii toward the margin. The distal two- I42 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. thirds of the mandible is narrow wedge-shaped ; the external surface is coarsely pitted. There are no teeth preserved. The pectoral elements are displaced, but the clavicles are subtriangular, and are strongly ridged toward the inner margin. The interclavicle is short spatulate, the narrow portion directed anteriorly; the posterior rounded. It is coarsely pitted medially, and coarsely and strongly radiate-ridged to the margin. The abdominal armature commences immediately behind the pectoral girdle. It consists of elongate, narrow, subcylindric scales, which meet on the median line, converging anteriorly. Small limbs are present. Measurements of Ptyonius marshii Cope. Type specimen: mm. Length of fragmentary skull 7 Width of skull as preserved 5 Length of entire specimen 68 Width across pectoral plates 8 Width across belly 7 Specimen No. 1098 G, American Museum: Length of specimen 32 Width of specimen 7 4 scutes of abdominal armature in 1 mm. Ptyonius nummifer Cope. Cope, Geol. Surv. Ohio, 11, pt. 11, pp. 374, 375, pi. xli, figs. 2 and 3, 1875. Moodie, Bull. Am. Mus. Nat. Hist., xxvi, p. 356, pi. 63, fig. 3, 1909. Type: Specimen No. 8546 G, American Museum of Natural History. No. 8614 G, same museum, is associated with the type specimen. Horizon and locality: Linton, Ohio, Coal Measures. Two well-preserved individuals display peculiarities which indicate specific dis- tinctness from the previously known species of Ptyonius. The abdominal rods are of the coarse type of those of P. marshii. The caudal fans are well developed, and not so wide as in P. pectinatus. The interclavicle is a discoid body of different form from that of P. marshii and I can not detect the clavicles. The sculpture consists of strong ridges, which radiate from the center to near the border. Immediately in front of this interclavicle is the head, which has a different form from that of the other known species. The interorbital width is two-thirds the long diameter of the orbit. The structure of the skull can not be made out. A slender, elongate hind limb is present in the second specimen, and a humerus is well preserved in the type. Measurements of the Type of Ptyonius nummifer Cope. (No. 8546 G, American Museum of Natural History. No. 8614 G is associated in the same species.) mm. mm. Length to beginning of caudal fans 65 Width of abdominal armature 8 Length of head 15 Length of a caudal fan 2.5 Length from muzzle to orbits 6 Length of femur 5 Length of interclavicle 7 Proximal width of femur 1.5 Width of interclavicle 8 Ptyonius serrula Cope. Cope, Proc. Am. Phil. Soc, 1871, p. 177. Cope, Geol. Surv. Ohio, 11, pt. n, p. 379, pi. xxviii, fig. 5; pi. xxx, fig. I, 1875. Type: Specimen No. 8615 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The specimens of this species indicate that the form was only about half the size of Ptyonius pectinatus. The interclavicle is narrower and more reticulately sculp- THE AISTOPODOUS MICROSAURIAN FAMILY PTYONIID^. I43 tured. The tail is relatively longer. Abdominal rods hair-like. Ribs distinct. Small limbs are present in one specimen. Measurements of Cope's Type of Ptyonius serrula. mm. mm. Length of specimen 95 Width of clavicle 1.5 Length of portion of skull preserved 3 Length of vertebra I Length of interclavicle 4.5 Width of vertebra from tip of neural spine to tip Width of interclavicle '. 2 of haemal spine 4 Length of clavicle 4 Another specimen (456 G, American Museum of Natural History) shows some of the same characters. There is not the slightest basis for the support of- this spe- cies, so far as I can observe. The ones mentioned by Cope are insufficient. It is in all probability a mutant or variety of Ptyonius pectinatus. Genus (ESTOCEPHALUS Cope, 1868. Cope, Proc. Phila. Acad. Nat. Sci., 218, 1868. Cope, Trans. Amer. Phil. Soc, xiv, 16. Cope, Proc. Phila. Acad. Nat. Sci., 1868, 217. Cope, Proc. Amer. Phil. Soc, 1871, 41. Cope, Geol. Survey Ohio, n, pt. 11, 380, 1875. Type : (Estocephalus remex Cope. Form slender and snake-like; caudal vertebras with elongated, dilated, sculp- tured neural and haemal spines. Cranium lanceolate. Teeth numerous, of nearly equal size. No pectoral shields known; abdomen protected by very numerous bristle-like rods, which converge forwards. A pair of weak posterior limbs ; branchi- hyal bones present. In the only well-preserved species the cranial bones exhibit no sculpture from the parietal region forward. The genus is not very distinct from Ptyonius, but it can not be united with that genus until more complete material is available. The species of the genus share with Ceraterpeton, Urocordylus (334), and Ptyonius, as well as Crossotelos (98), from the Permian of Oklahoma, the elongation, sculpture, and expansion of the neural and haemal spines. There are but 2 species, which Cope distinguishes by the following characters: I. Vertebras elongate; fan-like caudal processes narrowed. Size large; mandibular teeth of unequal lengths, with the apices turned backward (Estocephalus remex II. Species only known from cranial bones with teeth. Teeth equal, erect, with acute conic apices, II in 5 mm (Estocephalus reclidens (Estocephalus remex Cope. Cope, Proc. Phila. Acad. Nat. Sci., p. 217, 1868 (Sauropleura remex). Cope, Proc. Phila. Acad. Nat. Sci., p. 218, 1868 ((Estocephalus amphiumianus). Cope, Trans. Amer. Phil. Soc, xiv, p. 17. Cope, Geol. Surv. Ohio, vol. 11, pt. 11, p. 381, pi. xxvii, fig. 5; pi. xxxi, fig. 1; pi. xxxii, fig. 1; pi. xxxiii, fig. 2; pi. xxxiv, fig. 4, 1875. Moodie, Proc. U. S. Nat. Mus., 37, p. 27, 1909. Type: Specimen undetermined. The following specimens are to be found: Specimens of GLstocephalus remex Cope in the National Museum, Nos. 451 1, 4460, 4478. There is one specimen of (Estocephalus remex in the University of Chicago. Specimens of the species in the American Museum of Natural History: Nos. 121, 8322 G, 8694 G, no number, 8656 G, 8583 G, 8659 G, 19, 120, 8655 G, 8662 G, 8708 G, 8665 G, 112, 8663 G, 8581 G, 8658 G, 8660 G, 8700 G, 8469 G, 1102 G, 1 152 G 142, 8381 G, and obverse, 21, 8664 G, 8672 G, 8592 G, 8684 G. Horizon and locality: Linton, Ohio, Coal Measures. 144 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. This species is one of the most abundant of the Linton Amphibia. Cope based his description of the species on 9 specimens. There are more than two dozen avail- able at the present time, the majority of them being in the possession of the Amer- ican Museum of Natural History. There is a single specimen in Walker Museum of the University of Chicago and 3 in the United States National Museum. The numbers of all of these specimens are given above. The material consists, for the most part, of fragmentary portions of the vertebral column, but there are a few skulls more or less complete, though none are sufficiently well preserved for a com- plete analysis of the characters. The specimens indicate an animal slightly smaller than the modern Amphiuma means of the Mississippi River. It will not be necessary here to enter into a detailed account of each specimen, since this has been done by Cope, and a careful comparison of his descriptions with the originals indicates that his observations are correct. The species, as suggested in the discussion of the genus, is not clearly distinct from those of Ptyonius, and it has largely the characters of that genus. The cranium is long, slender, and wedge- F10. 31. — Restoration of CEstoccphalus. X I. shaped. The teeth are numerous both in the maxillary and in the mandible, one specimen indicating about 30 in a single series. They are all uniformly cylindrical, except at the extremity, where they are flattened and expanded so as to produce a longitudinal edge, which is carried backward on a recurvature of the apex. The bases are anchylosed equally and without enlargement, and no part of the shaft is striate or grooved. The upper surface of the cranium is narrow, with the median suture distinct. The skull surface, with that of the mandible, is smooth. Characteristic of the species are the remarkable length and slenderness of the fan-shaped neural and haemal spines, and the absence of an acute serration on their margins. In this species the spines have a laminiform expansion at the base in their plane. One specimen exhibits the pelvic region, including a portion of the tail. The ilium has an expanded anterior extremity and is directed backwards and somewhat inwards on either side of the vertebral column. The femur is nearly straight, short, contracted medially, and expanded distally. The tibia is shorter and is subcylindrical. Beneath the ilium the last chevron of the abdominal rods appears, the outer extremities rising on the base of the tail. The pectoral arch is almost unknown, and Cope based the distinction of Ptyo- nius and CEstocephalus on the absence of these plates in the latter genus — an uncer- THE AISTOPODOUS MICROSAURIAN FAMILY PTYONIIMs. I45 tain characterization. The fore limbs are indicated by a humerus. There were possibly from 75 to 100 vertebrae in the entire column. The animal was exclusively adapted to life in the water and was, without doubt, an excellent swimmer. There are preserved in one specimen portions of what seem to be hyobranchial elements. Measurements of CEstocephalus remex Cope. mm. mm_ Length of entire caudal series 195 Length of mandibular dental series 24 Width at ninth vertebra 21 Depth of mandible at middle ^ Width at thirty-sixth vertebra 2 Nine teeth in 5 Length of four phalanges in place 14 Length of longer teeth 2 Expanse of fan of proximal caudal 18.5 Length of first hsmal branchial 6 Length of ilium 1 1.5 Length of sixth vertebra from skull 7 Length of femur 11. 5 Width of centrum 3 Length of tibia 7 CEstocephalus rectidens Cope. Cope, Trans. Amcr. Phil. Soc., p. 268, Apr., 1874. Cope, Geol. Surv. Ohio, 11, pt. n, p. 386, pi. xxvii, fig. 3, 1875. Type: Specimen No. 9033, American Museum of Natural History, collection of J. S. Newberry. Horizon and locality: Linton, Ohio, Coal Measures. The species is indicated by a left dentary bone, with its teeth and external sur- face preserved. The latter is nearly smooth and without sculpture. The outer face is convex, and the general form is slender, but not curved upward at the extrem- ity. The extremity of the dentary does not show any evidences of teeth. Teeth straight and conic, apex acute, non-plicated. Cope also associated with this species a portion of a caudal series, consisting of 25 vertebrae. The centra are elongate and expanded at the extremities. The neural arches have a close union. The neural and haemal spines are fan-shaped and striated. The bases are quite narrow. Measurements of QSstocephalus rectidens Cope. mm. mm. Length of dentary 22 Length of 3 centra of the caudal series 8.6 Length of tooth line 15.2 Extent of neural and haimal spines 8.7 Depth of dentary at last tooth 2.7 Genus THYRSIDIUM Cope, 1875. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 365, pi. xxxi, fig. 2, 1875. Type: Thyrsidium fascicular e Cope. Established on a species which presents its principal peculiarities in the struc- ture of the vertebrae. Two specimens present inferior views of the spinal column, showing that the genus possesses, like Siren, enlarged diapophyses, but they are peculiar in their fan-like form. They resemble slightly the neural spines of the caudals of Ptyonius, but are present on the dorsal vertebrae. Whether the caudals of the present species possess ornamented neural spines the specimens do not indi- cate. The abdomen is protected by the usual hair-like rods arranged en chevron, the angle directed forwards. No indications of limbs can be discovered on the blocks. I46 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Without the cranial bones the affinities of this genus can not be determined; while it may be allied to Cocytinus, the vertebrae of that form are without peculiar diapophyses. Thyrsidium fasciculare Cope. Cope, Geol. Surv. Ohio, n, pt. n, p. 365, pi. xxxi, fig. 2, 1875. Type: Specimen No. 8552 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The best preserved example of this species includes 9 vertebrae and the corre- sponding ventral armature. The centra, seen from below, are much contracted in their form, presenting an obtuse median rib, which expands to the articular extrem- ities. In one or two instances the latter are divided by fracture, and the moder- ately concave form of the adjacent surfaces is displayed. The diapophyses are of complex form, but the details are concealed by the prevalent thin layer of coal which invests them. An inferior prominence runs parallel to the centrum; outside of this the process is obscurely trilobate and thickened, not flattened, as in the caudal ver- tebras of Ptyonius. Several ribs (fig. 8) of moderate thickness appear by the side of the diapophyses. Eleven abdominal ribs in 5 mm. The second specimen was originally referred to QLstocephalus rcmcx, as a pos- terior portion of its vertebral column, immediately preceding the caudal series. This reference appears to be incorrect, although the resemblance between the corresponding parts in the two genera is no doubt considerable, and the alternative of proposing a new genus and species was not at that time advisable. The neural spines are longer than high, and are nearly in contact at their mar- gins; each is marked by about 5 obtuse vertical ribs. A fractured section of the abdominal spines in place displayed at least six layers of them. The material on which the above account is based is imperfect. The spec- imen figured by Cope (Geol. Surv. Ohio, vol. 11, pt, 11, pi. xxxi, fig. 2, 1875) is undoubtedly a portion of the vertebral column of QLstocephalus remex. Nos. 4462 and 4480 of the United States National Museum may be representatives of Thyr- sidium fasciculare, but they are more probably QLstocephalus remex; if they are the latter this leaves the type as the only known specimen of the species. Measurements of Type of Thyrsidium fasciculare Cope. mm. Length of portion of vertebral column preserved 70 Width of ventral armature 18 Length of a vertebra 9 Width of same vertebra 4 CHAPTER XX. THE MICROSAURIAN FAMILY MOLGOPH1D/E, FROM THE COAL MEASURES OF OHIO AND MAZON CREEK, ILLINOIS. Family MOLGOPHID^ Cope, 1875. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 357, 1875. Cope, Bull. U. S. Nat. Mus., No. 1, p. 11, 1875. Type of family : Molgophis. Body long, serpentine, a few species apparently limbless, ribless, and with abdominal armature lacking. Vertebras elongate, neural and haemal spines short or absent. Ribs long, heavy, and broad. The vertebrae seem to bear the charac- teristic marks of the family. One species has the skeleton reduced to a lanceolate skull and a string of about 50 slender vertebras, all the rest of the skeleton being absent. The family is very poorly known, but was apparently of wide distribu- tion in North America and confined to this continent. The representatives of the group are known from Iowa, Illinois, and Ohio. Four genera are assigned to the family, but future discoveries will undoubtedly demand revision of the present classification. The genera are Molgophis, Pleu- roptyx, Phlegethontia, and Erpetobrachium. The distinguishing characters of these genera are apparent from the descriptions of the various forms. The skeletons of the species are too incompletely known to allow the establishment of a tabular key to the genera. Genus MOLGOPHIS Cope, 1868. Cope, Proc. Phila. Acad. Nat. Sci., p. 220, 1868. Cope, Trans. Amer. Phil. Soc, xiv, p. 20, 1869. Cope, Geol. Surv. Ohio, it, pt. 11, p. 368, 1875. Cope, Trans. Amer. Phil. Soc, xv, p. 263, 1874. Type: Molgophis macrurus Cope. Cope (123) gives the following: "The characters of this genus are: body long, serpentine, without dermal armature, so far as known; vertebras long and broad, with very prominent zygapophyses and moder- ate neural spines; ribs large, curved. No limbs or cranium can be ascribed to the type of the genus. The ribs are long, and though the head is not bifurcate, there appears to be both tubercle and head on the dilated extremity. Where crushed they display a large median vacuity. "This genus differs from Ophiderpeton Huxley (334) in the characters of the dorsal vertebrae, which, in their projecting zygapophyses, resemble those of Amphiuma. The lack of ventral armature distinguishes it from CEstocephalus, while its well-developed ribs separate it from Phlegethontia.'" Molgophis macrurus Cope. Cope, Proc. Phila. Acad. Nat. Sci., p. 220, 1868. Wyman, Am. Jour. Sci. and Arts, p. II, fig. I, 1858 (refers to a batrachian reptile). Cope, Trans. Amer. Phil. Soc, xiv, p. 20, 1869. Cope, Trans. Amer. Phil. Soc, xv, p. 263, 1874. Cope, Geol. Sun'. Ohio, 11, pt. U, p. 368, pi. xliii, fig. I, 1875. Type: Specimen No. 8617 G, American Museum of Natural History, collec- tion of Dr. J. S. Newberry. Horizon and locality: Linton, Ohio, Coal Measures. 147 148 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. This species is established by remains of 2 individuals, one embracing 16 and the other 14 vertebrae, with ribs. The neural arches, viewed from above, have a V-shaped outline posteriorly, from the fact that the broad zygapophyses meet on the median line and spread out distally over the broad anterior ones adjoining. The latter appear to be somewhat concave and to border the former exteriorly as well as inferiorly. The base of the neural spine extends to the posterior emargination, but not quite to the anterior. The breadth of the dorsal vertebrae above is equal from the emargination behind to the anterior margin of the anterior zygapophyses. The ribs are long for an amphibian, but not long for a reptile. They are well curved, chiefly near the proximal extremity. The longest found, measured by a cord, equals two and two-fifths vertebrae. These vertebrae, measured along the median line above, equal 11 lines; one of these is 3.6 lines in width above. This animal has been, like Amphiama, a snake-like amphibian, but was probably still larger. How near the affinities to this genus may be can not now be determined, owing to the want of many important parts of the skeleton, but it differs in the important feature of large, well -developed ribs. The size of the vertebrae would indicate a body of the size of the com- mon rattlesnake (Crotalus horridus) and too large for Br achy - dectes newberryi, which is only known from jaws. Molgophis brevicostatus Cope. Cope, Geol. Surv. Ohio, n, pt. 11, p. 369, pi. xliv, fig. 1, 1875. Moodie, Proc. U. S. Nat. Mus., 37, p. 27, 1909. Neura, spine Ribj Centrum Type: Specimen No. 8341 G, Amer. Mus. of Nat. History. Horizon and locality: Linton, Ohio, Coal Measures. Represented by portions of the vertebral column of several individuals. One of these includes 9 pairs of ribs, with verte- brae, and another 13 pairs. The vertebrae are subquadrate in section, and the concavity of the two articular faces is not deep. They support strong lateral ridges separated by deep concavities. The heads of the ribs are somewhat contracted, and the shafts present outward a tubercular angle at a dis- tance of one-fourth the length from the head. The remaining part of the shaft is stout, nearly straight, and gradually con- tracts to an obtuse extremity. It possesses a narrow medullary cavity. In none of the specimens is there any trace of abdominal armature, but abundant remains of the contents of the abdominal cavity, in proper position, are preserved on the blocks. This species is more massive than Molgophis macrurus, and the ribs are shorter, thicker, and less curved. Ventral scutellae are present in one specimen of this species. There are a number of specimens. They have the following numbers at the American Museum: 158, 1 100 G, no number, 8341 G (type), no number, 8466 G; and 4477 in the U. S. National Museum. Fig. 32. — Drawing from Cope's figure of Molgo- phis brevicostatus Cope. X0.5. THE MICROSAURIAN FAMILY MOLGOPHID^E. I49 Measurements of Type Specimen of Molgophis brevicostatus Cope. mm. mm. Length of 7 vertebra 105 Length of a rib on a curve 24 Length of 1 centrum 16 Greatest thickness of same 2.5 Diameter of same vertically 11 Molgophis wheatleyi Cope. Cope, Gcol. Surv. Ohio, II, pt. n, pp. 369, 370, pi. xlv, fig. 1, 1875. Cope, Trans Am. Phil. Soc, xv, p. 263, 1874. Type: Specimen No. 1101 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. A critical study of the type specimen of this species does not reveal anything essentially different from the description of Cope. The following is taken from his report (123): "Established on a specimen which exhibits about twenty-five vertebrae with ribs, and the posterior portion of the cranium. No traces of abdominal scales or rods, thoracic shields, or limbs are visible. By such negative characters it is referable to the genus Mol- gophis, although the definition of this genus is incomplete. The present batrachian may, indeed, be ultimately found to be an Ophiderpeton, to which it also bears some resemblance. "The specimen is that of an animal of very much smaller size than the M. macrurus. The vertebra are of moderate length, with a low neural spine, and centrum angular at the sides and truncate at the articular extremities when in place. The ribs are rather short, slightly curved, apparently hollow and intercentral in position. Although the ver- tebral centra are ossified, the elements of the cranium have a larval appearance. These consist of two parallel bony plates, which resemble the fronto-parietal bones of the frog; they are slightly separated from each other, but do not inclose a fontanelle. A wedge- shaped bone extends from the outside of the front of each of these, acuminate behind, and widening anteriorly in the position of a postfrontal bone. In front of the posterior border of each parietal, on its outer side, a bony enlargement arises which contracts out- ward and forward into a narrow element which curves forward beneath the postfrontal. These look like an anteriorly directed quadrate with articular bone, such as seen in the larvae and some adults of existing batrachians. These determinations will require con- firmation from additional material. In the meantime it is evident that the present specimen can not be referred to any of the other species herein described. The elements of the cranium are entirely smooth with no sign of sculpture, and in this respect the present species is unlike any of the other known from the Carboniferous." The vertebrae are not so clearly marked as one is led to believe from Cope's figure. Measurements of the Type of Molgophis wheatleyi Cope. mm. mm. Length of entire specimen 63 Width of a vertebra. . 1.5 Length of portion of skull preserved 9 Length of a posterior rib 5 Posterior width of same 8.5 Width of rib 5 Length of a vertebra 1.5 The species is dedicated to Charles M. Wheatley, of Phcenixville, Pennsylvania, one of the original investigators of the Linton deposits. It is a part of the New- berry Collection. Additional material of this species is represented by specimens Nos. 7 and 8699 G of the American Museum of Natural History. They are both very unsatisfactory. They consist of molds of the vertebral column, with in one case an enlargement at one end which may represent the head, and if such, the specimen probably repre- sents a distinct species. The impression, No. 7, contains molds of about 30 ver- 150 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. tebrae which are very similar in form to those exhibited by the type of the species. To the vertebrae are articulated short, curved ribs of a slender nature. The ver- tebras themselves are short and somewhat constricted in the middle. The other impression, 8699 G, contains impressions of about 20 vertebrae, appar- ently immature, though one can not be entirely sure as to the nature of the struc- tures. They are covered over with a thin layer of carbonaceous material which is impossible to remove satisfactorily. The two specimens remind one of what Hux- ley has written in regard to the forms of Microsauria (334) from Kilkenny, Ireland. Measurements of Nos. 7 and 8699 G (Molgophis whf.atleyi). mm. mm. Length of No. 7 87 Length of vertebra 2 Length of head mold 18 Length of rib 3 Posterior width of head 6 Length of specimen No. 8699 G S3 Genus ERPETOBRACHIUM Moodie, 1912. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 353, 1912. Type: Erpetobrachium mazonensis Moodie. The generic characters are apparent in the greatly elongated fore limb, in the exceptionally broad scapula, the long radius and ulna, which slightly exceed the humerus in length, a character hitherto unknown among Carboniferous Amphibia. Erpetobrachium mazonensis Moodie. Moodie, Kans. Univ. Sci. Bull., vi, No. 2, pp. 353-354, pi. 2, fig. 2; pi. 8, fig. 3, 1912. Type: Specimen No. 799 (222), Yale University Museum. Horizon and locality : Mazon Creek shales, near Morris, Illinois. The scapula of the present form is exceptional in its shape. It resembles an asymmetrical pyramid, the anterior side of the lower edge of the bone being con- tracted so that the anterior edge of the element is arcuate. Its top is very thin and possibly terminated in a broad cartilage. The lower end is thick and heavy and the articular surface is, apparently, well formed, though somewhat obscured. The element identified as a clavicle is lying on its edge and has the proportions of the clavicle of Mazonerpeton costatum. The exterior edge is somewhat rounded and small. A portion of another element which I suppose to represent the coracoid lies alongside the humerus, although its form is quite obscured. (Plate 3, fig. 3.) The humerus has a remarkably well-formed head. In perfection of formation it corresponds well with that of the higher reptiles. This surface can even be divided into an anterior and a posterior articulation. The element projects posteriorly for the distance of 1 mm. from the surface of the shaft. The shaft immediately below the head is somewhat flattened and has an ovoid section. Further on it becomes flattened, a part of which is probably due to pressure during fossilization. The elements of the forearm are both preserved and are approximately equal in size. They are remarkable in that they exceed the humerus in length, although they are not so heavy as that element. They are greatly elongate and slender, with the middle of the shaft only moderately contracted. The articular surfaces are well formed and both bones were hollow, as was also, apparently, the humerus. The ulna may be represented by the most posterior of the two elements, though the relations of the elements may have been reversed (fig. 15, D). THE MICROSAURIAN FAMILY MOLGOPHID^. 151 The base of the left wing of an orthopterous insect possibly allied to Paolia gitrlcyi Scudder lies between the radius and ulna. The nodule also contains impres- sions of plants, a portion of a frond of a Neuropteris and the impression of one of the Cordaites. Lying next the radius is a slender elongate element which may be a rib or a portion of a metacarpal. If a rib, it indicates that the animal belongs among the Branchiosauria. The fragment is only half as long as the radius and is entirely too obscure to base any conclusions. The other characters of the specimen point quite strongly to its microsaurian affinities. The structure of the articular surfaces of the limb bones alone would indicate the microsaurian relationship of Erpetobrachium. It may be provisionally asso- ciated in the family Molgophidae with such forms as Molgophis brevicostatus Cope, Molgophis {Pleuroptyx) clavatus Cope, and Molgophis macrurus Cope from the Coal Measures of Linton, Ohio. Measurements of the Type. mm. mm. Length of scapula 14 Diameter of shaft 2 Distal width 6 Distal width 3 Proximal diameter 3 Length of radius 25 Length of clavicle (?) 24 Proximal width 4 Length of humerus 25 Diameter of shaft 3 Length of ulna 24 Width of distal end 4 Proximal width 4 Genus PLEUROPTYX Cope, 1875. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 370, pi. xlii, fig. 1; pi. xliv, fig. 2, 1875. Cope, Proc. Phila. Acad. Nat. Sci., p. 16, 1875. Moodie, Proc. U. S. Nat. Mus., 37, p. 27, 1909. Type: Pleuroptyx clavatus Cope. The specimens on which the species of this genus repose do not exhibit crania. The 5, probably 6, specimens which represent them offer various views of the ver- tebral column, and in none is there any trace of ventral or thoracic armature. Limbs can be ascribed to them with probability only. The vertebrae are of moderate length, with well-developed zygapophyses, and a short and not very elevated neu- ral spine in the dorsal region, which is not sculptured in any way. The generic character is seen in the ribs. These are rather short and very stout and support an ala on the posterior or convex border, which expands downwards, and then sud- denly contracts to the shaft. The extremity of the latter is broad and truncate, and includes a medullary cavity, which is only partially fitted with cancellated tissue. Pleuroptyx clavatus Cope. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 370, pi. xlii, fig. 1 ; pi. xliv, fig. 2, 1875. Cope, Proc. Phila. Acad. Nat. Sci., p. 16. 1875, Moodie, Proc. U. S. Nat. Mus., 37, p. 27, 1909. Type: Specimen No. 8617 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The general appearance of the species of Pleuroptyx is that of the Molgophis, so far as known, but nothing resembling the peculiar structure of the ribs is seen in any other. There is no assurance that the genus is distinct from Molgophis. 152 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Humerus Ulna. Parts of two individuals express the typical characters of this species, while a third only differs in being considerably smaller. A fourth may very probably be referred here, and another, bearing several elements of a leg, should be most likely associated with the last mentioned. The ribs are considerably narrowed near the head, and appear to possess a low tubercular process some distance below it. The shaft is curved throughout; the laminar expansion is quite thin; while the distal end is expanded and concave, perhaps for the attachment of car- tilage, although no trace of this remains on the shale. The neural spines have short bases, oblique anterior and nearly straight pos- terior borders, with obtuse extremity. I perceive no essential dif- ference in the smaller specimen, which is one-third less than the types. The limb is appropriate in its proportions to the present species, and may be described in this place. The first segment is one-third longer than the second, and has a transversely expanded head. The shaft is stout, the distal extremity not expanded and concave. The second segment is stout, more ex- panded proximally than distally, the proximal end truncate and slightly concave. A bone, much displaced, lies near it, and is fig. 33. probably ulna or radius; it is as stout as the first, the end not expanded. Of metatarsals there are 2, three-fifths the length of the second bone of the leg, and of phalanges 2, of 2 digits each. The proximal are three-fourths the length of the metatarsals, and indicate elongate toes. The obverse of the specimen is preserved, and contains no addi- tional toes or phalanges. Phalanges Fore limb of a member of the Mol- gophida-, possibly Pleuroptyx clavatus Cope. X0.75. Measurements of the Type of Pleuroptyx CLAVATUS Cope. Length of vertebral centrum 14 Depth of a vertebral centrum 9 Depth of entire vertebra 22 Length of neural spine 8 Height of a neural spine 6 Length of a rib on the curve 42 Width of a rib at ala 9 Width of a rib at extremity 5 Length of first segment of leg 38 Length of second segment of leg 24 Length of metapodial bone 10 Length of first phalanx 7 Hitherto only two portions of the dorsal series and a left limb have been assigned to this species. The present specimen (No. 4479, U. S. Nat. Mus.) thus proves of interest in determining that the creature was long-tailed, like CEstocephaius, Ptyo- nius, and Phlegethontia, but unlike the first two genera the neural and haemal spines are not elongate nor marked with radiating lines. The neural spines are indistinct and if developed at all were very low and short. The centra are short, cylindrical, and thick. They gradually decrease in size to where they are lost, since the portion preserved does not represent the entire length of the tail. There may have been 15 more vertebrae distally and 5 more proximally, thus making about 75 caudal vertebrae, as Woodward (630) has deter- mined obtains in Ceraterpeton galvani Huxley. THE MICROSAURIAN FAMILY MOLGOPHID^. 153 The ribs are continuous throughout the length of the tail preserved and have precisely the same structure as is found in the dorsal region with the possible excep- tion that the posterior alar expansion is not so well developed in the caudal ribs. The ribs are decidedly fan-shaped and articulate by a single head with a short transverse process. They are distinctly curved like all microsaurian ribs. Measurements of Specimen of Pleuroptyx clavatus Cope. (No. 4509, U. S. National Museum, Linton, Ohio, Coal Measures.) mm. mm. Length of tail as preserved 105 Length of rib 5 Length of anterior vertebra 1.5 Width of rib I Diameter of anterior vertebra 1 The above-described specimen represents what I suppose to be the posterior part of the body of Pleuroptyx clavatus Cope. The characters of the ribs and ver- tebras are the same. The fragment is interesting, since it gives an insight into the form of the body, which was slender, conforming thus to other long-tailed micro- saurs. Length of specimen, 65 mm. ; width of specimen, 30 mm. There is still a third example of this species among the collections belonging to the National Museum (No. 4484). The specimen includes a badly crushed pos- terior portion of a skull and a series of about 16 crushed vertebrae, with several pairs of ribs and ventral scutes. Very little can be said of the skull save that the maxilla of the right side was long and bore from 15 to 20 teeth, of which 9 are preserved more or less completely. The mandible is likewise crushed and one can not determine its elements. Por- tions of 2 or 3 teeth are preserved. The form of the mandible is long and slender. The ventral scutes are of the pectoral region. They are long, slender, and thread- like. They are not closely packed, but I count 12 in a distance of a millimeter. So far as can be determined the vertebrae are the same as has been described for other specimens. They are short and heavy. The ribs show, for the most part, the same characters as the type specimen. Measurements of Third Specimen (No. 4484, U. S. Nat. Mus.). mm. mm. Length of specimen 60 Length of anterior vertebra 2 Length of preserved portion of skull 15 Length of rib 4 Length of tooth 25 Width of rib I Posterior width of mandible 3 Genus PHLEGETHONTIA Cope, 1871. Cope, Proc. Am. Phil. Soc., p. 177, 1871. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 366, 1875. Type: Phlegethontia linearis Cope. This is one of the most interesting genera of the present series. It rests chiefly on a single specimen of one species, which is not perfect, but which displays the following characters: Head elongate-triangular; body and tail extremely elongate, the dorsal vertebrae without ribs, and the caudals without dilated spines ; no ven- tral armature nor limbs. As a great portion of the length is presented, and no ventral rods or scales are visible, and as this character is confirmed by a second specimen, it probably belongs to the genus. The pectoral shields are also wanting in the spec- 154 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. imen, but as there is a considerable vacuity behind the skull of the specimen, it may be that these were lost with other parts. Chevron bones are not observable on the caudal vertebrae. This form is a true amphibian snake. Phlegethontia linearis Cope. Cope, Geol. Surv. Ohio, II, pt. H, p. 367, pi. xliii, fig. 2, 1875. Type: Specimen in the American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. In the only specimen the dorsal vertebrae are much involved anteriorly, so that the length is not readily ascertained. There is an outline of a triangular object which may represent the skull of this specimen, although it is so far removed from the vertebrae that there is some doubt as to whether it belongs with the vertebrae or not. Indeed, there is even doubt whether it is a skull. The vertebrae have longitudinal diapophysial keels, and have a zig-zag interlocking of neural arches. The latter are distinctly turned outward. The vertebrae are very numerous, and the tail very attenuated. The number preserved is about 60. The total length of the coils unwound is about 295 mm., or 11 coils in 8 lines; but there are interrup- tions not measured and confusions not unraveled. This is the most elongate and slender of all the species of the Carboniferous Amphibia. The vertebrae are apparently ribless and there are no evidences of limbs or pectoral plates. It may be said that the body consists entirely of skull and vertebrae. Measurements of the Type of Phlegethontia linearis Cope. Entire length of skull (?) 18 mm. Length of single vertebra 2.50 mm. Width of same 8 mm. Height of vertebra 1 .50 mm. Length of vertebral column as preserved. . . 295 mm. Estimated length of body 15 in. No. 8370 G, American Museum of Natural History, shows a few vertebrae. Phlegethontia serpens Cope. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 367, pi. 32, fig. 2, 1875. Type: Specimen No. 1102 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. This amphibian is much larger than the last, approaching nearly in its dimen- sions the Molgophis macrurus. It is represented by a series of 22 vertebrae, which, like those of Phlegethontia linearis, are devoid of ribs, abdominal armature, dilated neural spines, etc. The series when complete must have been very long, as there is little difference in size between the first and the last of the 22. They are marginate fore and aft, and much contracted medially, owing to the transverse expanse of the diapophyses. There may be, indeed, a diapophysial element beneath these, but, if so, the two are indistinguishable. They are connected by longitudinal impressions, indicating the existence of the tendinous bands in the longitudinal muscles seen in Amphiuma, or the osseous spicules seen in the same situation in birds. The neural spines, as indicated by their narrow bases, occupy the length of the neural arch, and remind one of Amphiuma. Width of one of the vertebrae, 3 lines. CHAPTER XXI. THE MICROSAURIAN FAMILY SAUROPLEURID^, FROM THE COAL MEASURES OF OHIO. Family SAUROPLETJRIDvE Hay, 1902. Hay, Bull. U. S. Geol. Surv., No. 179, p. 419, 1902. The present family is an association of related forms due to similar structure of vertebrae, ribs, ventral scutellation, and limbs. There is no character in the skull which would indicate a separation of the genera here included, at least in the light of present knowledge. The family may be characterized as : Subaquatic or terrestrial vertebrates with a typically amphibian development of the ventral armature ; ribs intercentral, as in all members of the order; skull elongate and slender or broad and obtuse; cranial and dermal elements of the pectoral girdle sculptured; lateral-line canals indicated in one genus, Saurerpeton; limbs well developed, with well-developed digits and ungual phalanges claw-like; body usually slender, broad in Saurerpeton; the ribs broad and heavy; the vertebrae relatively stout; ventral armature highly developed, reaching the height of specialization among the Microsauria ; scutellse consisting of rods, plates, or stout bristles. The family is represented by 5 genera: Sauropleura, Saurerpeton, Ctenerpeton, Leptophr actus, and Eurythorax, the association of the last two genera being provisional. The genera may be distinguished as follows: I. Pectoral elements sculptured, clavicles triangular, interclavicle diamond-shaped, ventral scutellae rods, arranged en chevron with anterior angle Sauropleura II. Pectoral elements slightly sculptured, cranium broad, obtuse and sculptured, ventral armature broad imbricated plates extending on to the throat Saurerpeton III. Limbs, skull, arches and dorsal vertebra? unknown, caudal vertebrae with fan-shaped neural and haemal spines which may indicate relationship with Ptyonius and CEstocephalus, but in those genera the ventral armature is weakly developed; ventral scutelte curved rod-like plates arranged en chevron with anterior angle, marked in abdominal region by distinct rounded pits Ctenerpeton IV. Known only from fragments of the skull, teeth large and fluted; association in family provisional. . Leplophractus V. Known only from a single interclavicle of peculiar form which resembles that of Saurerpeton; association in the family provisional Eurythorax The members of this family are confined to the deposits of the Coal Measures at Linton, Ohio. Ctenerpeton, and possibly Sauropleura, were highly developed swimmers, but the strength of the limbs as exhibited, especially by Sauropleura and Saurerpeton, indicates that they had not entirely forsaken the land. Genus SAUROPLEURA Cope, 1868. Newberry, Proc. Phila. Acad. Nat. Sci., vm, p. 98, 1856 {Pygopterus scutellalus). Cope, Trans. Am. Phil. Soc, p. 22, 1869. Cope, Geol. .Surv. Ohio, it, pt. 11, p. 402, 1875. Hay, Bull. U. S. Geol. Surv., No. 179, p. 419, 1902. Type: Sauropleura scutellata Newberry. Vertebrae and ribs well developed; limbs 4, rather large; 5 digits in the fore- foot; carpus cartilaginous. Ventral armature of closely arranged rhomboidal scuta, arranged in lines, which are closely placed in chevrons, with the angle anterior. 155 I56 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. In none of the species of this genus have the usual 3 thoracic shields (clavicles and interclavicles) been observed. The abdominal scuta?, on the other hand, are much like those of Saurerpeton, being, however, smaller. The species formerly described by Cope in the genus Colosteus are included in Sauropleura, where they find their closest allies. There are 7 species belonging in this genus: Sauropleura digitata Cope, 5. new- berryi Cope, S. foveata Cope {Colosteus), S. scutellata Newberry {Colosteus) (type of genus), 5. pauciradiata Cope {Colosteus), S. longidentata Moodie, S. enchodus Cope {Anisodexis) . The species described by Cope as Sauropleura latithorax is regarded as belonging to a distinct genus, Saurerpeton. Sauropleura scutellata Newberry. Newberry, Proc. Phila. Acad. Nat. Sci., p. 98, 1856 (Pygoplerus sculellalus). Cope, Proc. Phila. Acad. Nat. Sci., p. 215, 1868. Cope, Trans. Am. Phil. Soc, p. 22, 1869. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 402, 1875. Type: Specimen No. 8669 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. (Plate 14, fig. 3.) This species was first described by Newberry as a fish belonging to the genus Pygopterus. Cope later placed it under the genus Colosteus and clearly showed its amphibian characters. The genus Colosteus was, however, based on a misconcep- tion. Cope in 1897 and Hay (317) referred the species to Sauropleura, where it is retained. The species is represented by a single individual preserved on a block of coal from Linton, and is also indicated by an interclavicle and its obverse; this element is of larger size than that of the type and was referred by Cope to Colosteus pauciradiatus. The characters of the plate are, however, so identical with those of the interclavicle in the type specimen that it is unhesitatingly referred to the pres- ent form. The type specimen consists of the supero-lateral view of a crushed cranium with the anterior part of the body, exhibiting the interclavicle and the ventral scutella- tion. No limbs have been observed in this species. The mandibles are crushed across the cranium in such a way as to obscure its structure. The boundary of the left orbit is doubtfully determined as being a little back of the median line of the skull. There are small teeth present on the mandibles, but their number can not be determined. The cranial elements are sculptured with radiating grooves and ridges, but these are weakly developed. The snout is broad and but little narrower than the base of the skull. (Plate 21, fig. 5.) The interclavicle, somewhat displaced, is the only element of the pectoral girdle preserved. It is peculiar in the possession of a backward extension which shows a beveled edge. The plate is ornamented by radiating grooves and ridges which are strongly developed. The larger specimen of an interclavicle shows the same characters as the one described, and it differs only in being about twice as large. There are no traces of limbs. The ventral armature of the body is rather weak as compared to that of Sau- ropleura pauciradiatus, but it is still composed of closely packed scutes arranged en chevron. The character of the ventral armature and the sculpturing of the inter- HIE MICROSAURIAN FAMILY SAUROPLEURIDyE. 157 clavicle are taken as the principal specific characters. From the other species of the genvis the present form differs in its more slender elements of the ventral armor and in the form of the skull. The ribs are not clearly defined. Measurements. mm. mm. Median length of skull 70 Length of mandibular tooth 1.5 Posterior width of skull 40 length of entire specimen 150 Anterior width of skull 20 Width across belly, maximum 47 Length of jaw 50 Length of interclavicle 27 Anterior width of jaw 7 Width of interclavicle 13 Posterior width of jaw 12 Width across posterior extension 4 Sauropleura digitata Cope. Cope, Proc. Phila. Acad. Nat. Sci., p. 216, 1868. Cope, Trans. Am. Phil. Soc., xiv, p. 15, 1869. Cope, Geol. Surv. Ohio, n, pt. 11, p. 403, pi. xxxvii, fig. 1, 1875. Type: Specimen No. 8004 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. (Plate 20, fig. 4.) This species is represented by a single individual which is quite distinct from other members of the genus. The specimen has been spread over a surface of the coal, and exhibits ventral armature, dorsal region with ribs, and anterior and pos- terior limbs. Skull and caudal region not present. The ventral armature is arranged in parallel lines directed obliquely forwards and continuous on the median line, forming there a chevron. The individual ele- ments are oat-shaped, and acuminate at both ends. They are moderately imbri- cate in an antero-posterior section. On the pectoral region between the fore limbs the series of scutellae assume different directions, forming chevrons directed back- wards, and forming with those of the belly a complete X. The humerus, ulna, and radius are rather stout, and of a size relative to the body, as in common types of existing lizards; the ulna and radius separate. The carpus is cartilaginous; the digits are 5 well-developed fingers having phalanges in the following numbers, commencing on the inside: 3, 4, 5, 6, 5. The last phalanx of the second is obscured, and it is not positive that the number is as given; it is more probable that it was 4 than 3. The outer toe was more slender than the others; the distal phalanges of all the toes are stout, as in modern Caudata. The ribs are long and curved as in reptiles, and judging by their distances the vertebrae are short ; the latter are not well-defined, but there is no indication of prom- inent spines of any kind. The pelvic bones and portions of the hind limbs are pres- ent, but so obscured and confused as not to be easily made out. Enough remains to show that the hind limbs were longer than the fore. Thirteen ribs on one side and twelve on the other are preserved, with short ribs in the sacral region. The specimen is very indistinct and it is difficult for one to be sure of all the characters described by Cope. Measurements of the Type of Sauropleura digitata Cope. mm. mm. length of specimen 115 Length of radius 10 Greatest width 80 Length of ulna 9 Diameter of ventral scute .75 Length of metacarpal 4 Length of rib 15 ' Length of fourth digit of hand 22 Width of rib I Length of interclavicle 20 Length of humerus 20 Width of interclavicle 1 1 I58 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The interclavicle is diamond-shaped, with surface punctate and edges radiately grooved at a distance of 1.5 mm. from the edge. The hand on the right side of the specimen contains 3, 3, 3, 6, 3 phalanges. The vertebras are all imperfectly preserved. Other specimens of this species are 2567 (48), 8376 G, 8704 G, American Museum of Natural History. Coal Measures of Linton, Ohio. Collected by Dr. J. S. New- berry. Sauropleura newberryi Cope. Cope, Geol. Surv. Ohio, II, pt. II, p. 404, pi. xxxvii, figs. 2 and 3; pi. xli, fig. 5, 1875. Type: Specimen No. 8612 G, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The type specimen of the species exhibits a portion of the posterior part of the skull, a considerable part of the body, with the fore limbs and abdominal scutellae. No vertebrae can be definitely discovered and the ribs are not distinctly visible. The cranial fragment is the upper surface of the tabulare and adjacent elements, and a broad band of the posterior parts of these is seen to be smooth, and is pre- ceded by a slightly roughened surface. The abdominal scutae are diamond-shaped, and are thin and light, not massive, as in S. scutellata, and are sometimes marked with a median longitudinal keel. The fore limb is large, especially the humerus, which is much dilated distally, and has a strong crest on the outer side from near the proximal end. The ulna and radius are much shorter, and more dilated prox- imally than distally; they are well separated. No phalanges are preserved.. The species is represented by other specimens, all of which are unsatisfactory in determining the structure of the form. A skull, apparently complete, is crushed out flat, so that little can be said of structure. The teeth are rather long, straight, acute, and striate at the base. The orbits are long and narrowed in front. Measurements of Sauropleura newberryi Cope. (.No. 8612 G, and two unnumbered specimens, American Museum of Natural History.) mm. mm. Length of humerus 35 Width at anterior angle of orbits 40 Proximal width of humerus 8 lnterorbital space 13 Distal width of humerus 14 Width of orbit 13 Length of ulna 19 Length of orbit 27 Proximal width of ulna 8 Length of a maxillary tooth 4 Posterior width of skull 67 Diameter of same at base 2 Sauropleura pauciradiata Cope. Cope, Trans. Am. Phil. Soc., xv, p. 275, 1874. Cope, Geol. Sun'. Ohio, n, pt. 11, p. 408, pi. xl, figs. 1-2, 1875. Type: Specimen No. 8671 G, and obverse, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. The species was founded on a median and a lateral plate of 2 individuals. The clavicle is here associated with S. scutellata and the interclavicle thus remains as the type specimen. There are other remains associated by Cope with S. scutellata which are here shown to have closer affinities with the S. pauciradiata. The clavicle is a right-angled triangle, the inner and thin edge concave poste- riorly, the posterior convex. The ridges and grooves are well developed on the specimen. The visceral side of the element is smooth. THE MICROSAURIAN FAMILY SAUROPLEURID^E. 159 ,, i.& '/ v->: &&tfgU^ -^ 1. Photograph of type specimen of Erpetosaurus (Tiit/itanits) nu/inlns Cope, from the Coal Measures of Linton, Ohio. X 1.3. Original in American Museum of Natural History. 2. Photograph of type specimen of Erpetosaurus tabulalus Cope, from the Coal Measures of Linton, Ohio. X 2. Original in the Zoological Collections of Columbia University. 3. Photograph of the impression of Stegops divaricate Cope, from the Coal Measures of Linton, Ohio. X 2. The specimen figured is in the American Museum of Natural History. Its obverse is in the collections at Walker Museum, I'niversitv of Chicago. 4. Type and only known specimen of .Vi< rrrpr/on caiufatittH Moodie, a branchiosaur from the Coal Measures shales of MilOB Creek, Illinois. X 2. Original in collections at Walker Museum, University of Chicago, THE TEMNOSPONDYLOUS AMPHIBIA. l8l Eryops sp. indet. Case, 1908. Case, Annals Carnegie Mus., iv,p. 234, pi. 59, 1908. A dorsal vertebra is very probably from this genus. The specimen consists of a nearly perfect vertebra, lacking only the anterior zygapophysis and the upper por- tion of the neural spine (plate i8, fig. 2). It shows no character that would war- rant its separation from the genus, and indicates a medium-sized individual. The zygapophyses have clean-cut articular faces. The pleurocentra are thickened above, with just well-defined articular faces, which were applied to faces on the neural arch posterior to the origin of the transverse process. The intercentrum is of the familiar half moon-shape, thick and heavy below, and thinner toward the extremities; the anterior edge is marked near the top by the indentation found on the intercentra of Eryops. Height of the vertebra from the middle of the lower face of the intercentrum to the middle of the neural canal, 0.035 m- ; width of intercentrum 0.026 m. The second recognizable specimen is a neural spine from the caudal series. This is without question a portion of the skeleton of an Eryops. Similar spines were described by Cope as Eryops (Epicordylus) erythrolithicus, but later discoveries seem to show that similar characters occur in other species of the genus as well. The apex of the spine is bifurcate; the space between extremities is concave and per- fectly smooth ; below the sides of the spine are rather rugose and marked with ridges. The lower portion of the spine is elongated anteroposteriorly and the edges are marked with sharp, double ridges. Three ribs also belong, in all probability, to the genus Eryops. The head of each rib is broad and the articular edge is divided between two faces which meet at an angle somewhat greater than a right angle; the two faces are continuous. The shaft is somewhat flattened and in the undistorted specimens is gently curved. The length of the largest rib is about 0.07 m. Other than these specimens there are several small intercentra (94) and the neural spine of a caudal vertebra from some undetermined amphibian. Family MACRERPETIDjE Moodie, 1909. Moodie, Bull. Am. Mus. Nat. Hist., xxvi, art. xxv, p. 354, pi. lix, fig. I, 1909. It has seemed necessary to propose a new family for the reception of the single species Macrerpeton (Tuditanus) huxleyi Cope. The characters exhibited by this species are so different from those offered by other members of the Carboniferous Microsauria that it is clearly distinct. In its cranial characters and the position of the orbits it approaches most nearly to Eryops megacephalus Cope from the Per- mian of Texas. In some of its characters the present form shows a similarity to Dasyceps bucklandi Lloyd (324), from the Permian of Kenilworth, England; more especially is this similarity found in the form of the skull, the size and shape of the teeth, and the posterior position of the orbits, and their wide removal from the border of the skull. Only a fragment of the skull has hitherto been known, but re- peated study of this fragment (123) has disclosed the wide diversity (462) of its characters. An almost complete skull, described below, substantiates the charac- 1 82 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. ters based on the fragment. Another species is here added to the genus, based on a portion of a mandible and a portion of the skull. The family, Macrerpetidae, may be defined (465) as follows : Skull larger than in any other known microsaurian, unless Baphetes proves to be microsaurian; cranial elements sculptured with pits and coarse grooves; lacrimal element present, teeth large, curved inwards and fluted; mandible heavy; orbits located far back on the skull and near the median line so that the interorbital space is about half the space from the outer edge of the orbit to the border of the skull, thus approaching the condition known in Eryops; the ribs (?) are strong, heavy, and curved, with an incipient tubercle. Genus MACRERPETON Moodie, 1909. Moodie, Jour. Geol., xvn, No. I, pp. 72-74, fig. 17, 1909. Type: Macrerpeton huxleyi Cope. The genus Macrerpeton was proposed for the reception of the amphibian species described by Cope as Tuditanus huxleyi (123). This form he placed provisionally in genus Tuditanus, since it seemed to present the same type of sculpturing of the cranial elements similar to that found in T. radiatus Cope. But this species has been removed from Tuditanus and placed in a new genus, Erpetosaurus (462). A close study of the type specimen of Tuditanus huxleyi Cope shows (465) great variation and distinction from any of the species described from Linton, Ohio, or indeed from any Carboniferous form thus far known. The specimen represents the left side of the face, and the characters exhibited by the fragment are supported by more complete material (No. 2933, Am. Mus. Nat. Hist.). The skull shows a close approach to the higher labyrinthodonts in its shape. The orbits are far removed from the border of the skull. The arrangement of the bones of the skull resembles that of Capitosaurus from the Keuper of Europe. The jaw is for the most part slender, with a pronounced downward inflection at the coro- noidal part. The teeth are heavy and strong and are curved backwards. They have the strong longitudinal fluting which is characteristic of the labyrinthodonts. Another character which is distinctive is the pattern of cranial sculpture. This consists of inosculating pits and grooves of a coarse character and compares favor- ably with the sculpturing of Triassic labyrinthodonts. If Macrerpeton really rep- resents a labyrinthodont form of Amphibia it is the oldest of the known Labyrintho- dontidae, since it seems probable that the Eosaurus vertebrae came from a higher horizon. Macrepeton huxleyi Cope, 1874. Cope, Trans. Am. Phil. Soc, xv, p. 274, 1874. Cope, Geol. Surv. Ohio, 11, pt. 11, p. 397, pi. xxxiv, fig. 2, 1875. Lesley, Dictionary of Fossils, p. 1237, 1890. Moodie, Jour. Geol., xvn, No. 1, p. 72, fig. 17, 1909. Moodie, Bull. Am. Mus. Nat. Hist., xxvi, art. xxv, p. 354, pi. lix, fig. 1, 1909. Type: Specimen No. 119, American Museum of Natural History. Collection of Dr. J. S. Newberry. (Plate 26, fig. 2.) Horizon and locality: Linton, Ohio, Coal Measures. The first part of the following account of the species, Macrerpeton huxleyi Cope, is a quotation of Cope's description (123) of the type specimen, and the second part deals with the description of the new material. Cope says the species is — MOOOII Pl.AT£ 26 2 1. Photograph of type specimen of Erpetosaurus tuberculatus Moodie, from the Ohio Coal Measures. X 1. Original in American Museum of Natural History. Z. Photograph of type of Macrerpeton huxleyi Cope, from the Ohio Coal Measures. X 1. Original in American Museum of Natural History. THE TEMNOSPONDYLOUS AMPHIBIA. 1 83 " Represented by a considerable portion of the face and muzzle of a single individual. A portion of the left mandible, supporting three teeth, remains in place, and almost the entire boundary of the right orbit is preserved. ' The fragment indicates a much larger species than any other referred to the genus, and, next to the Le ptophractus obsoletus, the largest of the Batrachians of the Ohio Coal Measures. Without more complete remains, it is not easy to determine its generic relations finally. " The form of the head is probably elongate, and the muzzle neither very obtuse nor elongate. The orbit is rather small, and near the middle of the length of the specimen, which is, however, incomplete at both ends. The sculpture of the surface of the head pos- terior to the orbits, as well as round their borders and for some distance in front of them, consists of a rather coarse pitting. On the middle line, between the orbits and on the muz- zle, the intervals become narrower, and are confluent into transverse ridges or a delicate reticulation. The surface of the mandible displays a coarse reticulation. " The teeth are stoutly conic, and with delicately striate grooved cementum. They are slightly recurved. " This species differs from the T. radiatus and T. obtusus in the absence of the area into which the sculpture is thrown. " Longitudinal diameter of orbit, 19 mm.; length of alveolar border supporting three teeth, 13 mm. ; diameter of base of tooth, 3 mm. ; eight pits in 10 mm. " Dedicated to Professor T. H. Huxley, facile princeps among English systematists, and an important contributor to the knowledge of the extinct Batrachia." The following discussion of the cranial elements, based on the writer's studies (462, 465) of the type, may be appended to Professor Cope's original description. The sutures bounding a few of the elements have been made out in part. The prefrontal element seems well assured. It lies well in front of the orbit, much as in the skull of Capitosaurus from the Keuper of Europe. The lacrimal is, apparently, a very large bone, though its entire extent is not assured. The maxilla is a long, narrow element on the border of the skull. The suture separating this from the lacrimal and jugal is quite clear. The teeth which the maxilla undoubtedly bore are hidden by the remains of the mandible, which lies partly on the edge of the skull. The jugal is a very large element and its boundaries seem well assured. Its size and relations recall the condition in Capitosaurus. It forms a part of the external boundary of the orbit. The lateral suture of the post- orbital is evident and is, as shown in the figure, somewhat curved. The remaining elements preserved on the fragmentary skull can not be accurately determined, though their probable position is indicated in plate 30, fig. 2, the lettering being based on the arrangement of these elements in Capitosaurus. The lower jaw is poorly preserved, but what remains shows evidence of being sculptured somewhat after the manner of the cranial elements. It bore strong recurved teeth which are longitudinally striate. Measurements of the Type Specimen of Macrerpeton huxleyi Cope. mm. mm. Length of portion preserved 120 Length of jaw, as preserved 75 Maximum width of specimen 58 Width of jaw at widest part 11 Length of orbit 20 Length of longest tooth 8 Width of orbit 14 Width of tooth at base 4 1 84 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. DESCRIPTION OF ADDITIONAL MATERIAL OF MACRERPETON HUXLEYI. The additional material of this species which has come to hand consists of an almost complete skull (American Museum No. 2933, two portions); another frag- mentary skull (American Museum No. 8572 G and 8532 G); a portion of an interclavicle (American Museum No. 8006) ; two incomplete vertebrae (American Museum No. 8007) ; and another fragmentary element possibly representing a scapula of this species (American Museum No. 8008). The skull has essentially the shape outlined (462) from a study of the fragmen- tary type specimen. The muzzle was drawn slightly too broad, but otherwise the restoration is fairly accurate. The specimen is distorted and imperfect, but enough is preserved to give a good idea of the shape and something of the structure of the skull. A portion of the obverse is preserved. The back part of the skull is broken, so that the occiput can not be studied. The length of the skull is one and two-fifths the greatest breadth (across the orbits). The cranial elements are deeply marked with pits and short, shallow grooves. On the left mandible these pits are in a very distinct row, the operculo- mandibular lateral line. ij £?*'** Macrerpeton deani new species. Type : Specimen No. 2934, American Museum of Natural History. Horizon and locality: Linton, Ohio, Coal Measures. (Plate 21, figs. 1,2.) The material for this species consists of the posterior half of the left mandible and a portion of the right antero-lateral surface of the skull, both incomplete. The reasons for regarding the species as distinct are the large size of the specimens and the manner of the sculpture, as well as the shape of the posterior end of the mandible. The present species is the largest amphibian of the Linton, Ohio, Coal Measures, exceeding in skull length that of Macrer- peton huxleyi by twice. The largest skull of Macrerpeton huxleyi which has so far come under my notice is 120 mm. in median length. There are 3 skulls of this species known, all of approxi- mately the same size. The skull of Macrerpeton deani must have reached or exceeded a foot in median length. The only species with which it can at all be com- pared are Eobaphetes kansensis Moodie and Baphetes planiceps Owen, but it is clearly distinct from all other genera of Linton Amphibia. It is possible that when better known Macrerpeton, Eobaphetes, Baphetes, Erpetosaurus, and possibly Dend- rerpeton will form a natural group of early labyrinthodont-like Amphibia. The mandible is similar in structure to that of the labyrinthodonts, with the elements marked by radiate flutings. I can detect no evidences of a lateral-line Fig. 40. — Mandible of Macrerpeton deani new species, from Linton, Ohio. X 0.75. THE TEMNOSPONDYLOUS AMPHIBIA. 1 85 canal, such as is clearly marked in Macrerpeton huxleyi Cope by a series of rounded pits, occupying the usual position of the operculo-mandibular lateral-line canal The teeth, of which 6 are preserved, are minutely striate, with smooth apices. They are dissimilar in size, showing a variation of 2 or 3 mm. in length. The sculpture is a coarse fluting, with no indications of the sharply marked pits of Macrerpeton huxleyi Cope. The fragment of a skull preserved shows characters of the sculpture which are identical with those of the mandible. The bones are so crushed that it is impossible to tell the limits of the elements. I believe a portion of one orbit is represented on one corner of the block. The cranium appears to have been broad, and the fragment preserved, which is only about one-sixth of the skull, is larger than the entire cranium and mandibles of Macrerpeton huxleyi. The specific distinctness of the form can not be doubted, although it is a mat- ter of regret that it is founded on so small a portion of the osteology of the animal. The species is proposed in honor of Dr. Bashford Dean, to whom I am greatly indebted for many kindnesses during the past 5 years in connection with my studies on Carboniferous Amphibia, particularly in the loan of the entire Newberry col- lection of Linton, Ohio, Amphibia. Measurements of the Type of Macrerpeton deani Moodie. mm. Length of portion preserved 115 Greatest width 50 Ix'nglh of tooth 9 Width at base 4 Length of angular 95 Diameter of angular 25 Measurements of specimen No. 8535 G, American Museum of Natural History, associated with the above in the type description: Length of preserved portion 140 Diameter of orbit 22 Family ANTHRACOSAURIDjE Cope, 1875. Cope, Bull. U. S. Nat. Mus., I, p. 10, 1875. Lydekker, R., 1890, Cat. Fossil Reptilia and Amphibia, p. 157. Skull usually triangular and more or less angulated, with the cranial sculpture well marked, the occipital condyles ossified, and the palatine foramina very small and placed far back; dentine of the teeth more or less complexly plicated. A ventral armor of elongated dermal scutes, and probably a sclerotic ring. Bodies of vertebras fully ossified in the adult ; intercentra present or absent. According to Atthey's figure (11) of the skull of the type genus, the palatine bears teeth which are situated immediately on the inner side of the maxilla, as in Masto- donsaurus (242). In the typical forms there is no postarticular process to the mandible. The North American species of this family are: Eosaurus acadianus Marsh, Eobaphetes kansensis Moodie, Dendrerpeton acadianum Owen, Dendrerpeton oweni Dawson, Platystegos loricatum Dawson, Baphetes planiceps Owen, Baphetes minor Dawson. There is but little assurance that any of these species belong in this family. They are put there provisionally, pending future discoveries. Huxley suggests the relationship of Eosaurus and Anthracosaur.us (Quart. Jour. Geol. Soc, xix, 1863, p. 65; Scientific Memoirs, 11, p. 566). 1 86 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Genus BAPHETES Owen, 1854. Owen, Quart. Jour. Geol. Soc. London, x, p. 207, pi. ix, 1854. DAWSON, Air-breathers of the Coal Period, pp. 10-16, pi. ii, 1863. Type : Baphetes planiceps Owen. Known only from an incomplete skull, which is large, broader than long; squa- mosals prolonged into obtuse horns. Teeth rather large, heterodont, arranged in a single row. Orbits placed well forward, frontals small, surface bones sculptured. Baphetes planiceps Owen. Owen, Quart. Jour. Geol. Soc. London, x, p. 207, pi. ix, 1854. Dawson, Air-breathers of the Coal Period, pp. 10-16, pi. ii, 1863. Type : Specimen in the British Museum of Natural History. Horizon and locality: Near Pictou, Nova Scotia (Coal Measures). The parts preserved include the premaxillaries, nasals, and portions of the frontal, prefrontal, and maxillary bones. The fossil is embedded in a mass of Pictou Coal from Nova Scotia and consists of the anterior extremity of the cranium (plate 22, fig. 6) and with the exterior surface of the bone embedded in the matrix, and its substance, for the most part, reduced to a thin layer by abrasion of the exposed inner layer. It displays accurately the contour of the fore part of the upper jaw, which was broad, obtuse, and rounded. The premaxillaries, which show some obscure traces of a symphysial suture at the median line, anterior to the nasal or naso-palatine vacuities, extend outwards, on each side, for an extent of 2.5 lines and there join the maxillaries. Traces of round alveoli for teeth, some of which are 2 lines in diameter, are visible on the alveolar border of the premaxillaries. The alveolar border is continued by the max- illary bone for an extent of 4.5 inches beyond the premaxillary border, and this border shows still more distinct traces of alveoli, of a circular form, about a line in diameter and rather close set in a single series. The fore part of the orbit is very unequivocally displayed, the smooth inner or under surface of the bone forming that part being entire; and this shows the fore part of the orbit to be formed, partly by the maxillary, partly by the lacrimal or prefrontal bone in close sutural union therewith, a structure which does not exist in any recent or fossil fish with a dentig- erous superior maxillary bone. Where the substance of the bone has been detached so far as to expose the external layer in contact with the coal, as, e.g., on the frontal and part of the prefrontal bones, the external surface of those bones is shown to have been impressed by subhemispherical or elliptical pits, from 1 line to 1.5 lines in diameter, and with intervals of half of that extent. This coarsely pitted character agrees with that presented by the other surface of the similarly broad and flat cranium of the labyrinthodonts. From the characters above specified, therefore, I conclude that this fossil is the fore part of the skull of an extinct family of the labyrinthodonts. It agrees with them in the number, size, and disposition of teeth ; in the proportions and mode of connection of the premaxillaries, maxillaries, nasals, prefrontals, and frontals, and in the resultant peculiarly broad and depressed character of the skull. The traces of THE TEMNOSPONDYLOUS AMPHIBIA. 1 87 the nostrils are less definite and satisfactory than the remains of the orbits, but the latter appear to be decisive against the piscine nature of the fossil. The fossil also presents the same well-marked external sculpturing as in the labyrinthodonts; and among the genera that have been established in that family, the form of the end of the muzzle, or upper jaw, in the Pictou coal specimen best accords with that in the Capitosaurus and Metopias of von Meyer and Burmeister (80). Measurements of the Skull of Baphetes planiceps Owen. (Type in the British Museum of Natural History, London.) mm. mm. Approximate median length of skull 136 Width of skull across orbits 97 Width of skull across base of horns 150 Diameter of orbit 21 Estimated width across tips of horns 186 Diameter of large tooth alveolus 7 Width of horn at base 31 Diameter of small tooth alveolus 2 Estimated length of horn from base 80 Pictou Coal, near Pictou, Nova Scotia, Canada, collected by Dr. J. William Dawson, 1850, and presented by him to the Geological Society of London. Baphetes minor Dawson. Dawson, Canadian Nat. and Jour. Sci., n. s., 1870, v, pp. 98, 99. Type: Specimen in the Peter Redpath Museum, McGill University. Horizon and locality: Coal formation of Nova Scotia. The species was based on a lower jaw of an amphibian, of which a cast had occurred in the coarse sandstone of the coal formation between Ragged Reef and the Joggins Coal Mine. It measured 6 inches in length; its surface was marked on the lower and posterior part with a network of ridges inclosing rounded depressions. The anterior part of the jaw contained about 16 teeth, some of which remained in the matrix. These were stout, conical and blunt, with large pulp cavities, and about 32 longitudinal striae, corresponding to the folds of the dentine. Dawson states that this jaw resembles most closely those of Baphetes and Dendrerpeton, but more especially the former. He regarded it as distinct from Baphetes planiceps, and proposed for it the name Baphetes minor. Eosaurus acadianus Marsh. Marsh, Am. Jour. Sci. (2), xxxiv, pp. 1-16, pis. i, ii, 1862. Agassiz, Am. Jour. Sci., xxxm, p. 138, 1862. Marsh, Quart. Jour. Geol. Soc, xix, pp. 52-56. Hay, Cat. Fossil Vertebrates (Bull. U. S. Geol. Surv. No. 179, p. 421, 1902). Type: Specimen No. 1648, Yale University Museum. Horizon and locality: South Joggins, Nova Scotia (Coal Measures). The genus and species are founded on two vertebral bodies of the stereospondy- lous type from the Coal Measures of the South Joggins, Nova Scotia. Marsh's description (404) is as follows: "The general form of the vertebrae is cylindrical, but their sides are compressed ob- liquely, which gives to the contour of the centra a subhexagonal appearance. They are much flattened in the direction of the anteroposterior diameter, which has to the transverse i88 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. diameter the proportion of i to 3. Both the articular terminal facets are deeply and equally concave; but from the center to the margin the surfaces are convex, and this convexity is greatest near the center. * * * The cavities for the reception of the intervertebral matter begin immediately from the margin, and are considerably deeper than the corresponding parts of the Ichthyosaurus, indicating a greater degree of flexibility in the vertebral column. The margins of the vertebras are some.what raised, as if they had yielded to a forcible com- pression applied longitudinally ; and hence the lateral surfaces of the centers are concave in an anteroposterior direction. This concavity is greater in the upper half of the vertebra and was undoubtedly more marked originally than at present, since the appearance of the margins indicates considerable abrasion. The non-articular surfaces of the centra are smooth and regular; and the external fibres of the osseous tissue are singularly reticulated. (Mi) \v\ ♦*c F Fig. 41. — Nova Scotian Amphibia. A. Oblique lateral view of vertebra? of Eosaurus acadianus Marsh, a, pits for articulation of neuropophy ses ; b, rudimentary transverse process on right lateral surface of centrum. B. Oblique view of vertebra?, a, pits for articulation of neuropophy ses ; 6, rudimentary process on lateral surface. C. Posterior view of nearly perfect centrum. D. Transverse section of same vertebra, showing deep concavities of articular terminal facets. E and F. Microscopic sections near surface, showing lacuna? arranged around an Haversian canal. Magnified 200 diameters. (All figures after Marsh. X 0.75.) ' The neuropophyses are not anchylosed to the centrum, as in the mammalia, nor con- nected to it by sutures, as in the crocodiles ; but their union with the vertebra is indicated by two pits, which served for their articulating surfaces. These depressions are situated on the superior surface of the centrum intermediate between the anterior and posterior mar- gins of the extremities. They are circular in form and sink directly into the body of the ver- tebra; instead of being elongated longitudinally and raised by ridges, as in Ichthyosaurus. The pits are about a line in depth, and in the more perfect of the fossils are not in their origi- nal position. The floor of the spinal canal is narrow, being but 5 lines in breadth. A rudi- mentary transverse process, or exogenous tubercle, is sent off from each lateral surface of the centrum, at points equidistant from the extremities of the vertical diameter. Their THE TEMNOSPONDYLOUS AMPHIBIA. 1 89 position is near the margin of the anterior articular surface, and the edges of these para- pophyses make the transverse diameter of this extremity somewhat greater than that of the corresponding posterior facet." Measurements of the More Perfect Vertehra of Eosai ms acadianvs (after Marsh). mm. Transverse diameter of centrum on anterior surface 59 Same on posterior surface 57 Same including the parapophyses 63 Vertical diameter of anterior surface P4 Anteroposterior diameter on superior surface 21 Same on inferior surface ny Same between centers of articular facets 2.5 Length of pits for articulation of neural arch 7 Breadth of same 7 Depth of same 2 Distance between centers of same II Same at centers of parapophyses 36 Collected by 0. C. Marsh at the South Joggins, Nova Scotia. Marsh regarded (404) the vertebrae as representing a new type of ichthyosaurian (2), but there can be no doubt that the vertebras belong to some form of the Amphibia, since the description applies equally well to them. In this connection mention must be made of a large rib from the Linton beds preserved in the U. S. National Museum. Only the proximal third of the rib is preserved, but it represents some large form of the Stegocephala. The rib is strongly curved backward, is heavy, and has an incipient tubercle. A cross-section shows that a longitudinal groove occupies the median line on the exposed surface of the rib. This may, however, be due to compression and thus indicate that the rib was hollow. The rib as pre- served measures: length, 102 mm.; maximum width, 22 mm.; minimum width, 14 mm. (Nos. 4490, 4489, U. S. National Museum.) Genus E0BAPHETES new name. Type: Eobaphetes kansensis MoOdie. The new name is proposed to replace the generic term Erpetosuchus used for the species E. kansensis described by the writer (Proc. U. S. Nat. Mus., 39, p. 491, 191 1), and which later was found to be preoccupied by Newton (Phil. Trans. Roy. Soc. London, 185, p. 573, 1894 b). The genus is very readily distinguished by two prominent characters — the short, uniform dentition and the presence of two elongate, oval, internal mandibular fora- mina on the inner side of the jaw. The genus may be further distinguished by the great depth of the posterior portion of the jaw and the slender anterior part, as well as by the ornamentation, which is typically the rough tuberculated labyrinthodont sculpture on the anterior end of the mandible. This changes gradually to longi- tudinal grooves and ridges of a rather small size on the posterior portion, a very unusual arrangement for a labyrinthodont. These characters are sustained by those of the skull fragment, in which the den- tition is uniform and the sculpture very similar to that of the mandible. The ribs are long, curved, and solid, as in other labyrinthodonts. I90 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. The internal surface of the mandible shows much similarity to that of the croco- diles and alligators of the present day. The resemblance is not due to homology of structures, but must be regarded as a parallel development of similar characters. Eobaphetes kansensis Moodie. Moodie, Proc. U. S. Nat. Mus., 39, pp. 491-494, figs. 1 to 3, 191 1 (Erpetosuchus). Type: Specimen No. 6699, U. S. National Museum. Horizon and locality: Coal Measures of Washington County, Kansas. The species is represented in the collections of the U. S. National Museum by a fragment of the skull, with portions of two ribs (Cat. No. 6699, Vert. Pal. U. S. Nat. Mus.) and the larger part of the left ramus of the mandible (Cat. No. 6680, Vert. Pal. U. S. Nat. Mus.). The mandible was preserved in a large block of coal which contained the impression of the back portion of the mandible from which the bone had been weathered. It was possible to remove the bone and make a plaster cast of the impression. This shows in a very satisfactory manner all of the char- acters of the external surface. Skull. — Only a portion of the left maxilla, with 14 teeth, and a part of the nasal are preserved. The skull seems to have been laterally crushed and the right side of the skull has been crushed flat under the left. It has not seemed feasible to remove the skull from the matrix. The teeth are uniform, rather short, bluntly conical, curved backward, and coarsely striate. They are somewhat crowded, the bases being separated from each other by only a fraction of a millimeter. The maxilla and portion of the nasal are coarsely sculptured with elongate pits and ridges. A portion of the infraorbital lateral-line canal is preserved. It is simply a rounded groove with three short branches. It lies near the middle of the maxilla. Mandible. — It has been possible to study both sides of the mandible. The left ramus was preserved in the coal, with its inner face exposed. This face is broken by two large oval openings, the internal mandibular foramina. This is the term used by Reynolds for the openings on the inner surface of the alligator jaw. So far as I can ascertain, no other known labyrinthodont mandible displays this char- acter in such a marked degree. Dr. Branson has figured in Anaschisma browiri Branson from the Triassic (49) of Wyoming the inner surface of the left ramus, on which there are likewise two openings but differently situated. A similarity be- tween the two mandibles is observed in that the suture separating the prearticular and angular touches the posterior edge of the posterior foramen. Several of the sutures are well preserved and they have been indicated in the drawing (fig. 42). The pillar separating the two foramina is cut by the suture separating the angular and prearticular very much as in Anaschisma, with the difference that in the latter form the angular and prearticular are not approximated. I believe I detect the suture as represented separating the anterior end of the angular from the dentary and splenial. I am assured of the portion near the anterior fora- men and also of the part near the tip of the ramus. This shows the angular to be a very elongate element, running very nearly the entire length of the mandible, much THE TEMNOSPONDYLOUS AMPHIBIA. 191 as in Anaschisma and other labyrinthodont genera. The splenial is a small, slender element located farther forward, where it has been shoved by the large-sized internal mandibular foramina. The prearticular is a rather long, broad element, of which only a portion is preserved. I am not sure as to the location of the suture for the dentary, unless it is represented by the line bounding the roughened area near the teeth. If this is true, the dentary is a large element, since it extends well down upon the outer side of the jaw. The dentary possesses evidences of 26 teeth, a few of which are completely preserved. Most of them are, however, represented either by bases or by impressions in the coal. The teeth are very similar to those of the maxilla, though slightly larger. The characters given for the maxillary teeth will suffice for those of the dentary. * 1 \ - Fig. 42. A. Outer view of mandible of Eobaphetes kansensis Moodie, from the Coal Measures of Washington County, Kansas. Original in U. S. National Museum. X 0.33. Cat. No. 6680. o, angular; particular; d, dentary. B. Portion of skull of Eobaphetes kansensis Moodie, from the Coal Meas- ures of Washington County, Kansas. Original in U. S. National Museum. Cat. No. 6699. X 0.33. Lateral-line canal represented by heavy broken line, n, nasal; m, maxilla. C. Inner surface of mandible of Eobaphetes kansensis Moodie, from the Coal Measures of Washington County, Kansas. X 0.33. a, angular; p, prearticular; r, articular; s, splenial. The markings of the inner surface are as indicated in the drawing. The back portion of the angular shows a few radiating lines. The dentary is roughened in two portions: one near the teeth, the other at the tip, where there is a cartilaginous roughening for union with its mate. The remainder of the inner surface is relatively smooth. The outer surface shows at the anterior end the typical labyrinthodont sculptur- ing, which becomes slight grooves and ridges posteriorly. I detect evidences of the operculo-mandibular lateral-line canal throughout the entire length of the mandible. Its location is indicated by the heavy broken line. The suture between the dentary and angular is quite clear. The suture separating the dentary and splenial joins the angular suture about midway of the length of the jaw. Measurements of Skull Fragment of Eobaphetes kansensis Moodie. (Cat. No. 6699, U. S. Nat. Mus.) mm. rom. Total length of portion preserved 109 Length of tooth 10 Maximum width of maxilla 45 Width of tooth at base 4 1 hickness of maxilla 7 19-2 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Measurements of Left Ramus of Mandible of Eobaphetes kansensis Moodie. (Cat. No. 6680, U. S. Nat. Mus.) mm. mm. Total length of jaw, as preserved 305 Length of anterior internal mandibular foramen. ... 56 Greatest width 79 Greatest width 15 Least width 24 Least width 7 Length of angular 132 Length of posterior internal mandibular foramen. . . 77 Width of angular 45 Greatest width 28 Length of largest tooth 10 Least width 14 Width of largest tooth at base 6 Length of bridge 16 Length of most posterior tooth 6 Width of bridge 8 Width of most posterior tooth at base 4 Ribs. — There are portions of two dorsal ribs preserved on the block of coal with the skull. These show characters very similar to those exhibited by the rib ascribed to Macrerpeton huxleyi Cope, and also those of Metoposaurus diagnosticus von Meyer (242) and Anaschisma. The ribs are solid, heavy, curved, and have a longitudinal groove on the middle of each side. The heads of the ribs in the present specimen are obscured and nothing can be said of them except that they appear to be large. Measurements of Ribs of Eobaphetes kansensis Moodie. mm. Length of preserved portion 130 Width at distal end 18 Thickness of rib 5 Genus DENDRERPETON Owen, 1853. Owen, Quart. Jour. Geol. Soc. London, ix, p. 66, 1853. Type: Dendrerpeton acadianum Owen. The genus is characterized by Dawson as follows (Phil. Trans., 1882, p. 635): " Lizard-like, with anterior and posterior extremities nearly equal; the skull somewhat elongate with small orbits, and the nostrils placed at the front. The cranial bones sculp- tured. The teeth plicated at the base, more especially on their inner sides. A series of large teeth on the palate. The body was covered above with imbricated horny scales and had lappets or pendants at the sides. The abdomen was protected by thin bony scales semi- elliptical or oat-shaped in form, and arranged in a chevron pattern. There was probably also a thoracic plate. Two species: D. acadianum and D. oweni. "Type: D. acadianum Owen." Dendrerpeton acadianum Owen. Owen, Quart. Jour. Geol. Soc. London, ix, p. 66, 1853. Dawson, Air-breathers of the Coal Period, p. 17, 1863. Dawson, Acadian Geology, 3d ed., p. 362. Dawson, Phil. Trans. Roy. Soc. London, pt. 11, p. 642, pi. 40, figs. 46 to 51; pi. 44, figs. 129 to 137. 1882. Type: Specimens Nos. 434-438, British Museum of Natural History. Horizon and locality: Coal formation at South Joggins, Nova Scotia. (Plate 6.) This species has been fully described and figured by Dawson (Air-breathers of the Coal Period, pp. 17-30, pi. in, figs. 1 to 30, 1863), who gives a detailed account of the discovery of the material of this species by himself and Sir Charles Lyell. He says, in part : "In form, Dendrerpeton acadianum was probably lizard-like; with a broad flat head, short, stout limbs and an elongated tail; and having its skin, and more particularly that of the belly, protected by small bony plates closely overlapping each other. It may have attained the length of 2 feet. The form of the head is not unlike that of Baphetes, but longer in pro- THE TEMNOSPONDYLOUS AMPHIBIA. I93 portion ; and much resembles that of the labyrinthodont reptiles of the Trias. The bones of the skull are sculptured as in Baphetes, but in a smaller pattern. The nostrils are small, and near the muzzle ; the orbits are circular, and separated by a space of more than their own diameter. In the upper jaw there is a series of conical teeth on the maxillary and inter- maxillary bones. Those on the intermaxillaries are much larger than the others, and have the aspect of canines or tusks. Within this outer series of teeth, but implanted apparently in the same bones, there is as in Archegosaurus a second series of teeth, closely placed, or with intervals equal to the diameter of one tooth. These inner teeth are longer than the others, implanted in shallow sockets, to which they are anchylosed, and have the dentine plicated, except toward the point. A third group of teeth, blunt at the points, largely hol- low in the interior, and with the dentine quite simple, appears in detached bones, which may represent the vomer. Only a part of this formidable armature of the teeth appears in the skull, as the bones of the roof of the mouth have been removed, adhering to the opposite side of the matrix ; but the fact of the occurrence of two sets of teeth was ascertained by Professor Wyman, from the original specimens, and is manifest in the fragment * * * while the other teeth, supposed to be vomerine, appear in fragments which must, from their size and collocation, have belonged to Dendrerpeton. It will be observed that all of these teeth are anchylosed to the bone ; and that those of the vomer are thinly walled and simple, the outer series on the maxillaries and intermaxillaries simple and flattened, while the inner series of teeth are conical and plicated. In the lower jaw there was a uniform series of conical teeth, not perceptibly enlarged toward the front; at least this is the case in the only specimen at present in my collection; which is, however, merely an imperfect cast in hard sandstone. ' ' The scapular and sternal bones seem to have been well developed and strong, but only portions of them are known. The fore limb of the adult animal, including the toes, must have been 4 or 5 inches in length, and is of massive proportions. The bones are hollow, and in the case of the phalanges the bony walls were thin, so that they are often found crushed flat. The humerus, however, was a strong bone, with thick walls and a cancellated structure toward its extremities; still, even these have sometimes yielded to the great pressure to which they have been subjected. The cavity of the interior of the limb-bones is usually filled with calc-spar stained with organic matter, but showing no structure; and the inner side of the bony wall is smooth, without any indications of cartilaginous matter lining it. ' ' The vertebrae, in the external aspect of their bodies, remind one of those of fishes, expanding toward the extremities, and being deeply hollowed by conical cavities, which appear even to meet in the center. There is, however, a large and flattened neural spine. The vertebrae are usually much crushed, and it is almost impossible to disengage them from the stone. * * * in its long neural and hasmal spines, reminds us of the caudal vertebrae of those batrachians and reptiles which have tails flattened for swimming, and probably indi- cates that this was the case with Dendrerpeton. The ribs are long and curved, with an expanded head, near to which they are solid, but become hollow toward the middle ; and the distal extremities are flattened and thin-walled. The posterior seems to have been not larger than the anterior, perhaps smaller. The tibia is much flattened at the extremity, as in some labyrinthodonts, and the foot must have been broad, and probably suited for swim- ming or walking on soft mud, or both. That the hind limb was adapted for walking is shown not merely by the form of the bones, but also by that of the pelvis, the best preserved speci- men of which I have represented (208, pi. in, fig. 28). "The external scales are thin, oblique-rhomboidal or elongated-oval, marked with slight concentric lines, but otherwise smooth, and having a thickened ridge or margin; in which they resemble those of Archegosaurus, and also those of Pholidogasier . * * * The microscopic structure of the scales is quite similar to that of the other bones, and different from that of the scales of ganoid fishes * * * ." 194 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. Dendrerpeton oweni Dawson. Dawson, Quart. Jour. Geol. Soc. London, xvm, p. 460. Dawson, Air-breathers of the Coal Period, p. 32, 1863. Dawson, Acadian "Geology, 3d ed., p. 368. Dawson, Phil. Trans. Roy. Soc. London, pt. 11, p. 643, pi. 44, figs. 131, 138, 139, 1882. A smaller species than the preceding. The form (plate 13) is fully described by Dawson (Air-breathers of the Coal Period, p. 32, pi. iv, 1863) as follows: "Among the reptilian remains found in erect trees at the South Joggins, there have occurred several portions of skeletons which, from their sculptural cranial bones, plicated teeth, and the forms of their scales and limb-bones, I have referred to the genus Dendrer- peton, but to individuals of much smaller size than the full-grown specimens of D. acad- ianum. It did not occur to me to suppose that these were specifically distinct from the larger individuals, until I observed that bones of this kind, contained in the collections sent by me to the Geological Society, or represented in the figures drawn to illustrate one of my papers, were referred by Professor Owen, in his notes on these specimens and figures, in the Journal of the Geological Society, to the genus Hylonomus; which is quite distinct from Dendrerpeton, as will be explained in sequel. ' ' I was thus induced to reexamine all the specimens in my collection and the result has been to establish a strong probability that there is in reality a second species of Dendrer- peton, smaller than D. acadianum, and differing from it in several points. This species I propose to name D. oweni. It differs from D. acadianum in the following particulars : (1) its much smaller size; (2) its long and hooked teeth (it will be seen that these teeth differ very markedly in their proportions and form from those of the larger species) ; (3) the greater plication of the ivory in the intermaxillary teeth (in D. acadianum these teeth are, on the outside, simple almost to the base, and plicated on the inner side, while in this species they are plicated all around like the inner maxillary teeth) ; (4) the form of the skull, which has the orbits larger in proportion, and is also shorter and broader. On the other hand, when we have described the species Hylonomus, it will be seen that this animal, except in size, differs from them quite as widely as does D. acadianum. ' ' The distinctness of D. oweni is further confirmed by the fact that I possess small jaw-bones of Dendrerpeton,- about the size of those of this species, but having the teeth similar in form to those of the larger species ; these I suppose to have belonged to young individuals. "On examining the figures (208, pi. iv) it will be seen that the bones of the skull were corrugated as in the large Dendrerpeton, but with a smaller pattern. The forms of the jaw- bones also, and of the vertebras, ribs, scapular bone, bones of the limbs, and bony scales, are very similar, and indicate that in general form this creature was not far removed from its larger relative. The bones of the foot especially deserve attention. This is the most per- fect foot of Dendrerpeton hitherto found; and I have enlarged it in the figure (208) in order more distinctly to show its parts. It presents three long toes with traces of a smaller one at each side, so that there were probably five in all. If these toes be compared with the foot- prints on the slab discovered by Dr. Harding, it will be seen that they very closely corre- spond, though the toes of the present species are much smaller. The footprints are precisely those which we may suppose an animal of the size of Dendrerpeton acadianum would have made if, as the bones found render in every way probable, this larger species had a foot similar to that of D. oweni. I suppose, for this reason, that these footprints are really those of Dendrerpeton acadianum and that this species continued to exist from the time of the lower Coal Measures to the period when those higher beds of the series, in which its bones are found at the Joggins, were deposited. The present species must have lived in the same places with its larger relative, but may have differed somewhat in its habits. Its longer and sharper teeth may have been THE TEMNOSPONDYLOUS AMPHIBIA. 195 better suited for devouring worms, larvae, or soft-skinned fishes, while those of the larger Dendrerpeton were better adapted to deal with the mailed ganoids of the period, or with those smaller reptiles which were more or less protected with bony or horny scales. REMAINS OF SKIN AND HORNY SCALES. ' ' In one of my earliest explorations of the reptile-bearing stumps of the Joggins, I observed on some of the surfaces patches of a shining black substance, which on minute examination proved to be the remains of cuticle, with horny scales and other appendages. The fragments were preserved; but I found it impossible to determine with certainty to which of the species whose bones occur with them they belonged, or even to ascertain the precise relations of the several fragments to each other. I therefore merely mentioned them in general terms, and stated my belief that they may have belonged to the species of Hylono- mus * More recently other specimens have been obtained, and I have undertaken the detailed examination of the whole. I shall now endeavor to describe the principal or most continuous fragments, and afterward to consider the probabilities of their having belonged to certain of the reptiles entombed with them. I do this here, rather than under the titles of these several animals, on account of the uncertainty which still rests on the assignment of certain portions of this cuticle to the species in question, and which renders it more con- venient to consider these peculiar remains in one place and to compare the different portions with each other. "(i) One of my specimens is a flattened portion of cuticle 2.25 inches in length. The greater part of the surface is smooth and shining to the naked eye, and under the micro- scope shows only a minute granulation. A limited portion of the upper and, I suppose, anterior part is covered with imbricated scales, which must have been membranous or horny and generally have a small spot or pore near the outer margin, some having in addition smaller scales or points on their surfaces. In contact with the upper part of this specimen there were many fragments of the skull of Dendrerpeton oweni. "(2) Another portion of the cuticle, similarly marked, appears to preserve the form of the posterior part of the body and tail of the animal, and also a mark representing the point of attachment of the hind leg; near to which, and along the dorsal ridge, is a portion of the skin covered with much smaller scales. This was found in close proximity to a mass of bones of Dendrerpeton oweni, mingled with some of Hylonomus lyelli. "(3) A third and still larger surface of integument with similar markings has upon it a number of vertebras and detached bones of the small reptile Hylonomus wymani, to be described in the sequel; for which species, however, it would be much too large a covering. "(4) Another well-preserved fragment, less than 2 inches in length, exhibits very dif- erent markings. It is nearly covered with very small imbricated scales, thicker than those on the specimens previously described. On either side of what seems to have been the mid- dle line of the back, there is a series of pointed flat horny processes, which probably formed a double spinous crest. Without these there are tufts of strong bristles, and exteriorly to these last are rows of flat, thick, horny plates, transversely wrinkled. Near to these was a row of conical truncated tubercles. Sections of these appendages show them to have been horny and attached to the cuticle. None of them have bony structure. "(5) Near this last portion of cuticle, and possibly belonging to it, are pointed and probably membranous appendages, marked on each side with rows of scales not overlapping and each with a pore in its center. The manner in which these appendages are bent and wrinkled shows that they must have been soft, except at the tips, which seem to have been hard and horny, and they are arranged in series, as if originally placed along the sides of the * Journal of Geological Society, vol. xvi. I96 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. neck or abdomen, or both. The use of these appendages is not easy to conjecture. They remind us of the gular pouches of iguana, and of the lateral expansions of some geckos and of the Draco volans. Possibly they formed lateral parachutes, aiding the animal in moving over soft mud, or perhaps in leaping and swimming. "(6) Some other fragments appear to have belonged to a different species from either of the foregoing. The best-preserved specimen, which is about 1 inch in length and half an inch in breadth, is covered with very small imbricated scales. It is crossed by 6 or 7 obscure ridges, which both at the bottom and along a mesial line project into points covered with larger scales. A row of large scales with round pores connects these along the lower side. If, as seems probable, this fragment belonged to the side of the trunk or tail, it would perhaps indicate a division of the subcutaneous muscles into an upper and lower band, as in the newts. A separate fragment with transverse horny ridges and another with a longer lobe, similar in structure to those above mentioned, may perhaps be referred to the same animal. A larger patch of skin presents similar imbricated scales, but without a mesial line, and with an edging of larger scales. ' ' Six species of reptiles have left their bones in the repositories containing these remnants of cuticle. Of these, Dendrerpeton acadianum, was an animal of too great size to have been clothed with integument of this character and of such dimensions. Hylono- mus aciedentatus and Hylerpeton dawsoni are each represented by only a single specimen, and these did not occur in proximity to any of the portions of cuticle, except that the appendages were found near a specimen of the former. Of the three remaining species, Dendrerpeton oweni, from its size, the number of specimens found, and the juxtaposition of their bones to the fragments of cuticle, appears to have the best claim to the integu- ment included under Nos. 1, 2, and 3; and in this case, while the creature had its throat, and perhaps its abdomen, armed with bony scales, its upper parts and tail, as well as its limbs, had a uniform covering of small, thin imbricated horny scales, in the manner of many modern reptiles. " If the remaining portions of integument, Nos. 4 and 5, as would seem likely, belonged to two species, both of smaller dimensions, there would seem little reason to doubt that these were Hylonomus lyelli and H. wymani. In this case, both of these species must have possessed a highly ornate covering of horny scales and appendages, comparable with that of many of the modern lizards, while there seems good reason to believe, as stated in a pre- vious paper, that they were in part protected by bony scales somewhat like those of Den- drerpeton. These points, however, we shall consider more in detail under the sections which refer to the species of Hylonomus. Before leaving these curious specimens of ancient skin, the most ancient I suppose known to exist, it is of interest to observe that the thicker portions, when broken across, have the aspect of jet, or of pure shining coal, and thin slices, under the microscope, have the same rich brown colour with that material, though rather more translucent. When burned, fragments of the substance give a strong flame, and a bituminous and ammoniacal odour. We have thus an example of the production of coal from animal membrane, no doubt gelatinous and horny in the first instance, but which has proved itself capable of the same chemical changes that have been experienced by the vegetable matter buried with it. In order that this substance should be preserved in this way, it would be neces- sary that it should either be kept dry and hard, or that it should be immediately buried in matter impervious to air and kept moist. The latter conditions are the more probable. The preservative qualities of the peaty vegetable matter imbedded with it must be con- sidered ; and it is possible that these hollow stumps, partly filled with fragments of Sigillaria bark, may have formed natural tan-pits, in which animal membranes would be preserved in a manner impossible in ordinary sediments. If this were the case, we may yet find an entire reptile, preserved as a flattened mummy, in one of these strange repositories." THE TEMNOSPONDYLOUS AMPHIBIA. 197 Genus PLATYSTEGOS Dawson, 189S. Dawson, Proc. and Trans. Roy. Soc. Canada, 1895, xn, p. 77 (sec. iv). Type: Platystegos loricatum Dawson. Dawson's description is as follows : "Head broad and short, orbits very large, cranial bones deeply sculptured; teeth strongly plicated and curved, with sharp edges at apices, especially the inner palatal teeth, which are very large. Many minute teeth on the vomerine bones; vertebrae ossified, bicon- cave ; limb bones imperfectly ossified, short ; lower surface protected with a thoracic plate and thick, densely imbricated bony scales in transverse rows; body above with thin, round scales, concentrically marked." Platystegos loricatum Dawson. Dawson, Proc. and Trans. Roy. Soc. Canada, xn, p. 77, 1895. Dawson describes the species: " Head about 8 cm. long; when flattened 9 cm. broad across parietal foramen; squa- mosal and temporal bones projecting backwards in points much behind the condyles; parietal foramen small, orbits large; length of longest tooth seen 7 mm.; cranial bones closely and deeply pitted; humerus with very thin bony walls, cartilaginous within, 3.5 cm. long." Erect tree, coal formation, at the South Joggins; collected by P. W. McNaugh- ton. Type in Peter Redpath Museum at McGill University. Dawson regarded the form as of uncertain relationship. CHAPTER XXV. THE STEREOSPONDYLOUS AMPHIBIA FROM THE COAL MEASURES OF NORTH AMERICA. DEFINITION OF THE ORDER STEREOSPONDYLIA ZITTEL, 1887. Zittel, Handbuch der Paleontologie, Bd. m, Abth. I, p. 397, 1887. Large terrestrial vertebrates; largest of the class. Skull equal to one-fourth or one-third of the entire body in at least one species, Metoposaurus diagnosticus von Meyer (242). Lateral-line canals always present (458) on the skulls as deeply impressed grooves which, in life, were possibly roofed over by a cartilaginous or other connective-tissue membrane. The sensory organs undoubtedly being sup- plied by the superficial ophthalmic branch of the trigeminal nerve, branches of which pierced the cranial elements near the grooves, no evidence of such openings in the bottoms of the grooves ; the condition probably being analogous to Hydrola- gus colei and other chimaeroids. Vertebras stereospondylous, with well-developed neural arches from which projected the well-developed zygapophyses, sometimes slightly amphiccelous and pierced for the notochord, such forms being uncertainly placed in the group. Tail unknown, possibly short. Limbs and girdles well developed (243), phalangeal formula unknown; carpus osseous and tarsus un- known. Pectoral girdle composed of osseous scapulae, clavicles, interclavicle, coracoid (?); clavicles and interclavicle ventrally sculptured. Pelvic girdle com- posed of osseous pubis, ischium, and ilium (242), the pubis a small plate, in life largely cartilaginous, the three uniting by cartilaginous union to form the acetabu- lum. Ventral armature unknown, possibly wanting. Range : Pennsylvanian to Upper Triassic. Distribution: North America, Europe, Asia, Africa, and Australia. Family MASTODONSAURIDjE Huxley, 1863. Huxley, Quart. Jour. Geol. See, xix, p. 65, 1863. Lydekker, R., Cat. Fossil Reptilia Amphibia, pt. iv, p. 141, 1890. Skull triangular, and more or less elongated, with the cranial bones very strongly sculptured, the occipital condyles ossified (49), and large palatal vacuities; dentine of teeth with very complex plications (502) ; no bony rings (242) in scle- rotic; and no ventral scutes. Bodies of vertebrae (49) fully ossified in the adult. There are large palato-vomerine tusks on the inner side of the maxillary teeth; and the palatines run parallel to the maxilla. The mandible has a large post- articular process; and there is a small inner series of mandibular teeth. In the type genus the pubes are separate from the ischia, and do not enter into the formation of the acetabulum; and the sacral ribs form kidney-shaped disks (393). Represented in North America by a single tooth from the Carboniferous of Kansas. Described by Williston as Mastodonsaurus sp. (Kans. Univ. Quart., vi, pp. 209-210, pi. xxi). Represented in the Triassic of Wyoming by Anaschisma browni, Branson (49). Mastodonsaurus sp. indet. Williston, Kans. Univ. Quart., vi, p. 209, 1897. The specimen preserved comprises the entire crown of a single vomerine (?) tooth, 38 mm. in length by 14 mm. in diameter at base (pi. 2 1 , fig. 6) . The immediate tip had been partly worn away in life, but was acuminate. It is composed of a dense 198 THE STEREOSPONDYLOUS AMPHIBIA. 199 blackish material, with the exterior smooth, shining black. It has about 20 narrow flutings, nearly straight, running from the base to the tip, separating shallow grooves. A transverse section of the base shows a narrow pulp-cavity not more than 5 mm. in diameter which extends in about the same proportional width to beyond the middle of the tooth, and in all probability to near the apex. The cross-section of the tooth throughout is nearly or quite circular. A hemisection of the tooth was made near the middle, showing a structure most remarkably like that of Mastodonsaurus; so nearly alike, in fact, that there is no dif- ference from the large figure given by Owen of a section of Mastodonsaurus (502). The discovery of this tooth in the Kansas Coal Measures is of great interest, proving, as it does, the presence of true labyrinthodonts from a lower horizon than elsewhere recorded. The discovery of Eobaphetes kansensis Moodie in the Carbo- niferous of Washington County (473) would seem to indicate another labyrinthodont. The tooth from Louisville was possibly not the first evidence of labyrintho- donts in North America, since the discovery by Marsh of Eosaurus acadianus from the Coal Measures of Nova Scotia, possibly a member of the Stereospondylia, ante- dates this discovery 30 years. The specimen is preserved in the Museum at the University of Kansas. AMPHIBIAN FOOTPRINTS FROM THE COAL MEASURES. Footprints may be said to be fairly common in the Coal Measures of North America. Especial attention has been given to the classification of these objects by G. F. Matthew (408-413), Dawson (208-210) and others. Hay (317, pp. 538- 553) has given a catalogue of all the species described from the Coal Measures of North America, to which the reader is referred for further information in regard to these interesting evidences of former animal activities. The writer has not been interested in the taxonomy of footprints, but has studied such as have come to his notice (465). A description of the species Dromopus agilis Marsh (fig. 43) is given here, because there is a large slab in the University of Kansas Museum which has not been figured. Since the chief interest in the present contribution is mor- phology, footprints are thus scantily dealt with. Leidy (374) , Dawson (207) , Moodie (465, pi. lxiv, fig. 1) and others have given various brief descriptions of Coal Meas- ures footprints, probably all of which are evidences of Amphibia which are other- wise unknown. Dromopus agilis Marsh. Marsh, Jour. Sci. (3), xlviii, p. 82, pi. ii, fig. 3; pi. iii, fig. 3, 1894- Hay, Bull. U. S. Geol. Surv., No. 179, p. 543, 1902. Type: Specimen in the Yale University Museum. Horizon and locality: Osage limestone (Coal Measures), near Osage City, Kansas. In 1894 Professor Marsh described a collection of footprints which he had secured from Professor B. F. Mudge, of Manhattan, Kansas, who had collected them in Osage County, Kansas, in a rock quarry, having purchased a large quantity of rock from the quarrymen for that purpose. A preliminary note by Mudge (490) 200 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. was published in the Transactions of the Kansas Academy of Science and later copied in the American Journal of Science. Professor Marsh's description (406) of the remains is as follows: " The impressions are well preserved in a calcareous shale, which separates readily into thin slabs, each representing a surface of the beach at the time the footprints were made upon it. A few shells in the shale are sufficient to prove that the formation is marine (no shells are evident in the slab at the Museum of the University of Kansas, but the slab is quite arenaceous). Trails of annelids, or perhaps of other invertebrates, are seen on some of the surfaces. The footprints of vertebrate animals, however, are of paramount importance, and the large number and variety of these here recorded on a single surface, if they could be rightly interpreted, would form an interesting chapter of land vertebrate life in the Carboniferous, about which so little is at present known." Professor Marsh's description of Dromopus agilis is as follows: " The third series of footprints is of special interest, and indicates an animal very distinct from the two already described. The diagram rep- resents the impression of the phalanges sufficiently in de- tail to indicate (406) their number and general form. A striking feature in the fore and hind feet of this animal was the long, slender digits terminated by sharp claws. Another point of interest, as recorded in the footprints, is that the animal in walking swung the hind feet outward, and so near the ground that the ends of the longer toes sometimes made trails in the mud, marking accurately FlG 43._Footprints of Promopus agilis Marsh, from the Coal Measures of the Sweep Of the foot. This Osage County, Kansas. Original slab in University of Kansas Museum. would seem to indicate a Greatly reduced. comparatively short hind leg, rather than the long, slender one which the footprints themselves naturally suggest. " The animal which made these interesting footprints was probably a Lacertilian rather than an Amphibian, but there is also a possibility that it was a primitive Dinosaur." Further on Professor Marsh remarks (p. 84) : " So far as at present known, land vertebrate life began in the Carboniferous age, no footprints of other remains of this kind having been detected below the Subcarbo- niferous. That such remains will eventually be found in the Devonian, there can be no reasonable doubt, and perhaps even in the Silurian, if the land surfaces then existing can be explored." This last statement of Marsh's was, of course, partly demonstrated by the dis- covery of footprints in the Devonian rocks of Pennsylvania, which he described in 1896 as Thinopus aiitiquus. The footprints of Dromopus agilis Marsh which are THE STEREOSPONDYLOUS AMPHIBIA. 201 preserved in obverse in the University of Kansas Museum are of considerably greater length than those described by Professor Marsh. The measurements of one of the larger impressions are appended. There appear to be series of footprints of two different animals preserved on the large slab (5 by 7 feet), but their nature is essentially the same. Measurements of Large Footprint. mm. mm. Greatest length 200 Length of shortest toe 65 Greatest width 90 Width across heel 50 Length of longest toe 120 The slab is No. 5 of the University of Kansas Museum of Natural History, collected in 1873 by Professor B. F. Mudge at Osage, Kansas, and presented by him to the museum in 1875. The following list of Carboniferous amphibian (?) footprints is compiled from Hay's Catalogue of Fossil Vertebrates of North America. It is given here for the sake of completeness. Three types of amphibian footprints are described in the body of this work and they constitute the kind of material which an ichnologist has for the basis of his conclusions. The material is not very satisfactory for the morphologist, though much can be determined as to the foot structure. Genus and species. Allopus littoralis Anomccpus ? culbertsonii gallinidoides Anlhracopus ellangowensis Baropus tentus Batrachichnus plainvillensis. . . . Clieirotherium ? heterodactylum. reiteri Collettosaurus indiantensis Crucipes parvus Dromopus agilis Hylopus caudifer hardtngi logani minor trifidus sp. indet Limnopus vagus Xanopus caudutus Notalacerta missouriensis . . Nolamphibia magna Palaosauropus anliquior. . primcevus . . sydenensis . unguifer. . . Thenaropiis leptodactylus . . . ovoidaetylus . . . . pachydaityltts . . spharodactylus . Author. Horizon. Locality. Marsh Coal Measures. . . King Do King Do Leidy Do Marsh Do Woodworth Do Do Moore Do Cox Do Butts Do Marsh Do Dawson Do Dawson Do Dawson Do Dawson Do Dawson Do Do Marsh Do Marsh Do Butts Do Butts Do. Dawson. . . . Subcarboniferous Lea Coal Measures.. . Do Carboniferous (millstone grit). Coal Measures.. . Do Do Do Dawson. Dawson . King. King. King. King. Osage County, Kansas. Pennsylvania. Do. Do. Osage County, Kansas. Massachusetts. Pennsylvania. Do. Indiana. Missouri. Osage County, Kansas. Nova Scotia. Do. Do. Do. Do. Do. Osage County, Kansas. Do. Missouri. Do. Nova Scotia. Pennsylvania. Cape Breton Island. Nova Scotia. Pennsylvania. Do. Do. Do. BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA, WITH ESPECIAL REFERENCE TO THE AMPHIBIA FROM THE COAL MEASURES OF NORTH AMERICA. The following literature list, containing more than six hundred titles ( 1 824-1916), is believed to be fairly complete. It certainly includes all of the important contributions to the subject of the fossil Amphibia. A few brief bibliographies of fossil Amphibia have appeared from time to time, in special memoirs on various faunas, such as: Broili (58), Schwarz (541), Fraas (242), von Ammon (7), and Case (98). None of these have attempted a complete survey of the field. The references given below have all, or nearly all, been compiled from the original sources and every effort has been made to have them complete and accurate. 1. Abel, O. 1912. Grundziige der Paleobiologie der Wirbelthiere. 211-220, figs. 144-150. 2. Agassiz, L. 1862. Highly interesting discovery of new Sauroid remains. Amer. Jour. Sci. and Arts, Jan., xxxiii, 138. 3. Alberti, Fr. v. 1834. Beitrag zu einer Monographie des bunten Sandsteins, Muschelkalks und Keuper. 4. . 1864. Ueberblick ueber die Trias. 235-241. 5. Albrecht, P. 1883. Note sur le Basioccipital des Batraciens Anoures. Bull, de Musee Roy. d'Hist. Nat. de Belgique, n, 195-198, with figs, and a plate. 6. American Naturalist. 1878. A new fauna, xn, 327-328 (Ed.). 7. Ammon, Ludwig von. 1889. Die permischen Am- phibien der Rheinpfalz, Munich. 1-117, pis. 1-5, with extensive bibliography. 8. Andrews, C. W. 1895. Note on a specimen of Kera- terpeton galvani Huxley, from Staffordshire. Geol. Mag., Dec, iv, n, 81-84, fig. 9. Arldt, Theodor. 1907. Die Entwicklung der Konti- nente und ihre Lebewelt. 333, Karte 10. 10. . 1909. Die Stegocephalen und ihre Stellung unter den Wirbelthieren. Naturw. Rundschau, xxiv, 353-355- 11. Atthey, T. 1876. On Anthracosaurus russelli. Ann. and Mag. Nat. Hist., Ser. 4, xvm, 146-167, pis. 8-1 i. 12. Bailey, W. H. 1866. On the new discovery of fossil reptiles in the Carboniferous of southern Ireland. Geol. Mag., 11, No. 20, 84. 1875. Description of a new species of Laby- 13 14- 15- 10. 17. rinthodont Amphibian from the Coal at Jarrow Colliery near Castlecomer, Kilkenny. Rept. Brit. Assn. Adv. Sci., 62. (Describes Anthracosaurus edgei.) — ■ — ■. 1884. Some additional notes on Anthraco- saurus edgei Bailey. Rept. Brit. Assn. Adv. Sci., 96-97 (1883). Barkas, T. P. 1868. On the fauna of the low main Coal Seam, Northumberland. Geol. Mag., v, 580. . 1869. Unusual forms of Ctenoptychius. Geol. Mag., Jan., vi, 43, 2 woodcuts. . 1869. On a supposed mammalian tooth from the Coal Measures. Monthly Micros. Jour., 11, 104, figs. 1-5. 18. Barkas, T. P. 1869. On the discovery of a molar of a large reptile in the Northumberland Coal Meas- ures. Ann. and Mag. Nat. Hist., m, 419. (Pter- oplax cornuta.) 19. . 1869. Reptile Remains and Climadoxus. Ann. and Mag. Nat. Hist., iv, 438-440. 20. . 1873. A manual of Coal Measure paleon- tology. London. 21. Barrell, Joseph. 1906. Origin and significance of Mauch Chunk shale. Bull. Geol. Soc. Amer., xvm, 460. 22. Baur, George. 1886. Die alteste Tarsus (Archego- saurus). Zool. Anz., No. 216, 9 Jahrg., 104-106. 23. . 1886. The oldest Tarsus (Archegosaurus). Amer. Nat., xx, 173-174. 1886. Ueber die Homologien einiger Schadel- 24 25- 26. 27- 28. 29. 30- 31- 32. 32a 33- knochen der Stegocephalen und Reptilien. Anat. Anz., 1, No. 13, 348-350. . 1886. Ueber die Morphologie der Wirbel- saule der Amnioten. Biol. Centralb., vi, Nr. 1 1-12, 322-342, 353-363- . 1887. On the morphology of ribs. Amer. Nat., Oct., 942-945 (Archegosaurus). . 1887. Nachtriigliche Notiz zu meinen Be- merkungen ueber die Homologie der Schadel- knochen der Stegocephalen und Reptilien. Anat. Anz., 2, No. 21, 657-658. . 1888. Morphogenie der Carpus und Tarsus der Wirbelthiere. Pt. I, Batrachia. Jena, Verlag von Gustav Fischer, i-88, pis. 1-3, figs. . 1894. Bemerkungen liber die Osteologie der Schliifengegend der hoherer Wirbelthiere. Anat. Anz., x, 315-330. . 1896. Bemerkungen iiber die Phylogenie der Schildkroten. Anat. Anz., xn, 561-570. (Dis- sorophus.) . 1896. The Stegocephali. A phylogenetic study. Anat. Anz., 20ten Marz, xi, No. 22, 657- 673, with bibliography and eight figures. — . 1897. Archegosaurus. Amer. Nat., xxxi, 975-980. . 1897. Ueber die systematische Stellung der Microsaurier. Anat. Anz., xiv, 148-151. Bayer, Franz. 1880. Palaeobatrachus bohemicus von Meyer, aus der Braunkohle von Freudenhain. Sitzber. d. k. bohm. Ges. Prag,, 291-298, 1 taf. BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA. 203 34. Beasley, H. C. 1900. Notes on the type specimen of Cheirotherium herculis Egerton. Proc. Liver- pool Geol. Soc., xlii, 81, pi. v. 35. Bernard, Felix. 1895. Elements de Paleontologie. Batraciens, 740-760, with figs. 36. Beyrich, Ernst. 1850. Ueber einige organische Reste der Lettenkohlenbildung in Thueringen. Zeit. d. deutsch. geol. Gesell., 11, 165. 37. Bieber, V. 1880. Ueber zwei neue Batrachier der Bohm. Braunkohlenformation. Sitz. d. k. Akad. d. Wissen., Wien., lxxxii, abth. 1, 117, taf. iii, fig. 1. 38. Binney, E. W. 1883. Trans. Manchester Geol. Soc, vi, p. 42. 39. Blanchard, R. 1 88 1. Sur les Glandes cloacale et Pelvien et sur la Papille cloacale des Batraciens urodeles. Zool. Anz., iv, 9, 34. 40. Blanford, W. T. 1884. Address to the geological section of the British Association at Montreal, 21 pp.; also Nature, xxx. 41. Bolkay, S. J. de. 191 1. On a Pleistocene predecessor of Rana fusca Ros. Mittheil. aus d. Jahrb. k. Ung. Geol. R., xix, H. 3, 155-160, figs. 1-7. 42. Bolton, H. 1896. The animal life of the Lancashire Coal Measures. Trans. Manchester Micros. Soc, 123-135 (1895)- 43- . 1904. The paleontology of the Lancashire Coal Measures. Trans. Manchester Geol. Soc, xxviii, 378-420, 578-650, 668-689. 44. Boulenger, G. A. 1890. On the presence of ptery- goid teeth in a tailless batrachian (Pelobates cul- tripes), with remarks on the localization of teeth on the palate in batrachians and reptiles. Proc Zool. Soc. London, 664-666. -. 1904. On the characters and affinities of the Triassic reptile Telerpeton eligense. Proc Zool. Soc. London, 470-480, pi. xxx. 1910. Les Batraciens et principalement ceux 45- 46. d'Europe. 1. Paris. 47. Bradley, Frank H. 1870. Geology of Grundy County. Geol. Surv. Illinois, iv, 196. 48. Branco, W. 1887. Weissia bavarica, nov. gen. et sp. von Stegocephalia, ein neuer Stegocephale aus dem unteren Rothliegenden. Jahrb. d. k. Preuss. Geol. Landesanstalt f. 1886, 22-39, ta^- '; a's° Neues Jahrb. f.Mineral., Geol. u.Paleon., 1888, 117. 49. Branson, E. B. 1905. Structure and relationships of American Labyrinthodontidje. Jour. Geol., xin, No. 7, 568-610, figs. 1-19. 50. . 1910. Amphibian footprints from the Mis- sissippian of Virginia. Jour. Geol., xvin, No. 6, 356-358, 1 fig- 51. Braun, M. von. 1 841. Bericht tiber die Versamm- lung deutscher Naturforscher und Aertze. Braun- schweig, 74-75. (Trematosaurus.) 52. Braun, M. 1878. Ueber aussere Hilfsorgane bei der Begattung von Triton viridescens. Zool. Anz., I, 124. 53. British Museum. 1905. A guide to the fossil rep- tiles, amphibians and fishes in the department of geology and paleontology in the British Museum (Natural History). 8 plates, 116 figs., 8th ed. 54. Brodie, P. B. 1859. On the occurrence of footsteps of Cheirotherium in the Upper Keuper of War- wickshire. Quart. Jour. Geo!. Soc, xvi, 278. 55. Broili, F. 1899. Ein Beitrag zur Kenntniss von Eryops megacephalus Cope. Paleontographica, xlvi, 61-84, pis. 8-10. 56. 57- 58. 59- 60. 61. 62. 63- 64. 65- 66. - Broili, F. 1902. Beitragc zur Kenntniss von Diplo- caulus Cope (D. magnicornis). (Paleonto- graphica, li, 8.) Vorlaufigc Mittheilung, Cen- tralb. f. Mineral., Geol. u. Paleon., 536-541, with figs. . 1904. Stammreptilicn. Anat. Anz., xxv, No- 23, 577-587- — . 1904. Permische Stegocephalen und Rep- tilien. Paleontographica, li, 1-120, 5 pis. . 1905. Beobachtungen an Cochleosaurus bc- hemicus. Paleontographica, L, 1-16, pis. 1-2. . 1906. Ein Stegocephalen Rest aus den Bayrischcn Alpen. Centralb. f. Mineral., Geol. u. Paleon., No. 18, 568-571. 1908. Ueber die Rachitomen Wirbel der Stegocephalen. Monatsberichten d. deutsch. geol. Gesell., 60, Nr. 8/10, 235-240, figs. 1-7. • 1908. Ueber Sclerocephalus aus der Gas- kohle von Nurschan und das Alter dieser Ablager- ungen. Jahrb. d. k. k. Geol. Reichsanstalt, lviii, Heft 1, 49-69, Taf. 1. . 1908. Systematische und Biologische Be- merkungen zu der permischen Gattung Lyso- rophus. Anat. Anz., xxxm, Nr. 11/12, 290-298. 1909. Reviews of Literature. Neues Jahrb. f. Mineral. Geol. u. Paleon., Bd. 11, Heft 1, 132-138. . 1913. Ueber zwei Stegocephalenreste aus dem texanischen Perm. Neues Jahrb. f. Mineral., Geol. u. Paleon., Bd. 1, Heft 2, 96-100, Taf. IX. — . 1913. Unser Wissen iiber die altesten Tetra- 67. 68. Go- To. 71- 72. 73- 74- 75- 76. 77- poden. Fortschr. d. naturwissenschaftlichen For- schung, Halle, Bd. vm, 51-93, figs. 14-62. Bronn, H. G. 1852-54. Lethea Geognostica, mit Atlas. Bd. 1 (2), Kohlen Periode, von F. Roenier. Broom, R. 1903. On a new Stegocephalian (Ba- trachosuchus browni) from the Karoo Beds of Aliwal North, South Africa. Geol. Mag., n. s., Dec, iv, x, No. 473-499. 1-2. figs- . 1904. On a new South African Labyrinthc- dont (Cyclosaurus albertyni). Records of the Albany Museum, 1, No. 3. -, 1907. On the geological horizons of the vertebrate genera of the Karoo Formation. Records of the Albany Museum, 11, No. 1. -. 1908. On a new Labyrinthodont (Rhine- suchus whaitsi) from the Permian beds of South Africa. Ann. So. African Museum, iv, pt. viii, May 1908, 373. -. On the nomenclature of the elements of the amphibian shoulder girdle. -. 1910. A comparison of the Permian reptiles of North America with those of South Africa. Bull. Amer. Mus. Nat. Hist., xxvin, art. xx, 197- 234- 191 3. On the structure of the mandible in the Stegocephalia. Anat. Anz., Bd. 45, No. 2/3, 73-78, figs. 19 1 3. Studies on the Permian temnospondy- lous Stegocephalians of North America. Bull. Am. Mus. Nat. Hist., xxxn, 563-595, 21 figs. Brown, Barnum. 1908. The Conard fissure, a Pleistocene bone deposit in northern Arkansas, with description of two new genera and twenty new species of mammals. Mem. Amer. Mus. Nat. Hist., ix, pt. iv, 206, pi. xxii. Browne, M. 1893. On some vertebrate remains not hitherto recorded from the Rhaetic beds of Britain. Rept. Brit. Assn. Adv. Sci., 748. 204 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. 78. Brownrigg, W. B. 1865. Jour. Roy. Geog. Soc. Ireland, I, pt. 2, 145. 79. Budgett, J. S. 1902. On the structure of the larval Polypterus. Trans. Zool. Soc. London, xvi, pt. vil, Oct.; also Works, 154-176, pis. x-xii, 1907. 80. Burmeister, H. 1840. Die Labyrinthodonten aus dem bunten Sandstein von Bernburg. 4 plates (same 1849) (?). 81. . 1850. Die Labyrinthodonten aus dem Saar- briicken Steinkohlengebirge. Quarto, pis. 1-4. 82. Butts, Edward. 1891. Recently discovered foot- prints of the amphibian age in the Upper Coal Measure Group of Kansas City, Missouri. Kans. City Scientist, v, No. 2, 17-19, figs. 83. . 1891. Footprints of new species of amphib- ians in the Upper Coal Measure Group of Kansas City, Missouri. Kans. City Scientist, V, 44, figs. 84. Carus, J. Victor. 1875. Labyrinthodontia. Hand- buch der Zoologie, Bd. I, 485-486. 85. Case, E. C. 1898. The significance of certain changes in the temporal region of the primitive reptilia. Amer. Nat., xxxn, No. 374, 69. . 1900. The vertebrates from the Permian bone bed of Vermilion County, Illinois. Jour. Geol., viii, 698-729, 5 plates. 1902. On some vertebrate fossils from the 86. 87. 88. 89. 90. 91- 92. 93- 94. 95- 96. 97- 98. 99- Permian beds of Oklahoma. 2d Biennial Rpt. Dept. Geol. and Nat. Hist. Territory Oklahoma, 1901-1902, 62-68. 1902. Paleontological notes. Jour. Geol., x, 256-261. -. 1903. New or little-known vertebrates from the Permian of Texas. Jour. Geol., xi, 394-402. 1907. Additional description of the genus Zatrachys Cope. Bull. Amer. Mus. Nat. Hist., xxiii, art. xxix, 665-668. . 1907. Revision of the Pelycosauria of North America. Pub. No. 55, Carnegie Inst. Washing- ton, Stegocephalia, 159. -. 1908. On the value of the evidence furnished by vertebrate fossils of the age of certain so-called Permian beds in America. Jour. Geol., xvi, No. 6, 572-580. . 1908. Notes on the skull of Lysorophus tri- carinatus Cope. Bull. Amer. Mus. Nat. Hist., xxiv, 531-533, fig. 1. -. 1908. Description of vertebrate fossils from the vicinity of Pittsburgh, Pa. Annals of the Car- negie Museum, iv, Nos. iii-iv, 234-241, pi. lix, Apr. 1908. A great Permian delta and its verte- brate life, with restorations by the author. Pop. Sci. Month., lxxiii, No. 6, 557-568. . 1910. New or little-known reptiles and amphibians from the Permian? of Texas. Bull. Amer. Mus. Nat. Hist., xxvm, art. xvn, 163-181. 191 1. A revision of the Cotylosauria of North America. Pub. No. 145, Carnegie Inst. Washington. . 191 1. Revision of the Amphibia and Pisces of the Permian of North America. Pub. No. 146, Carnegie Inst. Washington, 1-148, pis. 1-25, 51 text-figs. . 1912. Paleozoic Reptilia and Amphibia. Bull. Geol. Soc. Amer., 23, 200-204. 1915. The Permo-carboniferous beds of North America and their vertebrate fauna. Pub. No. 207, Carnegie Inst. Washington, 1-176, pis. 1-24. 101. Chamberlin, T. C. 1900. On the habitat of the early vertebrates. Jour. Geol., vm, 400-412. 102. , and R. D. Salisbury. 1906. Geology, earth history. Amphibia, 11, 606-610. 103. Clark, C. T. 1847. On the neighborhood of Bom- bay and certain beds containing fossil frogs. Quart. Jour. Geol. Soc., in, 221-224. (See Owen, 1847.) 104. Colliery Guardian. Description of claspers of Amphibia. Mar. 10, 1 87 1. (Barkas.) 105. Cope, E. D. 1865. On Amphibamus grandiceps, a new Batrachian from the Coal Measures. Proc. Acad. Nat. Sci. Phila., 134-137. See also Geol. Surv. 111., 11, 135, fig., pi. 32, 1866. 106. . 1866. Remarks on extinct vertebrates of the Mesozoic Red Sandstone. Proc. Acad. Nat. Sci. Phila., 249. (Mastodonsaurus durus.) 107. . 1866. Supplement to the description of vertebrates. Geol. Surv. 111., n, 135-141, fig., pi. 32, fig. 8. (Amphibamus.) 108. . 1866. On the structure and distribution of the genera of the arciferous Anura. Jour. Acad. Nat. Sci. Phila., 2d ser., v, 67-112. 109. . 1866. On the families of Raniform Anura. Jour. Acad. Nat. Sci. Phila., 189. no. . 1868. Synopsis of the extinct Batrachia of North America. Proc. Acad. Nat. Sci. Phila., 208-221. in. . 1868. Two Batrachians from North Caro- lina. Proc. Acad. Nat. Sci. Phila., xx, 211-212. (Pariostegus myops.) . 1869. Synopsis of the extinct Batrachia, Reptilia, and Aves of North America. Trans. Amer. Phil. Soc, xiv, 1-252, pis. 1-14. . 1 87 1. Observations on extinct Batrachia. Proc. Acad. Nat. Sci. Phila., 53. 1 87 1. Supplement to the synopsis of the 112. 113- 114. "5- 116. 117. 118. 119. extinct Batrachia and Reptilia of North America. Proc. Amer. Phil. Soc., xn, 41-52. 1871. Observations on the extinct Batra- chian fauna of the Carboniferous of Ohio (Linton). Proc. Amer. Phil. Soc, xn, 177. — . 1872. Carboniferous reptiles of Ohio. Amer. Nat., vi, 46. . 1873. Observations on the distribution of certain extinct Vertebrata in North Carolina. Proc. Amer. Phil. Soc, xn, 210-216. . 1873. On some new Batrachia and Fishes from the Coal Measures of Linton, Ohio. Proc. Acad. Nat. Sci. Phila., 340-343. . 1874. Supplement to the extinct Batrachia, Reptilia and Aves of North America. I. Cata- logue of the air-breathing Vertebrata from the Coal Measures of Linton, Ohio. Trans. Amer. Phil. Soc, xv, 261-278. 1875. Check-list of North American Ba- trachia and Reptilia, with a systematic list of the higher groups and an essay on geographical distribution based on the specimens contained in the U. S. National Museum. Bull. U. S. Nat. Mus., 1, 7-12. . 1875. Synopsis of the Vertebrata whose remains have been preserved in the formations of North Carolina. Appendix "b" by Kerr, W. C. Rept. Geol. Surv. N. C, Raleigh, 29-52, pis. v-viii. . 1875. The extinct Batrachia of Ohio. Proc. Acad. Nat. Sci. Phila., 16. BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA. 205 123- 124. 125- 126. 127. 128. 129. 130. I3i- 132. 133- 134- iss- 137- 138. 139- 140. 141. 142. 143- 144. Cope, E. D. 1875. Synopsis of the extinct Batrachia from the Coal Measures. Geol. Surv. Ohio, 11, pt. 11, 351-410, pis. xxvi-xlv. . 1876. On some extinct reptiles and Batra- chia from the Judith River and Fox Hills Beds of Montana. Proc. Acad. Nat. Sci. Phila., 357; Pal. Bull. 23. • 1876. Description of some vertebrate re- mains from the Fort Union Beds of Montana. Proc. Acad. Nat. Sci. Phila., 260. (Ccratodus hicroglyphus, an amphibian clasper?.) . 1877. On the fossil remains of Reptilia and fishes from Illinois. Proc. Acad. Nat. Sci. Phila., 405. 1877. On the Vertebrata of the Bone Bed of eastern Illinois. Proc. Amcr. Phil. Soc, xvn, 53-64- . 1877. Report on the geology of the region of the Judith River, Montana, and on vertebrate fossils obtained on or near the Missouri River. Bull. U. S. Geol. and Geog. Surv. of Territories, hi, art. xix, Batrachia, 573, 576. . 1877. Descriptions of Vertebrata from the Permian and Triassic Formations. Proc. Amer. Phil. Soc., xvir, 182-193. . 1877. Extinct Vertebrata from North Carolina. Proc. Amer. Phil. Soc, 17, 182. . 1877. A continuation of researches among the Batrachia of the Coal Measures of Ohio. Proc. Amcr. Phil. Soc, xvi, 573-578. . 1878. Description of extinct Batrachia and Reptilia from the Permian of Texas. Proc. Amer. Phil. Soc, xvii, 522-526. 1878. The vertebra of Rachitomus. Amer. Nat., xii, 633. (Eryops.) 1879. The relations of the horizons of extinct Vertebrata of Europe and North America. Bull. U. S. Geol. and Geog. Surv. of the Terri- tories, v, No. 1, 33-34. -. 1880. Extinct Batrachia. Amer. Nat., xiv, 609-610. Reviews Fritsch's and Wiedersheim's papers. . 1880. The structure of the Permian Ganocephala. Amer. Nat., xiv, 383-384. (Eryops.) . 1 88 1. The Permian Formation of New Mexico. Amer. Nat., xv, 1020-1021. (Eryops and Zatrachys.) . 1 88 1. Catalogue of Vertebrata of the Permian Formation of the United States. Amer. Nat., xv, 162-164. — ■. 1 88 1. Ein Uebergangsglied von den Am- phibicn zu den Reptilien (Cricotus). Auszug in Kosmos von Krause, Bd. 9, 230-231. . 1 88 1. On some new Batrachia and Reptilia from the Permian Beds of Texas. Bull. U. S. Geol. Surv. Terr., vi, art. II, 79-82. -. 1882. Third contribution to the history of the Vertebrata of the Permian Formation of Texas. Proc. Amer. Phil. Soc, xx, 447-461 ; Pal. Bull. 35, 451. 1882. The Rachitomous Stegocephalia. Amer. Nat., xvi, 335. . 1882. Permian Vertebrata. Amer. Nat., xvi, 925. . 1883. Fourth contribution to the history of the Permian Formation of Texas. Proc. Amer. Phil. Soc, xx, 628-636. 145. Cope.E.D. 1883. On some Vertebrata from the Per- mian of Illinois. Proc. Acad. Nat. Sci. Phila., 108-110. 146. . 1884. The Batrachia of the Permian Period of North America. Amer. Nat., xvm, 26-39, pis. ii-v, 7 figs. 1 884. Fifth contribution to the fauna of the 147- 148. 149. 150. 151- 152. '53- 154- 155- 156. '57- 158. «59- 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. Permian of Texas and Indian Territory. Proc. Amer. Phil. Soc, xxn, 28-30, pi. 1 ; Pal. Bull. 39. . 1884. Note on the phylogeny of the Verte- brata. Amer. Nat., xvm, 1255-1256. 1885. The Batrachia of the Permian Beds of Bohemia and the Labyrinthodon from the Bijori Group, India. Amer. Nat., xix, 592-594. 1885. Paleontological nomenclature. Geol. Mag., London (3), 11, 575. (Batrachia for Am- phibia.) . 1885. The Batrachia of the Permian Beds of Bohemia. Geol. Mag., Ixindon (3), n, 527. . 1885. Second continuation of researches among the Batrachia of the Coal Measures of Ohio. Proc. Amer. Phil. Soc, xxn, 405-408. . 1885. On the evolution of the Vertebrata, progressive and retrogressive. Amcr. Nat., xix, 244, Mar. 1886. On the structure and affinities of the Amphiumidx. Proc. Amer. Phil. Soc, xxm, No. 123, 442-445. July- 1886. The Batrachian intcrcentrum. Amer. Nat., xx, 76, 175. . 1886. Systematic catalogue of the species of vertebrates found in the Beds of the Permian Epoch of North America. Trans. Amer. Phil. Soc, xvi, 285-296, pis. ii-iii. . 1887. A contribution to the history of the Vertebrata of the Trias of North America. Proc. Amcr. Phil. Soc, xxiv, 209. (Eupclor durus.) — . 1888. On the intercentrum of terrestrial vertebrates. Trans. Amcr. Phil. Soc, xvi, 243- 253, pl- 1, 6 figs. 1888. The Ossicula Auditus of the Batra- chia. Amer. Nat., xxn, 464-467, pl. vi. 1888. The pineal eye in extinct vertebrates. Amer. Nat., xxn, 914-917, pis. xv-xviii. 1888. On the shoulder girdle and extremi- ties of Eryops. Trans. Amer. Phil. Soc, xvi, 362-367, with plate. 1888. On the relation of the hyoid and otic elements of the skeleton in the Batrachia. Jour. Morphol., 11, 297-310. . 1889. The Batrachia of North America. Bull. U. S. Nat. Mus., No. 34, i-vi, 7-514, pis. i-lxxxvi. . 1889. Synopsis of the families of Verte- brata. Amer. Nat., xxm, 849-877. (Published in 1898 as "Syllabus of Lectures.") -. 1892. A preliminary report on the verte- brate paleontology of the Llano Estacado. Geol. Surv. Texas, 4th Ann. Rept., 12-17. 1892. On the phylogeny of Vertebrata. Proc. Amer. Phil. Soc, xxx, 278-280. . 1892. On some new and little-known Pal- eozoic vertebrates. Proc. Amcr. Phil. Soc, xxx, 221-229, pls- vii-viii. . 1893. Fritsch's Fauna of the Gaskohle of Bohemia. Amcr. Nat., xxyii, 709. . 1895. Some new Batrachia from the Per- mian Beds of Texas. Proc. Amer. Phil. Soc, .xxxiv, 452-457, plate ix, Nov. 15. 206 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA. 170. 171. 172. 173- 174. 175- 176. 177. 178. 179- 180. Cope, E. D. 1895. A Batra chianarmadillo. Amer. Nat., xxix, 998, Nov. . 1 896. The Paleozoic reptilian order Cotylo- sauria. Amer. Nat., xxx, 303. (Describes Eosauravus copei Williston, oldest known reptile.) 1896. Permian land vertebrates with cara- paces. Amer. 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New species of reptiles from the Coal Formation of Nova Scotia. Quart. Jour. Geol. Soc, xvi, 273. (Describes Hylonomus.) i860. On the terrestrial Molluska, a chilog- 205. 206. nathus Myriapod and some new species from the Coal Formation of Nova Scotia. Quart. Jour. Geol. Soc, xvi, 268. . 1862. Additional remains of land animals in the Coal Measures of Nova Scotia. Quart. Jour. Geol. Soc, xvm, 5-7. BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA. 207 2060. 207. 208. - 209. 211. 212. 213- 214.- 2I.S- 216. 217. 218. 219. Dawson, J. W. 1862. Descriptions of specimens of fossil Reptilia discovered in the Coal Measures of the South Joggins, Nova Scotia. Quart. Jour. Geol. Soc., xvih, 238-244, pis. ix, x. . 1863. Note on the footprint of a reptile from the coal of Cape Breton. Canadian Naturalist and Geologist, vm, 430-431, 1 fig.; review in Neues Jahrb. f. Min., Geol. u. Paleon., 64. 752- . 1863. Air-breathers of the Coal Period. Montreal, i-iv, 1-81, pis. 1-6. 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On a recent discovery of Carbonifer- ous Batrachians in Nova Scotia. Amer. Jour. Sci. (3), xii, 440-447- 1878. Acadian geology. Supplement to 3d 223. edition, London, Amphibia, 57-65. . 1882. On the results of the recent explora- tions of the erect trees containing animal remains in the Coal Formation of Nova Scotia. Phil. Trans. Roy. Soc. London, clxxiii, pt. 11, 621- 654. Pis- 39-47- . 1882. On the results of recent explorations of the erect trees (Batrachia). Canadian Natu- ralist (2), x, 252-254. (Abstract.) . 1 89 1. On new specimens of Dendrerpeton acadianum, with remarks on other Carboniferous Amphibians. Geol. Mag. (3), vm, 145-156, 4 figs. . 1891. Note on Hylonomous lyelli, with photographic reproduction of skeleton. Geol. Mag., n. s., Dec, m, vm, 258-259, pi. 8. . 1 89 1. On the mode of occurrence of remains of land animals in erect trees of Nova Scotia. Trans. Roy. Soc. Canada, ix. sec. IV, 127-128. . 1894. Some salient points in the science of the earth. 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(Archego- saurus extremity.) 597. — — ■ — . 1877. Ueber Protriton petrolei von Fried- richroda in Thuringen. Zeit. d. deutsch. geol. Gesell., xxix, 202. 598. Wheatley, Chas. M. 1871. The bone cave of eastern Pennsylvania. Amer. Jour. Sc (3), I, 384-385. 599. White, David. 1906. Geological position of the principal insect-bearing localities of the American Paleozoic, in " Handlirsch's Revision of American Paleozoic Insects. " Proc. U. S. Nat. Mus., xxix, No. 1441,664-667. 600. . 1907. Report of the field work in the coal districts of the state. Bull. No. 4, Illinois State Geol. Surv., 201 . (Gives correlation of the Mazon Creek Beds.) 601. Wiedersheim, Robert. 1876. Die altesten Formen des Carpus und Tarsus der heutigen Amphibien. Morphol. Jahrb., Bd. 2, 421-423, Taf. xxix, fig. 2. 602. . 1878. Ueber Labyrinthodon ruetemeyeri — Anatomie der Gehirn. Abhandl. d. schweizer paleontol. Gesell., v, 1-56, mit 3 tafeln. -. 1879. Die Anatomie der Gymnophionen, 603. pp. i-ioi, Tafeln i-ix. (Refers to extinct Am- phibia in connection with phylogeny.) 604. . 1886. Lehrbuch der vergleichenden Anat- omie der Wirbelthiere. 2 Auflage. 605. ■ and W. N. Parker. 1897. Elements of the comparative anatomy of vertebrates. Trans- lated by W. N. Parker. Labyrinthodontia, 242, fig. 193 c. 606. Wilder, H. H. 1909. History of the human body. Amphibia, 541-542. BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA. 217 607. Williston, S. W. 1897. Some vertebrates from the Kansas Permian. Kans. Univ. Quart., ser. A, VI, No. 1, 53, fig. Cricotus. 608. . 1897. A new Labyrinthodont from the Kansas Carboniferous. Kans. Univ. Quart., vi, No. 4, ser. A, 309, fig. and pi. (Refers tooth and skull fragments to Mastodonsaurus.) 1899. The Red-Beds of Kansas. Science 609. 610. 611. 612. 613. 614. 615. 616. 617. 618. 619. 620. 621. 622. (2), ix, 221. . 1899. Notes on the coraco-scapula of Eryops Cope. Kans. Univ. Quart., vm, No. 4, 185-186, pis. xxvii-xxx. (See Moodie (476).) 1900. Some fish teeth from the Kansas Cretaceous. Kans. Univ. Quart., ser. A, ix, No. I, Jan., pi. vi, figs. 13, 13a. . 1904. The temporal arches of the Reptilia. Biol. Bull., vil, No. 4, 175-192. . 1908. The faunal relations of the early vertebrates. Jour. Geol., xvn, No. 5, 389-402. Also, Outlines of geologic history, Willis and Salisbury, 1910, 163-175. . 1908. Lysorophus, a Permian Urodele. Biol. Bull., xv, No. 5, 229-240. Review in Rev. crit. de Paleozoologie 13th Annee, No. 2, 82. -. 1909. The skull and extremities of Diplo- caulus. Trans. Kans. Acad. Science, xxii, 122- 131, pis. 1-5. -. 1909. New or little-known Permian verte- brates, Trematops, new genus. Jour. Geol., xvn, No. 7, Nov., 636-658, 7 figs. 1909. Principal characters of the Chelydo- sauria, a suborder of temnospondylous Amphib- ians from the Texas Permian. A. A. A. S. Soc. Vert. Paleon. . 1910. Dissorophus. Jour. Geol., xvm, No. 6, 526-535. Pis- i-iii- . 1910. Cacops, Desmospondylus, new genera of Permian Vertebrates. Bull. Geol. Soc. Amer., xxi, 249-284, pis. 6-17. . 1910. New Permian Reptiles: Rachitomous vertebrae. Jour.Geol.,xvm, N0.7, 585-600, with figs. — — . 191 1. American Permian vertebrates. Am- phibia, 9-14, pis. xxxii, xxxviii. 1913. The primitive structure of the man- 623. 624. 624a. 624A. 6 2 4C. dible in amphibians and reptiles. Jour. Geol., xxi, 625-627, 1 fig. 1 9 14. The osteology of some American Per- mian vertebrates. Jour. Geol., xxii, No. 4,416, figs. 1914. Water reptiles of the past and pres- ent. Amphibia, 35, 47, and figures. - — ■ . 1914. Broiliellus, a new genus of am- phibians from the Permian of Texas. Jour. Geol., xxii, No. 1, 49-56, figs. 1-2. -. 19 1 4. Restorations of some American Permocarboniferous amphibians and reptiles. Jour. Geol., xxii, No. 1, 57-70, figs. 1-11. . 19 1 5. Trimerorhachis, a Permian temno- spondyl amphibian. Jour. Geol., xxiii, No. 3, 246-255, figs. 1-6. (i2^d. . 1916. Synopsis of the Amer. Permocar- boniferous Tetrapoda. Contrib. from Walker Museum, vol. I, No. 9, pp. 200-211, figs. 38-53. 625. Wiman, Carl. 1908. Ein Paar Labyrinthodonten- reste aus der Trias Spitzbergens. Bull. Geol. Inst. Univ. of Upsala, ix, 34-40, Tafeln 11, Nos. 17, 18. 626. •. 1913. Ueber das Hinterhaupt der Laby- rinthodonten. Bull. Geol. Inst, of Upsala, XII, 1-7, figs. 1-8. 627. 628. 629. 630. 631. 632. 633- °34- 635- 636. 637- 638. 639- 640. 641. 642. 643- 644- 645- Wiman Carl. 1914. LMierdieStegocephalenausder Trias Spitzbergens. Bull. Geol. Inst, of Upsala, xiii, 1-30, pis. i-ix, figs. 1-10, bibliography. Woodward, A. Smith. 1891. Catalogue of the fos- sil fishes in the British Museum, pt. 11, 474. (Note on the synonymy of Pygopterus, Colosteus, and Archegosaurus. ) . 1 89 1. On a Microsaurian (Hylonomus wildii) from the Lancashire Coal Field. Geol. Mag., n. s., Dec. in, vm, 211-213, with figs. . 1897. On a new specimen of the Stegoce- phalian Ceraterpeton galvani Huxley. Geol. Mag., Dec. iv, iv, No. 397, 293, with plate. 1904. On two new Labyrinthodont skulls of the genera Capitosaurus and Aphaneramraa (from the Trias of Spitzbergen). Proc. Zool. Soc. London, 170-176, with 2 pis. . 1906. The relations of paleontology to biology. Ann. and Mag. Nat. Hist., xvm, No. 106,312-318. 1905. Fishes and Labyrinthodonts. Paleon- tologica Indica, n. s., 11, Memoir No. 2, 10, 2 pis. -. 1909. On a new Labyrinthodont from Oil Shale, N. S. W. Rec. Geol. Surv. N. S. W., vm, pt. iv, 317-319. pl- 51- Woodward, H. B. 1887. The geology of England and Wales. The Northumberland Coal Field, 1 80. Wood worth, J. B. 1900. Vertebrate footprints on Carboniferous shales of Plain ville, Mass. Bull. Geol. Soc. Amer., xi, 449-454, pl. 40, fig. I. Wyman, Jeffries. 1843. Analogies which exist between the structure of the teeth of the Lepi- dostei (Gars and Gar-pikes) and those of the Laby- rinthodonts (extinct Amphibia). Proc. Bost. Nat. Hist. Soc, 1, 131-132; also Silliman's Jour- nal, 843. . 1853. Notes on the Reptilian remains (Dendrerpeton). Quart. Jour. Geol. Soc. Lon- don, ix, 64-66. -. 1856. On a batrachian reptile from the Coal Formation. Proc. Amer. Assn. Adv. Sci., 10th Meeting at Albany, 172-173. Reviews: Edinb. n. Philos. Jour. 1857, v, 360-361; Neues Jahrb. f. Mineral., Geol. u. Paleon., 1. 857, 340. 1858. On some remains of batrachian rep- tiles discovered in the Coal Formation of Ohio. Amer. Jour. Science, xxv (2), Mar., 158-164, fig. Review: Zeit. f. gesammt Naturwissensch., xin, 71 ; Neues Jahrb. f. Mineral., Geol. u. Paleon., 1858,340. Yakowlew, N. 1902. Neue Funde von Trias — Saurier auf Spitzbergen. Verhandl. Russ. k. Min. Gesell., xl, 180; Nachtrag, xli, 165. Zittel, Karl von. 1 887-1 890. Handbuch der Paleontologie, I Abth., Bd. in, 337~437. Amphibia. Miinchen u. Leipsic, R. Oldenbourg, also East- man's translation of above, 2, 1 14-139, figs. . 1888. Ueber Labyrinthodon ruetimoycri (Wiedersheim). Neues Jahrb. f. Mineral., Geol.u. Paleon., Bd. 11, 17. (Regards the form as a rep- tile.) -. 1899. Amphibien. Geschichte der Geologie und Paleontologie. Muenchen u. Leipzig, 1899. 827. Zwick, W. 1897. Beitrage zur Kenntniss des Baucs und der Entwickelung der Amphibienglie.lmassen, besonders des Carpus und Tarsus. Zeitschr. wissen. Zool., Bd. 63, 62-144, Taf. 4"5- AN INDEX TO THE BIBLIOGRAPHY OF FOSSIL AMPHIBIA. The following will be of assistance to those wishing aid in finding the literature on special phases, whether anatomical or geological, of the fossil Amphibia. The author's name and the number of his paper in the preceding bibliography are given in groups of classified subjects, beginning with distribution and ending with anatomy. The various anatomical notes of interest are especially widely scattered in papers which often deal with a variety of other subjects. I. Geological and Geographical Distribution. Devonian: North America: Marsh, 407. Europe: Lohest, 381; Weiss, 596; Thevenin, 566. Mississippian: North America: Branson, 50; Barrell, 21 ; Lea, 371- 372- Scotland: Huxley, 331-334; Atthey, II; Hancock and Atthey, 305. Coal Measures (Pennsylvanian) : North America: Agassiz, 2; Butts, 82, 83; Case, 86, 94; Cope, 105, 107, 115, 116, 118, 122, 123, 126, 127, 131, 145, 152, 167, 176; Dawson, 200-225; Eastman, 230; Hay, 316; Jaekel, 347; King, 356, 357; Marsh, 404, 406; G. F. Matthew, 408-413; Moodie, 458-465, 469-475, 478, 479, 482-486; Mudge, 490; Newberry, 495-498; Ray- mond, 531; Schwarz, 540, 541; Williston, 608; Wyman, 639, 640. Europe: Andrews, 8; Atthey, 11; Bailey, 12-13; Barkas, 15-20; Bolton, 42-43; Credner, 179-194 (see also under Permian); Davis, 199; Dunlop, 229; Embleton and Atthey, 235; Etheridge, 241; Fritsch, 251; Gaudry, 263-268, 278, 281, 282; Gergens, 291; Goldfuss, 297; Hancock and Atthey, 304-310; Lydekker, 394; von Meyer, 422, 426, 436; Oldham, 500; Owen, 507-509, 514, 515; Schwarz, 540; Thevenin, 565-568; Weiss, 597; Woodward, 629, 630; Woodworth, 636. Australia: Stephens, 557; Woodward, 634. Permian : North America: Broili, 55, 56, 58,63,65; Case, 86-93, 95-100; Cope, 129, 132-133, 137, 138, 140-144, 156, 169, 170-175; Emmons, 237; Gregory, 299; W. D. Matthew, 414, 415; Moodie, 457, 458, 476, 477; Sternberg, 558; Strickler, 564; Willis- ton, 607, 609, 610, 614, 615, 616, 617-624. Europe: Ammon, 7; Branco, 48; Broili, 59, 60, 61, 62, 66; Credner, 179-195; Deichmueller, 224, 225; Eichwald, 233; Emery, 236; Fritsch, 251; Gaudry, 258-261, 269-282; Geinitz, 287; Geinitz und Deichmueller, 289, 290; Goldfuss, 295, 296; von Huene, 324-327; Huxley, 328; Jaekel, 344, 345; LeRoy, 375; Lloyd, 379; Lortet, 382; von Meyer, 428, 436, 442; Roemer, 534; Traut- schold, 573; Twelvetrees, 576. Africa: Broom, 68-71-73, 74 Asia: Lydekker, 384, 386; Huxley, 333; Woodward, 633- Triassic : North America: Branson, 49; Cope, 106, ill, 121, 157; Emmons, 237; Leidy, 373; Lucas, 383. Europe: Alberti, 4; Burmeister, 80, 81; Fraas, 242- 245, 247a; von Huene, 323; Huxley, 329; Jaeger, 338-34i; Metcalfe, 417; von Meyer, 421, 425, 439; Miall, 453; Owen, 505; Quenstedt, 527; Seeley, 544; Storrie, 560; Ward, 588; Wieder- sheim, 602; Woodward, 631. Spitzbergen: Wiman, 625-627; Woodward, 631; Yakowlew, 641; Seeley, 547. Africa: Broom, 69; Huxley, 330; Owen, 517, 519-520. Asia: Lydekker, 384-391; Owen, 510, 511. Australia: Huxley, 330; Stephens, 554, 555. Jurassic: Europe: Dollo, 226. Comanchean : North America: Marsh, 405; Moodie, 480, 481. Europe: Vidal, 579, 580. Cretaceous: North America: Cope, 128; Hatcher, 314; Lambe, 365, 366; Williston, 6u. Tertiary: Europe: Beyrich, 36; Bieber, 37; Cuvier, 197; Fraas, 246, 247; Gunther, 301; Laube, 367-370; von Meyer, 424, 430, 444, 446, 447; Portis, 525; Scheuchzer, 535; Tschudi, 574-575; Walter- storff, 585-586. Pleistocene: North America: Brown, 76; Wheatley, 598. South America: Lydekker, 393. Asia: Clark, 103; Lydekker, 393; Owen, 506. Europe: Lydekker, 393. II. Phvlogenv of Amphibia. Arldt, 10; Baur, 22-25, 3'i Boulenger, 44; Branson, 49; Budgett, 79; Cope, 153, 166, 173; Davison, 198; Dollo, 228; Gadow, 256; Gaudry, 280, 282; Gregory, 299, 300, 300a; Haeckel, 312; Huxley, 337; Kingsley, 358; Moodie, 4896; Owen, 512; Pollard, 524; Quenstedt, 527; Thevenin, 567-568; Versluys, 578; Vogt, 581; Wiedersheim, 601. III. Ontogeny of Amphibia. Ammon, 7; Credner, 187; Fritsch, 251; Hay, 315; von Meyer, 422; Thevenin, 566. 18 INDEX TO THE BIBLIOGRAPHY OF FOSSIL AMPHIBIA. 219 IV. Structure and Morphology of Amphibia. I, at oral-line System of Sensory Organs: Andrews, 8;Baur, 31 ; Branson, 49; Fraas, 242, 247a; Huxley, 333; Mal- branc, 401 ; Miall, 450; von Meyer, 436; Moodie, 458, 465, 478, 488; Thevenin, 568. Pineal Eye: Cope, 160; Credner, 187; Jaekel, 346. Morphology of the Skull: Albrecht, 5; Baur, 24, 27, 29, 31; Boulenger, 44; Branson, 49; Broili, 55, 56, 66: Case, 85, 93, 98; Cope, 136, 177; Credner, 187-191; Fraas, 242, 247a; Fritsch, 251; Fuchs, 255; Gaupp, 283; Goodrich, 298; Jaekel, 347; Maggi, 397. 398: von Meyer, 428, 436, 439; Moodie, 458, 478; Owen, 51 1 ; Seeley, 546; Thevenin, 568; Thyng, 570; Willis- ton, 612, 614, 615, 616, 619; Wiman, 626; Wood- ward, 631. Brain: Wiedersheim, 602; Moodie, 487. Ear: Cope, 159; Broom, 75. Eye: Cope, 105, 107; Moodie, 478. Teeth: Broili, 58; Credner, 194; Fraas, 242; Fritsch, 251; Jaekel, 342; Owen, 502, 503, 504, 505; Seeley, 548; Strickler, 564; Tomes, 571-572; Williston, 608; Wy- man, 637. Gills: Budgett, 79; Credner, 187; Cope, 123; Fritsch, 251; Gaudry, 268; Thevenin, 566-568; von Meyer, 436- Vertebrae and Ribs: Baur, 25, 26; Broili, 61; Case, 98; Cope, 133, 136, 142, 148, 153; Credner, i79-<93: Fraas, 242; Fritsch, 251; Gadow, 256; Gegenbaur, 284; Jaekel, 344, 348; Lillie, 378: Marsh, 404: von Meyer, 436; Mivart, 456; Moodie, 468; Schwarz, 540, 541; Thevenin, 565-568; Williston, 620. Pectoral and Pelvic Arches: Broili, 62; Broom, 72; Cope, 161; Credner, 180-185; Fritsch, 251; Gregory, 299, 3000; Gegenbaur, 285; Jaekel, 347; Moodie, 478; Williston, 610, 616. Limbs, Carpus, and Tarsus: Baur, 22, 23, 28; Cope, 161 ; Credner, 180-185; Emery, 236; Fritsch, 251; Greg- ory, 299-3000 ; Jaekel, 347 ; Wiedersheim, 601 ; Zwick, 645- Clasping Organs: Barkas, 16; Blanchard, 39; Braun, 52; Hilton, 319; Moodie, 461; Fritsch, 251; Stock, 561; Newberry, 498. Muscles: Moodie, 464. Alimentary Canal : Moodie, 47 1 , 474, 478. Integument, Scales, and Structure of Bone: Jaekel, 343; Moodie, 464, 465, 478; Broili, 62; Dawson, 208. Footprints: Barrell, 21; Branson, 50; Brodie, 54; Butts, 82-83; Dawson, 207, 208; Fritsch, 251 ; Geinitz, 288; Hickling, 318; King, 356-357; Lea, 371, 372; Leidy, 374; Marsh, 406; G. F. Matthew, 408-413; Owen, 504; Moodie, 465; Pabst, 521; Smith, 549; Wood- worth, 636. INDEX. Page. Aistopoda Si 77 Alimentary Canal 25, 58 Amblyodon 178 A. probleraaticum 179 American Museum 1,8 Ames Limestone 12 Amphibamus 127 A. grandiceps 2, 7, 15, 128 A. thoracatus 132 Amphibamidae 127 Amphibia 3 Classification 46 Definition 49 Discovery in Carboniferous 6 Geographic and Geologic Distribution. ... 9 History of Classification 39 Anaschisma, Lateral Line System of 35 Anisodexis 164 Anthracosauridaa 187 Apoda 33 Archegosaurus 1, 7, 35 Arm 29 Atlas 27 Axis 27 Baphetes 188 B. minor 189 B. planiceps 6, 188 Baur, George 39 Brachydectes newberryi 177 Branchiosauria 4, 50 Branchiosauridas 51 Branchiosaurus 53 Branson, E. B 38, 192 Brown, N. H 12 Budgett, J. S 78 Cannelton Slates 10, 15 Carr, J. C 13, 14, 22 Case, E. C 9, 182 Caudata 67 Cephalerpeton ventriarmatum 133 Cercariomorphus parvisquamis 139 Clepsydrops Shales 9 Cocytinidas 67 Cocytinus gyrinoides 68 Cope, E. D 1,7,9,34.42,43, 131,207 Credner, Hermann 1 , 24, 52 Cricotidae 180 Ctcnerpeton alveolatum 168 Dawson, Sir J. W 7, 8, 19, 20, 32, 66, 194, 197 Dean, Bashford 1, 32, 187 Dendrerpeton I9> 194 D. acadianum 194 D. oweni 196 Page. Dermal Appendages 197 Devonian 10, 37 Diceratosaurus 8, 1 1 6 D. lajvis 121 D. punctolineatus 117 D. robustus 123 Diplocaulia 34 Dromopus 37, 201 D. aduncus 37 D. agilis 201 Eobaphetes 191 E. kansensis 192 Eosauravus 1 , 87, 1 76 Eosaurus acadianus 7, 1 89 Eoserpeton tenuicorne 1 24 Erierpeton branchialis 69 Erpetobrachium mazonensis 152 Erpetosaurus 97 E. acutirostris 108 E. minutus 105 E. obtusus 99 E. radiatus 98 E. sculptilis 106 E. tabulatus 101 E. tuberculatus no Eryopidae 182 Eryops 23, 34, 182 Euamphibia 46, 49 Eumicrerpeton parvum 57 Eurythorax sublasvis 1 72 Eye 25,131 Footprints 15, 201 Fritsch, Anaton 5, 54, 77, 78 Fritschia curtidentata 85 Gergens, Dr 6 Gurley, W. F. E 9, 12 Hay, O. P 8, 129 Hussakof, Louis 1, 16, 172 Huxley, Thomas H 3, 137 Hylerpeton 83 H. dawsonii 83 H. intermedium 84 H. longidentatum 84 Hylonomidae 79 Hylonomus 79 H. latidens 81 H. lyelli 80 H. multidens 81 H. wymani 81 Hylopus logani 6 Hyoid 25 Hyphasma laevis 70 220 INDEX. 221 Ichthycantlmte '73 Ichthycanthus 1 73 I. ohiensis 1 73 I. platypus 174 Ichthyerpeton squamosum 137 Ja:kel, Otto 8, 118 Joggins, The South (Nova Scotia) 19 Kammplatten 5 Kittanning Coal 15. l6 Lacoe, R. D. (Collection of) 1 Lateral Line System 32 Leg 30,95 Lepospondylia 76 Lcptophractus 169 L. dentatus 17l L. lineolatus I71 L. obsoletus 169 Linton, Ohio, Coal Measures 16 Amphibia of 18 Logan, William 6 Louisville, Kansas 10 Lydckker, Richard 44, 77 Lyell, Charles 19. 20 Macrerpetida; 183 Macrerpeton '84 M. deani lg6 M. huxleyi 184 Mandible 25, 103 Marsh, 0. C 10, 37. 189, 202 Mastodonsauridx 200 Mastodonsaurus sp. indet 200 Matthew, G. F 8, 22 Mauch Chunk 37 Mazon Creek Shales 7, I2 Amphibia of 13 Mazonerpeton 61 M. costatum 63 M. longicaudatum 61 Meyer, Hermann von 6 Micrerpeton 51 M. caudatum 52 Microsauria 4, 34, 7° Relation to Reptilia 77 Mississippian Amphibia 37 Molgophidae '49 Molgophis '-W M. brevicostatus '5° M. macrurus '49 M. wheatleyi '51 Morphology of Coal Measures Amphibia 23 Muscle 32 Myocommata 3° Nccturus 33. 56 Newberry, J. S. (Collections of) 1, 8, 17, 18 Nyraniids J37 CEstoccphalus 145 O. rectidens 147 0. remcx 145 Operculum 172 Osage City, Kansas 10 Palate 24, 104 Parabatrachus 7 Pectoral Girdle 29, 107 Pelion 7, 72 P. lyelli 73 Pcliontida? 72 Pelvic Girdle 29 Phlegethontia 155 P. linearis 156 P. serpens 156 Phcenix Tunnel, Pennsylvania 10 Pholidogaster 3 Pitcairn, Pennsylvania 12 Platystegos loricatum 199 Pleuroptyx clavatus 12, 19, 153 Proteida 67 Proterpeton gurleyi 12, 178 Ptyoniidae H1 Ptyonius I41 P. marshii 143 P. nummifer 144 P. pectinatus 141 P. serrula 144 P. vinchellianus 143 Raniceps. Ribs 7 27 Occiput Odonterpeton triangularis . 25 in Salientia 72 Saurerpeton latithorax 165 Sauropleura '57 S. digitata '59 S. (Anisodexis) enchodus 164 S. foveata l63 S. longidentata l62 S. newbenyi l6° S. pauciradiata '6° S. scutellata '58 Sauropleuridae 3°. '57 Scales 31.197 Schwarz, Hugo 8> 28 Sclerotic plates 24 Scutes ,62 Skin l6^,97 Skull 23 Smilerpeton acicdentatum 82 South Joggins, Nova Scotia 7. '9 Sparodus sp "™ Spondylerpeton spinatum ' 8' Stegopida; ' I3 Stegops "3 S. divaricata ' '4 Stereospondylia 5. 34. 200 Sternum 3° 222 INDEX. Tarsus 1/6 Teeth 25 Temnospondylia 5, 34, 180 Thinopus antiquus 10, 37 Thyrsidium fasciculare 147 Traquair, R. H 5 Triton walthi 50 Tuditanidas 86 Tuditanus 86 T. brevirostris 89 T. longipes 90 T. minimus 92 T. punctulatus 87 T. walcotti 94 Twin Mounds, Kansas 22 Udden, J. A n U record yliida? 116 Ventral Scutellai 30 Vertebrae 27, 181 Vertebral Column 27 Wiedersheim, Robert 30 Williston, S. W 2, 10, 200 Wyman, Jeffries 7 Zamenis flagellum 5 THIS BOOK IS DUE ON THE LAST DATE STAMPED BELOW RENEWED BOOKS ARE SUBJECT TO IMMEDIATE RECALL MAR " |yyg LIBRARY, UNIVERSITY OF CALIFORNIA, DAVIS Book Slip-50m-12,'64(F772s4)458 373008 Moodie, R.L. The coal measures Amphibia of North America. QE867 M?7 PHYSICAL SCIENCES LIBRARY LIBRARY UNIVERSITY Of CALIFORNIA DAVIS 373008 Moodie, R.L. The coal measures Amphibia of North America Call Number: QE867 M57