The coal measures Amphibia of North America

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;

</, body impression. X 3.

B. Eumicrer peton parvum Moodie. o, anus;/, femur; h, humerus; in, intestine; /, liver; st, stomach;

r, radius; u, ulna. X 3.3.

C. Larger specimen of Eumicrer peton parvum Moodie. a, anus; d, dorsal lateral line; h, humerus;

in, intestine; ml, median lateral line; si, stomach. X 2.6.

D. Skeleton of Erpetobrachium mazonensis Moodie. cl, clavicle; h, humerus; r, radius; sc, scapula;

11, ulna. X 2.

E. Rib of Mazonerpeton costalnm Moodie. X 2.5.

Originals in the Yale University Museum.

In another nodule (No. 804, Yale Museum) there is a single bone preserved which
resembles, to a great extent, a rib of the present species (fig. 15, E), although some-
what larger, and it has been provisionally identified as such. The element is very
slightly curved, but shows the expanded head of the rib of this species.

66

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

Measurements of the Type of Mazonerpeton costatum Moodie.

No. 800 (777), Yale University Museum: mm-

Length of portion of skull preserved 14

Length of right clavicle 16

Width of right clavicle 4

Length of dorsal region represented 30

Length of cervical rib 8

Length of dorsal rib 6.5

Length of caudal rib 3

Length of caudal portion of body preserved. . . 55

Length of mandible 15

Greatest width 6

Length of right humerus 10

Greatest width of humerus 2

No. 804 (332), Yale University Museum:

Length of rib 11

Width of head of rib 2

Diameter of shaft I

Sparodus sp. (?).

Dawson, Phil. Trans. Roy. Soc. London, pt. 11, p. 643, pi. 40, figs. 52 to 56, 1882.
Dawson, Proc. and Trans. Roy. Soc. Canada, xn, p. 75, 1895.

Type: Specimen in the Peter Redpath Museum of McGill University.

Horizon and locality: Coal formation at the South Joggins, Nova Scotia.

The material on which the above determination is based was collected in 1878
by Sir J. W. Dawson in the coal formation at the South Joggins, Nova Scotia.
Nothing has been collected since that date
that would give additional information as
to the nature of the form represented. I
give here Dawson's description of the
remains :

"In the coaly matter or mineral charcoal
at the base of tree No. 10 appeared a few frag-
ments of an animal which may possibly belong
to the above-named genus of Fritsch, though
I am by no means certain of this identification
or of the real nature of the animal.

"The skull is represented by a fragment of a maxillary or intermaxillary bone, with
blunt conical teeth. It is smooth or marked merely with microscopic dots. There is also a
fragment which may be a palatal bone studded with minute teeth.

"A few vertebrae associated with the above bones are long and narrow, with large zyga-
pophyses and long neural spines. Length of body (i.e., of the vertebra) about 3 millimeters.

"With these remains are a few bony scales different from those of any other species
found in these trees, and more resembling scales of Ganoid Fishes. They are somewhat
rectangular in form, enameled on the surface and beautifully sculptured with waving lines.

"In the same trunk were found some teeth and bones referable to Hylerpcton dawsoni,
and it is not impossible that the remains above referred to may have belonged to some crea-
ture devoured by that animal, and which would not otherwise have obtained admission to
the interior of an erect tree. The tree itself had been removed by the sea, all but a little
of the base, and this was in a very unsatisfactory state, so that doubt might even exist as to
the limit between the deposit in the interior of the tree and that under its base."

Fig. 15a. — Type material of Sparodus, consisting of a,
a tooth (X 25); b, four of the smaller teeth (in
maxilla?) (X 25); c, three bony scales (X 5); d,
fragment of a limb bone (X 2); e, avertebra(X 2).
(After Dawson.)

CHAPTER X.

ORDER CAUDATA DUMERIL, 1806. COAL MEASURES TO RECENT.

Naked-skinned, elongate, tailed salamanders, mud-puppies, efts, newts, etc.
External gills present or absent in adult condition, but always present in young.
Limbs short, with usually 4 digits on hand and 5 on foot, but this is subject to
much difference. Limbs never very stout. Carpus and tarsus cartilaginous. Skull
roof without the postparietal, postorbital, and supra temporal. Skull elements
never ornamented and never cut by the lateral-line canals. Vertebrae consisting
of a single element ; ribs short, attached to an elongate transverse process. Caudal
ribs seldom present. Parietal foramen lacking. No ventral armature. Fresh-
water inhabitants.

Suborder PROTEIDA Cope, 1868.

This order agrees generally with the Caudata, but presents one most important
feature of difference in the presence of the opisthotic. It is this point which gives
the Proteida its intermediate position between the extinct amphibians and the recent
species, and seems to indicate a connecting line from the Coal Measures down to
the present. The structure of the hyobranchial arches sustains this view.

The hyoid apparatus differs from that of other adult Caudata and resembles
that of their larvae in having three epibranchials, instead of one only. The second
basibranchial is also connected with the first, which is not the case with the other
Caudata. Three extinct genera are placed tentatively in this suborder.

Family COCYTINID.dE Cope, 1875.

Cope, Bull. U. S. Nat. Mus., No. 1, p. 12, 1875.

The present family, as here defined, includes the forms whose structure seems
to ally them with the modern salamanders. The character on which most depen-
dence is placed is that of the branchial apparatus, lacking in Hyphasma. The forms
are all incompletely known and the family will doubtless require revision on acqui-
sition of additional material. Three genera, each with a single species, are:

Cocytimts gyrinoides Cope. Linton, Ohio, Coal Measures. Based on the ventral impression of the

skull, with the well-developed branchial apparatus.
Erierpeton brattchialis Moodic. Mazon Creek, Illinois, shales. Based on impression of mandibles

and branchial apparatus.
Hyphasma Icevis Cope. Linton, Ohio, Coal Measures. Based on incomplete and obscure amphibian

body, lacking limbs.

Genus COCYTINUS Cope, 1871.

Cope, Proc. Am. Phil. Soc, 1871, 177.
Cope, Geol. Surv. Ohio, 11, pt. 11, 360, 1875.

Type: Cocytinus gyrinoides Cope.

Vertebra? and ribs osseous; teeth on the premaxillary bone, none on the maxil-
lary; hyoid elements largely developed, an axialhyal with basihyal on each side,
closely united with the corresponding ceratohyal, at the end of which is an element
in the position of a stylohyal; haemal or basibranchials 3, the anterior 2, each sup-
porting 1 pleural branchihyal, and the third supporting one also, the first haemal
branchihyal on the inner side of the ceratohyal, approaching the median line, and
with elongate pleural element.

67

68

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

">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.

L<i DECKER, Cat. Fossil Rcptilia and Amphibia, pt. iv, p. 196, 1890.
Moodie, Bull. Am. Mus. Nat. Hist., xxvi, art. xxv, p. 357, 1909.

Type of the family: Urocordylus wandesjordii Huxley.
Locality and horizon: Coal Measures of Kilkenny, Ireland.

Stout and long-tailed forms, with the tabulare cornua frequently much pro-
duced and pitted cranial bones; lateral lines well-developed; palate with teeth on
palatines, vomers, premaxillas, and maxillae, the two latter elements bearing conical
teeth and the others bearing short, stumpy cones, at least in one species; pineal
foramen well forward; nostrils and orbits in the anterior part of skull; scapula
peculiarly curved and pointed; other pectoral elements sculptured; neural spines
and chevrons of caudal vertebrae much dilated at their extremities, and pectinated ;
no caudal ribs ; vertebra in one genus apparently capped with a sculptured plate
as in Zatrachys; tail very long and tapering to a point, 50 to 80 caudal vertebrae;
dorsal region short; limbs well developed, with clawed digits; carpus and tarsus
cartilaginous; endochondrium well formed.

There are 4 genera which constitute this family: Urocordylus, from the Coal
Measures of Ireland; Ceraterpeton, from the Coal Measures of Ireland and Eng-
land; Diceratosaurus, from the Coal Measures of Linton, Ohio; Eoserpeton, from
the Coal Measures of Linton, Ohio.

These may be distinguished by the following characters :

I. Skull triangular, truncated behind, with rounded muzzle and aborted tabulare cornua, neural spines of
caudal vertebrae long, slender, and expanded in a fan-like manner; tail with about 80 vertebra;;
ventral scutes oat-like Urocordylus.

II. Skull parabolic and of great width, with short cornua projecting from the supratemporal ; tabulare cornua
nearly twice as long; neural spines of caudal vertebrae low and wide; ventral scutes oblong; caudal
vertebras about 50 Ceraterpeton.

III. Skull broad with obtuse snout, tabulare cornua absent, large, pointed posterior expansions from supra-

temporal, posterior table within the cornua truncate; vertebrae with an apical sculptured plate;
caudal vertebrae numerous, over 75; ventral scutellae bristle-like, arranged en chevron . . Diceratosaurus-

IV. Skull a broad oval, with large posterior projecting supratemporal horns, posterior table of skull between

cornua truncate without the small lateral projection from the supratemporal, orbits a long oval,
ribs long, curved and slender, tail unknown, possibly shorter than in other members of the
family; it is restored as short in Journal of Geology, XVII, p. 77, fig. 20, 1909, but this is uncertain;
the skull has all the characters of the family Eoserpeton.

The relationships of the family are not far to seek. They fall in immediately
with the Amphibamidas and Hylonomida? in being among the most reptile-like of
the Paleozoic Amphibia. The group is, however, distinctly amphibian in the pos-
session of 4 fingers, with the usual microsaurian phalangeal formula.

Genus DICERATOSAURUS Jaekel, 1903.

Jaekel, Ncucs Jahrbuch f. Mineral., Geol. u. Palcon., Bd. 1, p. 112, 1903.
.Moodie, Jour. Geol., xvil, pp. 63-69, figs. 13-15, 1909.

Type: Diceratosaurus punctolineatus Jaekel.

Orbits in the anterior two-thirds of the axial skull length, nostrils near to
the anterior end of the skull; pineal foramen in the center of the skull roof; skull

115

Il6 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

provided with tabulare cornua and a broad backwardly directed process ; quadrate
angle does not project on the border of the skull ; sculpture of the cranial elements
impressed as radial grooves; 12 presacral vertebrae, 1 sacral with expanded neural
spine which is sculptured at the top, with simple long, apparently separately ossi-
fied transverse processes ; extremities small ; foot with 5 digits ; phalangeal formula
2-3-3-4-3-

The most important differences between Diceratosaurus and Ceraterpeton, the
most nearly allied genus, is (in Diceratosaurus) in the more anterior position and
small size of the orbits, the backward extension of the quadrate region, and the
dorsal expansion of the vertebral spines. A further, and more important, difference
between the genera is in the location of the backwardly directed processes from the
skull. In Ceraterpeton they project backward from and are a portion of the tabulare
element, while in Diceratosaurus the projection consists almost entirely of squa-
mosal and supratemporal.

Diceratosaurus punctolineatus Cope.

Cope, Proc. Phila. Acad. Nat. Sci., 1875, p. 16.

Cope, Geol. Surv. Ohio, 11, pt. n, p. 372, pi. xli, fig. 4, 1875.

Moodie, Bull. Am. Mus. Nat. Hist., xxvi, art. xxv, p. 356, pi. lxv, 1909.

Type: Specimen No. 8606, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The species was first described by Cope as Ceraterpeton punctolineatum (122).
It was redescribed on more abundant material (plate 19) by Jaekel (347), and the
following is taken from the discussions of these two authors, checked by my own
observations on the type specimen. This shows a portion of the skull, consisting
of the squamosal and supratemporal with a projecting, convergent horn. The
sculpturing on the skull is similar to that on the pectoral plates, of which there are
three preserved (plate 14, fig. 4). The bones of the fore limbs are stout and
short. The ribs are only slightly curved. The character of the vertebrae can not
be ascertained. The sculpturing of the bones consists of radiating ridges, grooves,
and pittings.

Jaekel (347) described from the museum collection at Berlin 3 specimens of
this species, among which were 2 skulls. There were associated with these remains
some pectoral plates and limb bones with a nearly complete series of vertebras. A
modified translation of Jaekel's description follows:

The skull of the largest specimen has, including the horns, a length of 35 mm.
and a width of 30 mm. on the occipital border. From the anterior end to the pos-
terior border of the skull (exclusive of the horns) there is a length of 25 mm. The
pineal foramen lies about midway of this length. The orbits are rather large, al-
most circular, and lie about midway between the pineal foramen and the anterior
border of the skull. The nostrils, which lie anterior to the orbits, are a rather oblique
oval and narrowed on the lateral ends. The distance between them is about the
same as that between the orbits, which measures 7 mm.

The skull roof is sculptured with pits much like those of Archegosaurus. The
larger the bones, the rougher the sculpture. The bones of the middle of the skull
(that is, the parietals, frontals, premaxillae, nasals, and postparietals) are of the

MOODIE

PLATE 16

I • Type specimen of Diceratosaurus punctolincatus Cope, from Linton, Ohio, beds. X I . Drawn from photo-
graph. r/ = claviclc; A = humerus; ic = in terclavicle; mc = metacarpals; r-u = radius-ulna; r6 = rib;
sc = scapula; sq= squamosal; vs = ventral scutella?.

2. Skull of Sauropleura longidentata, Moodie, from the Coal Measures of Linton, Ohio. X I. Drawn from

photograph, fr = frontal ; m = maxilla ; n = nasal ; or - orbit ; par = parietal ; pp = postparietal ; tab = tabulare.

3. Mandible of Sauropleura longidentata, Moodie, from Coal Measures of Linton, Ohio. X i-5- Drawn

from photograph.

4. Type specimen of Sauropleura enchodus Cope, from the Coal Measures of Linton, Ohio. X 1.

5. Additional specimen of Diceratosaurus punctolineatus Cope, from the Coal Measures of Linton Ohio.

X 1. '= femur; j'/ = ilium; mc = metacarpal; mt = metatarsal; pA = phalanx; r = radius; < = tibia.

THE MICROSAURIAN FAMILY UROCORDYLID^E. 117

normal form and only show an unusual difference in that they are large in the
inverse order. The pineal foramen lies in the anterior third of the parietals ; that is
the primitive condition which occurs in the young forms of the Branchiosauridae.
The nostrils are inclosed by the premaxillae in front, in the median line by the nasals
and laterally by the maxillae. The jugals, by their backward prolongation, form,
behind the maxillae, the border of the skull, and only attain to some size on the
upper side of the cranium.

Jaekel's " perisquamosal " (see plate 15), which is of a doubtful nature, is not
indicated in the type specimen nor in the specimens of the other two species (462)
assigned to this genus. In D. robustus and D. Icevis the "perisquamosal" region is
easily separable into its component elements. The sutures between the elements
may have been indistinct in his specimen, but it is hardly conceivable that a union
of the skull bones would occur in one species and not in another of the same genus.
Jaekel's suggestion (347) that "Etwas mehr Wahrscheinlichkeit mochte ich der
Vorstellung beimessen, dass diese Ausbreitungen zum Schutz freier Kiemen dienten,
wie sie z. B. bei den Perennibranchiaten als baumformige Organe weit am Halse
herausragen" can hardly find acceptance with students of the Paleozoic Amphibia,
since there is not the slightest evidence that the Microsauria ever possessed external
gills and considerable presumptive evidence that they did not. His comparison
of the "perisquamosal" to the "Kiemendeckel" of the fishes is also very unhappy
on morphological grounds, since the elements of his "perisquamosal" form constit-
uent parts of the skull roof, which the operculum never does.

The palate of the skull (plate 15, fig. 2) has been determined by removing the skull
bones of one specimen. Anteriorly the premaxillae and maxillae are clearly recogniz-
able as large dentiferous elements. The premaxillae have 4 to 7 teeth, the short max-
illa has 3 to 4. All the teeth are of nearly equal size. Smaller teeth seem to be
indicated by impressions found between the larger ones. The vomers, which are
tolerably large, unite with the premaxillae behind and inclose at least half of the
palatine foramen on the inner side. They are furnished with small teeth, which in
the anterior part are very irregularly placed, but they are more regular posteriorly.

The palatines and transverse bones are questionably identified. They seem to
lie posterior and lateral to the vomers, but the sutures are indistinct. The large
parasphenoid seems well displayed and is more or less heart-shaped. There would
seem to be a slight indication of double occipital condyles. The pterygoids are
broad plates which inclose the parasphenoid and form the lateral boundary of the
palate. The cotyli are very indistinct, but appear as elongate grooves.

The pectoral girdle (plate 15, fig. 3) consists, apparently, of seven elements, three
paired and one unpaired. The unpaired element (the interclavicle) is truncate pos-
teriorly and acuminate in front, with its surface radially grooved and the anterior
borders beveled for articulation with the clavicles. The clavicles are triangular, as is
usual with the Microsauria. They are sculptured with radiate grooves and ridges,
with decided inosculations at the ossific center. The coracoids have only part of
their surface ornamented ; most of their surface is smooth for articulation with the
interclavicle and scapula. A long spine projects from the inner surface of the cora-

H8 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

coid. A pair of small elements lie one on either side of the clavicles and Jaekel (347)
interprets them as the cleithra. If they are cleithra they are unique among the
Microsauria. The pectoral girdle does not, however, indicate that Diceratosaarus
is "unique among all known quadrupeds."

Jaekel regards the limb which is preserved with the material as an arm (plate
15, fig. 4) , but there is no reason stated for his conclusion. It has all of the characters
of the leg and may be regarded as such. Only a part of the lower end of the femur is
preserved. The tibia and fibula are preserved as separate rod -like elements with one
of the bones longer and larger, probably the tibia. There are 5 toes which have the
customary phalangeal formula for a microsaurian foot of 2-3~3-4(?)-3. The
tarsals are unossified.

The vertebrae (plate 15) were perforated for the notochord, and are hour-glass-
shaped, with the neural arch thickened to support a heavy spine which bore a sculp-
tured plate. These apical plates occur in the dorsal region, but diminish toward the
caudal vertebrae. The number of the vertebrae in the dorsal region is very small and
in the tail very large. There are possibly 2 vertebrae in the cervical, 1 1 in the dorsal
series; the thirteenth carries the pelvis. There are over 100 vertebrae in the tail.

The ribs have an expanded head and the transverse processes of the vertebrae
are long.

The following account is based on the writer's study of the type specimen and
he is able to add several points of interest to a knowledge of the anatomy of the
type of this interesting microsaurian.

The type specimen consists of 1 1 consecutive vertebrae with a portion of the
skull, the greater portion of the pectoral girdle, parts of both fore limbs, ribs, and
ventral scutellae (plate 14, fig. 4). The species is represented in the collection by yet
another specimen, on which Cope based his Tuditamis mordax (plate 22, fig. 5), of
which he himself says : ' ' Further examination of the specimen on which the latter
(T. mordax) was founded leads to the belief that it is an imperfect cranium of Cerater-
peton (Diceratosaurus) punctolineatum Cope." The plates referred to are rather to be
regarded as elements of the pectoral girdle and I believe they represent the clavicle
and a portion of the interclavicle.

The skull of the present species is fully described by Jaekel. The type specimen
does not offer any evidence in support of Jaekel's "perisquamosal,'.' but rather
tends to the idea that he is incorrect in his assumption of the fusion of these elements
of the skull. The direction taken by the ridges and grooves on the elements pre-
served indicate a separation between the supratemporal and the squamosal. I do
not find that the grooves have the tendency to arise from a common center of ossi-
fication in the squamosal, as suggested in the figures of Jaekel. The horn which
projects backward from the squamosal is rather large and heavy for the size of the
skull, and after curving slightly inward ends in a blunt point and not sharply, as
Jaekel figures in his specimens. The vertebral column is indistinctly preserved and
I have nothing to add to Jaekel's account given above.

In the structure of the pectoral girdle my results are greatly at variance with
those of Jaekel. I do not find the remarkable elements which Jaekel has figured

THE MICROSAURIAN FAMILY UROCORDYLID/E.

119

(347) in his specimens. On the other hand, I find a normal microsaurian pectoral
arch (464), such as has been described for numerous other forms. There are present,
distinctly preserved in the type specimen, the scapulae, the clavicles, and the interclav-
icle, with the possibility of the coracoid. The peculiar element referred to by Cope
as resembling a "lacertilian pubis" is without doubt the left scapula of the animal
(plate 16, fig. 1 ) . Its form compares very favorably with that of Ceraterpeton as fig-
ured by Woodward (630). The coracoid may be represented by the fragment which
lies close to the scapula. The sculptured element lying next to the supratemporal
horn of the skull is the right clavicle preserved bottom side up. Of the other two
sculptured elements, one is the interclavicle, only a portion of which is preserved.
The left clavicle lies beside it. The clavicles in this species have a tendency to
assume the triangular shape so common in other species of Microsauria, and the

Fig. 24.

A. Skull of Diceratosaurus lozvis Moodie, from the Linton Coal Measures. X I. /.frontal:
j, jugal; mx, maxilla; n, nasal; or, orbit; par, parietal; pof, postfrontal; po, postorbital;
pf, prefrontal; pp, postparietal ; sq, squamosal; spt, supratemporal; qj, quadratojugal ;
pmx, premaxilla; tab, tabulare.

B. Reconstruction of skull outlines of Diceratosaurus robuslus Moodie, from the Coal

Measures of Ohio. X 0.75. fr, frontal; 7, jugal; or, orbit; par, parietal; pof, postfrontal;
po, postorbital; pp, postparietal; qj, quadratojugal; spt, supratemporal; tab, tabulare.

interclavicle, so far as can be determined, was shield-shaped. The upper sur-
faces of the pectoral elements are marked by grooves for the attachment of the
pectoral muscles.

The ventral scutellation is present in a small patch (plate 16, fig. i) near the
horn of the skull. The scutse are oat-shaped and take the usual form. The ribs
are not long, are rather stout, and beyond the proximal curve are nearly straight to
the obtuse tips. The heads of the ribs are so obscure that it is impossible to deter-
mine whether they were two-headed or not. They are expanded proximally and
there is a slight tendency to a division of the head.

Portions of both fore limbs are preserved. The right limb possesses the hume-
rus, separate radius and ulna, and 2 metacarpals. The other possesses only the
radius, 3 metacarpals, and a portion of a phalanx. The humerus is a very stout
bone and at once recalls that of Amblyrhynchus. The ends are expanded and there

120 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

are roughnesses on the bone for the attachment of muscles. The radius and ulna
are subequal in size. They are both expanded more proximally than distally. The
carpus was cartilaginous. An additional specimen of this species is figured on plate
1 6, fig. 5. This adds to our knowledge of the pelvis especially.

Measurements of the Type.

mm. mm.

Length of entire specimen 80 Width of various portions of ulna same as radius.

Length of tabulare horn of skull 20 Length of the only phalanx preserved 5

Width at base 4 Length of vertebra 5

Width at tip 2.5 Width of vertebra 4

Length of right humerus 16 Length of longest rib 17

Width at middle of shaft 3 Width of rib at widest part 1.5

Width at proximal end 5 Width of clavicle 18

Width at distal end 5.5 Length of clavicle 20

Length of radius 9 Length of interclavicle 25

Length of ulna 9 Width of interclavicle 16

Width of radius at proximal end 2.5 Length of single side of chevron scute 7

Width at middle 1.5 Width of same 25

Width at distal end 2

The specimen (No. 2566, Am. Mus. Nat. Hist.) on which Cope based his
Tuditanus mordax is composed of two plates of the above-described species.

Diceratosaurus laevis Moodie.
Moodie, Jour. Geol., xvn, No. I, p. 63, figs. 13, 14, 1909.

Type: Specimen No. 102 (8680 G), American Museum of Natural History,
where it forms part of the Newberry collection.

Horizon and locality: Linton, Ohio, Coal Measures.

The species is represented by an almost complete skull, which had been identi-
fied previously by Cope as Tuditanus radiatus. The specimen consists of the
impressions of the bones of the cranial roof, the bones themselves having disap-
peared. It is not probable that the dorsum of the skull was smooth. The details
in the structure of the skull have been ascertained quite definitely. There can be
no doubt that the arrangement of the elements is accurate, as shown in figtire 24 A.
The supratemporal, as in Erpetosaurus tabulatus Cope, is excluded from the parietal.

The form of the skull at once recalls that of the species D. punctolineatus, as
figured by Jaekel (see plate 1 5) . The orbits are located in nearly the same region
of the skull and the sutures separating the cranial elements are quite similar in the
anterior portions. The species D. loms is based on the divergent character of the
horn-like protuberances which project from the squamosals. The horns of D,
punctolineatus are convergent. The present skull is also smaller and the parietals
in D. leans are much larger than in the type species. In the type species, also, the
pineal foramen is located well forward in the parietal, while in the present form the
foramen is located well posterior.

The skull is almost rectangular. The nostrils are elongate ovals. The orbits
are circular and the distance between them is equal to two-thirds of the dimensions
of the orbit. They are located well forward in the skull and are bounded laterally
by the maxillaries. The nostrils have much the same character as in the type form,
being broadly oval.

The premaxillae are elongate transversely, being about twice as long as wide.
They are identical in shape and relations with the same elements in D. punctoline-
atus Cope. The nasal is nearly square and forms the interior boundary of the nos-
tril. The frontal is elongate in the median length of the skull and it is acuminate

THE MICROSAURIAN FAMILY UROCORDYLIOE. 121

posteriorly, where the acumination is inclosed by the parietal and postfrontal. The
parietals are by far the largest elements in the cranium. They form together an
oval which is elongate in the longitudinal diameter of the skull. They inclose
between them, in the median suture, the small pineal foramen. They are acu-
minate in front, with a broad truncate posterior base, where they are bounded by the
postparietals. The postparietal is nearly square, being somewhat wider than long.
It joins the tabulare and the parietal. The tabulare is elongate in the long diame-
ter of the skull. It ends anteriorly in a point which is inserted between the post-
orbital and the parietal, and bears a short protuberance posteriorly, much as does
the same element in the type species.

There are four elements which take part in the formation of the posterior border
of the skull. These are the postparietal, the tabulare, the squamosal, and the
supratemporal. It is very unusual for the supratemporal to reach the posterior edge
of the cranium. The prefrontal lies anterior to the orbit, of which it forms the ante-
rior border. The lacrimal has not been detected, although Jaekel (347) has indi-
cated it in his drawings of the skull of the type species. The maxilla is elongate and
forms the lateral border of the skull. No teeth have been detected, although they
were doubtless the same as Jaekel has figured in D. punctolineatus. The jugal is an
elongate element joining the maxilla posteriorly. Jaekel included this element in
his "perisquamosal," but the sutures are clearly evident in the present specimen
and there is no evidence of a structure at all similar to a "perisquamosal." The
postorbital is fully as large as the jugal which it joins, forming a part of the poste-
rior border of the orbit and ending posteriorly in a point which is inclosed by the
tabulare and the squamosal. The postfrontal with the foregoing element forms
the entire posterior border of the orbit and it likewise ends in a point inclosed by
the parietal and the postorbital. The quadratojugal has much the same shape and
relations as in D. punctolineatus, although it is located further back. The squa-
mosal is also elongate, as are most of the posterior cranial elements, and it also
has an acumination which is directed forward and is inclosed by the postorbital
and jugal. The anterior suture of this element is rather indistinct, but it is, I
believe, as represented (fig. 24). The element is elongate and is prolonged
posteriorly to form the horn, which ends in a blunt point and is not sharp, as in
the type species.

Jaekel (347) regards the species Dicer atosaur us punctolineatus Cope as being
unparalleled among known vertebrates in the possession of a "perisquamosal" ele-
ment. In closely allied species the "perisquamosal" is easily separated into its
component elements, and the morphology of the present skull would throw con-
siderable doubt on Jaekel's interpretation of the skull of the type species. Another
specimen, described below as another species of this genus, shows no evidence of
this fusion. So far as I can learn, there have been no cases of true fusion of cra-
nial elements correctly reported, unless it be that which possibly exists between
the two frontals in the skull of Diplocaulus. It was on the basis of such fusions
that Maggi (397) proposed to derive the interparietals of the primates from the
tabulare of the stegocephalians.

122 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

The posterior outline of the skull in the present specimen is not well preserved
and the outline as given may be slightly inaccurate. The indentation figured by
Jaekel in the posterior border of the skull of the type form is not present in the
species under discussion.

Measurements of the Type Skull of Diceratosaurus l^evis Moodie.

ram. mm.

Length of skull along median suture 37 Width of skull across the orbits 30

Length from muzzle to tip of horn 50 Interorbital width 6

Width between tips of horns, estimated 40 Length of nostril opening 2

Width of orbit 7 Width of nostril 1

Length of orbit 10 Diameter of the pineal foramen — I

Diceratosaurus robustus Moodie.

Moodie, Jour. Geol., xvn, No. 1, p. 67, fig. 15, 1909.

Moodie, Bull. Am. Mus. Nat. Hist., xxvi, art. xxv, p. 355, pi. lxiii, fig. 2, 1909.

Type: Specimen No. 8611 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The present species is indicated by the left portion of a cranium representing a
large individual. The characters of the skull are so clearly marked that it seems
worthy of description. The presence of horns as given in the restoration of the
skull (fig. 24, B) is based on the analogy with the other two species of this genus, in
both of which horns are present. The generic determination of the species is based
on the large size of the postorbital, which is essentially characteristic of the other
species of Diceratosaurus.

The characters which distinguish the species from others of the genus are the
large postorbitals and the small parietals, which are excluded from union with the
postf rentals on account of the large size of the frontal. In the other two known
species the frontal is small and the parietal comes forward to join the post frontal.
The present species exhibits a skull which is nearly twice as large as that of D.
IcBvis and nearly three times the size of the skull of D. punctolineatus.

The portion of the skull preserved shows the cranium to have had a rather acu-
minate snout, not blunt as in the type species. The orbit is an elongate oval,
although it has the same relative position in the skull as in the other species. The
nostril is indicated by an oval depression near the anterior edge of the skull. The
frontals, as indicated by the sutures present on the portion of the skull which is
preserved, are fully as long as the parietals. Whether they were as wide as is repre-
sented is uncertain. The postfrontals are very small bones, the sutures of which
are somewhat uncertain, although they can not be far from what is represented
(fig. 24, B) . The postorbital is large and elongate, and is distinctive of this species on
account of its unusual size, although it does not attain the same proportions as in
other members of the genus. The parietals are elongate and narrow. The pineal
foramen is represented by its lateral edge and its position is about midway of the
longitudinal diameter of the parietals. The narrow postparietal is represented by
its anterior border; as restored (fig. 24, B) it may be too long. The tabulare, also, is
represented by an anterior portion and it shows this element to have the position
and form which is typical of the form Diceratosaurus Icevis. Such other of the
cranial elements as are indicated are based on the relations discovered in D. Iceins.

THE MICROSAURIAN FAMILY UROCORDYLIOE. 1 23

The heavy line on the left of the drawing (fig. 24, B) represents the outline of the
preserved portion. The skull, as restored, may be a little too long, and the shape of
the horns is conjectural. In the orbit there are preserved 2 misplaced teeth show-
ing longitudinal fluting. The longest tooth is about 3 mm.

Measurements of the Type Skull of Diceratosaurus robustus Moodie.

mm. mm.

Median length of skull, estimated 67 Length of longest tooth 3

Posterior width of skull, estimated 78 Width of same tooth at base 1.5

Length of orbit 18 Length of postorbital 27

Width of orbit 12 Width of postorbital 14

Genus EOSERPETON Moodie, 1909.

Moodie, Jour. Geol., xvn, pp. 76-79, fig. 20, 1909.

Moodie, Bull. Am. Mus. Nat. Hist., xxvi, p. 355, pi. lxiii, fig. 1, 1909.

Type: Eoserpeton tenuicorne Cope.

The genus was proposed for the reception of a single species originally referred
by Cope to Ceraterpeton (C. tenuicorne). The species can not be placed in the genus
Ceraterpeton on account of the form and structure of the skull, which varies widely
from that of the type species, Ceraterpeton galvani Huxley, from the Kilkenny Coal
Measures of Ireland. The most important character in which the present species
differs from C. galvani is the peculiar form taken by the squamosal and by the posi-
tion of the "horn." These characters will be evident on referring to figure 25. No
undoubted remains of Ceraterpeton have been found outside the British Isles. Fritsch
referred (251) a species, previously described as Scincosaurus crassus, to this genus,
but Andrews (8), Jaekel (347) and Woodward (630) all unite in placing the species
in the genus where it was formerly described. Jaekel even says that the Scinco-
saurus has no horns, so far as he can determine. Cope referred 3 species (123)
from the Linton Coal Measures of Ohio to the genus Ceraterpeton, but it has been
shown elsewhere (347) that no one of them belongs in the genus, nor in fact do
they all belong in one genus.

Eoserpeton tenuicorne Cope.

Cope, Geol. Surv. Ohio, 11, pt. n, pp. 372-373, pi. xlii, fig. 2, 1875.
Cope, Proc. Am. Phil. Soc, xxn, p. 407, 1885.
Cope, Proc. Am. Phil. Soc, xxxvi, p. 85, pi. iii, fig. 2, 1897.
Moodie, Proc. U. S. Nat. Mus., vol. 37, p. 23, 1909.

Type : Specimen in the American Museum of Natural History. There are also
specimens Nos. 4472 and 4473 in the U. S. National Museum.

Locality and horizon: Linton, Ohio, Coal Measures.

The species was founded on a complete skull preserved on obverse sides of a
block of coal. Cope (123, pi. xlii, fig. 2) figured this skull. The figure is poorly
executed and does not do justice to the specimen, which is really well preserved.
In general the skull is oval, with the orbits located well towards the acuminate
snout. The interorbital space is equal to twice the width of the orbit. The pineal
foramen lies near the center of the skull. The quadrate angles are drawn out into
slender acuminate, longitudinally striate horns, processes from the squamosal.
The "horn" arises from an expanded base, which is a portion of the cranial element
at the postero-lateral angle of the skull. This character is taken as the distinctive

124

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

one of the genus. It is possessed by no other form of Carboniferous air-breathing
vertebrate, in association with the other characters of the form.

The boundaries of the premaxillse are indefinite, but what remains of the sutures
indicates that the elements were small. No teeth have been detected. The nasal
is likewise not clearly defined, but the frontal is an elongate element which occu-
pies the space between the orbits and joins the parietal posteriorly. The parie-

Measurements of the Tyj>e 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; </=pleurocentrum;
f=neural arch and spine. Originals in the Carnegie Museum at Pittsburgh.
After Case.

Photograph of amphibian footprints, Dromoput adiimus Branson, from the Mississippian
shales of Giles County, Virginia. X %. Courtesy of Dr. Branson. Original in the
Museum at Oberlin College.

Photograph of type of V'/iim/piis antitjiius Marsh, the amphibian footprint from the
Devonian of Pennsylvania. X %. Courtesy of Dr. Lull. Original No. 784, Vale
University Museum.

THE MICROSAURIAN FAMILY AMPHinAMID.4i.

133

Cephalerpeton ventriarmatum Moodie.
Moodie, Kans. Univ. Sci. Bull., vi, No. 2, pp, 350-352, pi. I, fig. 4; pi. 7, fig. 2, 1912.

Type: Specimen No. 796, of Yale University Museum.

Horizon and locality: Collected at Mazon Creek in 1871, near Morris, Illinois

The remains on which the present species
is based consist of an almost entire skull, 26
consecutive vertebrae, both fore limbs, 20 ribs
preserved on the right side of the body, and a
portion of the ventral armature (plate 4, fig. 4).

The skull is very broad posteriorly, its
width being one-third greater than its length,
with due allowance for crushing. A pineal
foramen is not preserved. The sutures bound-
ing the premaxillaries, the maxillae, the nasals,
the prefrontals, the frontals, a portion of the
parietals, the squamosal, the supratemporal,
the quadratojugal, and the quadrate (?) are
fairly well preserved. The arrangement of
these elements can be discerned by reference
to figure 29. The prefrontals are unusually
large and are triangular in shape. The supra-
temporal is also quite large. The surface of
the skull bones is smooth and there is nowhere
an indication of sculpture.

Portions of 4 sclerotic plates are preserved
in the right orbit. These measure 0.5 by 0.75
mm. The orbits are large and the interorbital
space is less than the transverse diameter of
the orbit. Thirteen teeth, apparently pleuro-
dont, are preserved on the left maxilla. They
are short, sharply pointed, smooth, and unequal. The first 2 left maxillary teeth
from the anterior end are short; then follows a tooth which is one-third longer
than these two; the fourth tooth is somewhat shorter than the third; the fifth
and sixth are still shorter and are practically equal in size, though somewhat larger
than the first two.

The right mandible is preserved almost entire, though so badly eroded that little
can be said of its structure. Impressions of 1 2 teeth are present on the mandible and
all are, apparently, equal. The cotylus seems to have been far posterior and an
angle of the mandible projected slightly back of the skull.

There remain only a few indefinite impressions of the cervical vertebrae. The
union of the skull with the vertebral column is obscured and lost. Impressions of
the dorsal vertebrae are well preserved, and wax molds made from these show
the structure of the dorsal vertebrae surprisingly well. They are long and cylin-
drical, with the median portions slightly constricted by a deep pit on each side of

Fig. 29. — Skeleton of Cephalerpeton ventriarmatum
Moodie. X I.

pf, prefrontal ; d, clavicle ; m, mandible ; h, humerus ;
j, jugal; mx, maxilla; or, orbit; ph, phalanges
of hand; par, parietal; po, postorbital; r,
radius; sp, sclerotic plates; u, ulna; vs, ventral
scutella;.

134 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

the low neural ridge, which takes the form observed in Thyrsidium, Molgophis, Phleg-
ethontia, Dolichosoma (fig. 8) and other genera. The vertebrae are strongly amphi-
ccelous and the notochord was probably persistent. The sides of the vertebras are
smooth.

The ribs are all intercentral in position; the anterior ones very broad near the
base, recalling the broadly expanded ribs described by Schwarz (540) for Scincosau-
rus, Ptyonius, Thyrsidium, and other genera. Posteriorly the ribs become slender
and cylindrical. They are all rather long and distinctly curved, with probably a
cartilaginous tip.

There is preserved a single element of the right side of the pectoral girdle. This
is, I think, the coracoid, an element which has hitherto escaped observation among
the American Microsauria. It is long and spatulate at both ends, with the median
portion apparently almost cylindrical, not unlike that described by Credner (181)
for the coracoid of Branchiosaurus, save that the lower end of the branchiosaurian
coracoid is acuminate. In the present form it is spatulate. Its relations with the
other elements of the pectoral girdle have never been satisfactorily determined.

The fore limbs are both partially preserved. The humerus of the right side is
complete. It is greatly elongated for a microsaurian. The form of the element is
not unlike that of a lizard, with the lower end of the bone spatulate and endochon-
drium well developed. Very little difference can be seen between the form of the
arm bones, which represent the radius and ulna. They are both elongated, with con-
stricted median portion and expanded truncate ends. The carpus is unossified and
the cartilage has left no trace of the elements.

The right hand has two metacarpals preserved, which are fully half as long as
the radius and ulna. They are separated some little distance from the ends of these
elements, though this may be due to post-mortem shifting. The carpus may, how-
ever, have been broad. On the left side are preserved portions of the humerus,
radius, ulna, and 3 metacarpals, lying close to the vertebral column. The carpal
space is not so large on the left as on the right. The ventral armature is well
preserved in a narrow patch about an inch in length. The chevron-shaped rods
are quite large, there being 2 of them in 1 mm.

Measurements.

mm. mm.

Entire length of fossil 98 Median width of a centrum 1.5

Length of skull 22 Length of rib 6.5

Width across base of skull 28 Width of rib at base 33

Long diameter of eye 10.5 Length of coracoid 8

Transverse diameter of eye 8 Width of coracoid at anterior end 2.5

Interorbital space 4 Length of carpal space 5

Length of mandible 26 Length of humerus 18

Depth of mandible at coronoidal region 3.5 Width of shaft I

Depth of dentary 2 Distal width of humerus 4

Length of long tooth 2 Length of radius and ulna 10.5

Diameter of long tooth at base .5 Length of metacarpal 6

Length of preserved portion of vertebral column ... 64 Length of ventral armature preserved 24

Length of a centrum 3 Number of rods in length of 5 mm 10

Type specimen of Ctenerpeton alveolatum Cope, from the Coal Measures of Ohio. X 1.33.
Original in U. S. National Museum.

MOOOIf

Skull of Erpetosaurus minutus Moorlie, from the Cannelton slates of Pennsylvania.

Original in U. S. National Museum. Enlarged X 3.3.
Skull and anterior part of body of Ptyonhis ftcclinatus Cope, from the Coal Measures of

Linton, Ohio. Original in U. S. National Museum. X 1.
Skeleton of' Eosauravus copei Williston, from the Coal Measures of Linton, Ohio. "The

oldest known reptile from North America" and closely related structurally to the

Microsauria. Original in U. S. National Museum. X 1.
l'art of the ventral scutellation and ribs of Sauroplevra iti»i/ata Cnpe, from the dial

Measures of Linton, Ohio. Original in American Museum of Natural History. X 1.

CHAPTER XVIII.

THE M1CROSAURIAN FAMILY NYRAN1ID/E, FROM THE COAL MEASURES OF OHIO.

Family NYRANIID.E Lydekker, 1890.

Lydekker, Cat. Fossil Reptilia and Amphibia, p. 166, 1890.

Skull with the palatines situated near the middle line, internally to the
vomers and pterygoids, and the palatine vacuities small and placed far back.
Vertebras (Ichthyerpeton) discoidal. Teeth less complex than in the Anthraco-
sauridae. A ventral armor present and the entire body covered with small cycloid
imbriated scales.

The type genus of this family was placed by Fritsch (251) with the Archegosau-
ridae, although its resemblance to Anthracosaurus was pointed out; it was subse-
quently made the type of a family by Lydekker (393) in 1890, and placed next the
Archegosauridae. Known from the Coal Measures of Bohemia, Ireland, and Ohio.

Two genera from North America, Ichthyerpeton and Cer car iomor phis, are
assigned tentatively to this family, both known from the Coal Measures (462) of
Linton, Ohio, and both with the body completely scaled. The distinguishing char-
acters are found chiefly in the shape and arrangement of the scales, the structure,
form, and size of the body, all of which are given full treatment in the discussion
below.

Genus ICHTHYERPETON Huxley, 1866.

Huxley, Trans. Roy. Irish Acad., xxiv, p. 195, pi. xxiii, fig. 1; Scientific Memoirs, in, p. 195, pi. 23, fig. 1,
1866.

The genus was founded by Huxley (334) for the reception of the species Ichthy-
erpeton bradleyce from the Kilkenny Coal Measures of Ireland. The remains of the
type specimen represent "the hinder moiety of the trunk, with the greater part of
the tail, of an animal whose scaly integument and laterally compressed, fin-like tail
might easily lead one to take it for a fish, were not its true position among higher ver-
tebrata settled at once by the digitate hind limb ; while its alliance with the labyrin-
thodonts is indicated by the delicate spicular ossicles, which form a rudimentary
dermal shield along the belly." (Huxley.)

Ichthyerpeton squamosum Moodie.

Moodie, Jour. Geol., XVII, No. I, p. 69, 1909.
Moodie, Proc. U. S. Nat. Mus., 37, p. 24, 1909.

Type: Specimens Nos. 4476 and 4459, U. S. National Museum.

Locality and horizon : Linton, Ohio, Coal Measures.

The present species is based on well-preserved remains from the Linton, Ohio,
beds. There are two specimens of the species preserved on blocks of coal and
together they represent the greater part of the length of the animal. The species is
located in the genus Ichthyerpeton, which was founded by Huxley (334, p. 351) on
remains from the Coal Measures of Ireland, on account of the character of the der-

135

!36 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

mal covering, which consists of small scales such as Huxley described in the form
from Ireland. The specific characters of this form are the small size of the rounded
scales, the attenuated tail, the apparent absence of limbs, the character of the ven-
tral scutellation, and the slightly curved condition of the ribs.

It is estimated, from the portions preserved, that the animal attained a length
of not less than 3 feet and its body was long and slender. It may have had an
appearance similar to the modern caudate genus Siren, though there were doubtless
4 limbs present instead of 2. The slenderness of the body is at variance with the con-
dition found in the type species Ichthyerpeton bradleyce Huxley, in which the trunk
was rather stoutly built. The character of the anterior portion of the body in the
present species can not be determined and the skull is wanting. There are no evi-
dences of anterior limbs, although the ventral scutellation preserved would seem to
include the pectoral region. No pectoral shields are preserved, nor are there any
traces of pelvic girdle or limbs.

The preserved portions on one block include nearly the entire tail and the pos-
terior region of the body, and on the other block the dorsal region of the body and
the anterior portion of the tail, so that the two specimens supplement each other
in an interesting manner. There are impressions of several vertebras preserved.
They are much the same in character as Huxley has described for the type species
(I. bradleyce). They are short and thick and were probably amphiccelous. There
are likewise preserved the remains of rather slender recurved ribs mingled in with
the remains of the ventral scutellation and distinguished from the elements of the
abdominal shield by their size and curvature. They are, apparently, single-headed,
but the character of their articulation can not be determined. The ventral scutel-
lation consists of fine continuous rods arranged in the regular chevron pattern.
They do not seem to be divided into oat-shaped scutes, as is the case with the form
described by Huxley. The ventral rods are closely packed for a distance of more
than 6 inches, but as they are scattered their exact arrangement can not be deter-
mined. They seem to have extended to the cloacal region, but there are no evidences
of the specialized clasping organs such as Fritsch (251) has described in the ven-
tral scutellae of Ophiderpeton. The scales, which are well preserved on the tail, may
have covered the entire body, since there are many scattered scales in the dorsal
region of one of the specimens. They are slightly oval, tuberculate, and measure
scarcely 1 mm. in their longest diameter. They show but slight evidences of having
been imbricated, though it is likewise possible that they were simply inclosed within
the integument, and somewhat separated from one another. The most posterior
part of the tail preserved seems to indicate that the tip was attenuated. It was prob-
ably flattened from side to side. We may thus regard Ichthyerpeton squamosum as
an elongate aquatic animal with a long, flattened tail, and since there were possibly
no limbs or very small ones, it would be an animal highly adapted for life in the
water. The present species is of interest because it represents an additional dis-
covery of the scaled Amphibia in North America. The species previously known
from the Linton, Ohio, deposits is Cercariomor phus parvisquamis Cope. Dermal
scales have also been observed in specimens of Amphibamus grandiceps Cope and

1.

4.

5.
6.

Mandible of Macrerpeton deani Moodie, from the Linton, Ohio, Coal Measures.
Original in American Museum of Natural History, No. 2934. X 0.6.

Portion of the skull of Macrerpeton deani Moodie, possibly of the same individual
as the mandible. From the Linton, Ohio, Coal Measures. Original in
American Museum of Natural History, No. 8535 G. X 0.4.

Type of CetvariemorphtU pandsquamis Cope, from the Linton, Ohio, Coal

Measures. Original in American Museum of Natural History. X 1.
An additional specimen of Cercariomoiphus pa/i'isi/naiiiis Cope, from the Linton,

( >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

,<W%;

Cope referred to S. scutellata the larger part of an individual with strongly
developed ventral scutellation. This he figured on plate xxxvi, fig. 2 (123).
Even a cursory glance will, however, suffice to show that the sculpturing of the
clavicle just described and that exhibited by the specimen there figured are identical.
Closer examination shows important differences between the species, the principal
distinctions being the strongly developed ventral scutellae in Sauropleura scutellata,
the difference in form and sculpture of the interclavicles, and the posterior extension
of the interclavicle in 5. scutellata which is wanting in S. pauciradiata. The sculp-
turing of the interclavicle in the form figured by Cope as S. scutellata, just referred
to, is identical with the sculpture of the clavicles, as would be expected. The ridges
on all of the pectoral elements of S. pauciradiata are strong and are rather few in
number, while in S. scutellata the sculpturing is more in the form of interrupted
grooves.

The specimen (plate 14, fig. 3) exhibits the great part of the body with one fore
limb. The skull is wanting. The belly was very broad and strongly protected by
broad, long scutes arranged en chevron. The scutes are close together and form a
compact ventral armor for the animal. The fore limb is very weak. The humerus is
represented by its distal end only. The radius and ulna are very short and weakly
developed in comparison to the size of the animal. The limbs could not have sup-
ported the animal on land and served, probably,
merely as organs of equilibration, for the animal
was undoubtedly aquatic. The fingers are not
all preserved and there is no carpus.

Numerous other remains formerly associated
with S. scutellata are here referred to S. pauci-
radiata, on account of the strongly developed
ventral armor, which is different from that of
the type of 5. scutellata. The remains do not,
however, add to our knowledge of the anatomy
of the forms, as they are very fragmentary.

Two skulls are provisionally associated with
this species. One of these skulls is figured by
Cope (123) on plate xxxm, fig. 1. It is there
referred to S. scutellata. That it can not, how-
ever, be referred to that species is manifest when
the teeth are observed. In the type of S. scutellata the teeth are very small, sharp
denticles, while in the skull under discussion the teeth are well developed and their
bases are longitudinally grooved. The teeth are elongate in the anterior part of the
skull and are shorter posteriorly. They are, however, all strong. The skull is acu-
minate and the orbit is located about midway of its length. The jaw is slightly longer
than the cranium. The structure of the cranium can not be determined in either
skull, and in one only the position of the orbits and the teeth.

Fig. 34-

A. Interclavicle of Sauropleura pauciradiata

Cope. X 1. (After Cope.)

B. Left clavicle of Sauropleura pauciradiata

Cope. X 1. (After Cope.)

l6o THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

Measurements of the Specimens of Sauropleura pauciradiata Cope.

ram. ram.

Length of clavicle, left 43 Space between ridges 2

Width of clavicle 25

Large specimen No. 8637 G and obverse (13) described as Colosteus sculcllatus Newb.

mm. ram.

Length of specimen, as preserved 117 Width of clavicle 18

Width of specimen, maximum 61 Length of interclavicle 40

Length of fore arm 9 Width of interclavicle 23

Length of hand 10 Width of a ventral scute 1.5

Length of metacarpus 5 Length of scute from angle to end 35

Length of clavicle 40

Skull No. 8666 C, plate xxiii,fig. I (Cope, 123), Colosteus scutellatus.

mm. mm.

Length of skull, as preserved 65 Width of mandible, maximum 10

Width of skull 4° Length of longest tooth 5

Diameter of orbit 6 Width of tooth at base 1.5

Length of mandible 70

Skull Nus. 8602 G and 860S G.

mm. mm.

Length of skull, as preserved 50 Diameter of orbit 15

Width of skull 60 Length of mandibular tooth 4.5

Other specimens associated with this species: Nos. 8668 G, 86740,85540,
8661 G. All specimens in the American Museum and all from Linton, Ohio.

SCUTES OF SAUROPLEURA.

There are found associated with the remains of the genus Sa u ro pleura a num-
ber of heavy scutes or scales. There are three of them on the same block as the
specimen of S. longidentata and are provisionally referred to the genus. There
are a number of scutes preserved separately, but they agree in their characters
with those discovered on the specimens. The scutes are elongate and usually acu-
minate at one end and having a broad base at the other. The acuminate end is
slightly bent to one side so as to present the appearance of a hook. Others are
shield-shaped and are quite large, while the majority of the hook-shaped ones are
small. The shield-shaped elements have a rounded boss near the center of the
plate and the edges are imbricated. Their nature and their proper location on the
animal are a puzzle. They may not belong to the genus, but have been noticed with
the remains of at least 3 species.

Measurements (in Millimeters) of Scutes Associated with the Specimens of Sauropleura.

No. 4513 U. S. National Museum:

Length 27

Width, maximum 14

Width, minimum 3

Scutes associated with the specimen of Sauropleura longidentata:

Width across base 3

Width across tip 1

Additional specimens are: Nos. 3, 8673 G and 8470 G, American Museum of
Natural History.

Sauropleura longidentata Moodie.
Moodie, Jour. Geol., 17, No. 1, pp. 74-76, figs. 18, 19, 1909.

Type: Specimen No. 8619 G, American Museum of Natural History.
Horizon and locality: Linton, Ohio, Coal Measures.

This species may be distinguished from other members of the genus by the large
size and shape of the cranium (462) and the broad mandible (plate 16, figs. 2, 3)

MOODII

FUkTS 22

1.

I.

Type of Leplophractns lineolatus Cope, from the Coal Measures of Linton, Ohio. Por-
tions of maxilla and mandible of left side with teeth. XI. Original in American
Museum of Natural History.

Type of Proterpelon gurleyi Moodie, from the Coal Measures of Illinois, near Danville.
Cervical of an otherwise unknown vertebrate. Neural spine to the right.
Original in Walker Museum, University of Chicago X 2.

Amphibian phalanx from the Coal Measures near Breeze, Illinois, of an unknown
species. The probable form of the element is represented. X 3.

Large rib of a stereospondylous stegocephalan otherwise unknown. The rib may
represent a species of Macrerpeton or may even belong with Macrerpeton deani
Moodie, but exact identification will have to wait for future discoveries. From
the Coal Measures of Linton, Ohio. Original in U. S. National Museum. X 1.

Type of Cope's species Tia/i/aniis mordax referred by him to the cranium, on account
of the sculpturing of the elements. We now know the specimen to be portions of
the interclavicle and clavicles of Diceralosaiirits ptinctolineattis, as Cope suggested
they might be. Original in American Museum of Natural History. X 1.

Skull of Raphetes planiceps Owen from the Coal Measures of Nova Scotia. 0.45.
Original in the British Museum, South Kensington. After Owen.

THE MICROSAURIAN FAMILY SAUROPLEURIM:. l6l

with its very long teeth. The skull of Sauropleura digitata Cope is not known, but
the body of that animal as preserved represents far too small a form for the skull to
be referred to that species. The skull is fully half as long as the dorsal region of S.
digitata Cope, so that an association of the remains would be incongruous. It differs
from the skull of S. scutellata Newberry in size and proportions. The skull of S.
scutellata is narrow, while in S. longidentata it is quite broad. The teeth of the latter
are also characteristic of the species, since in all other known species of this genus in
which the skull is preserved the large anterior tooth is wanting.

The bones of the skull show the coarse sculpturing of the larger species of Micro-
sauria. It consists more of radiating grooves than of pits. The skull, as restored
(462), is broadly ovate, with the posterior border truncate. The muzzle is broad
and the nostrils are, apparently, located near the anterior margin.

The posterior border of the orbits lies near the median transverse line of the
skull. They are circular and are removed some distance from the margin of the
cranium. Only the frontal and parietal can be determined with certainty.

The mandible is heavy and is provided with pleurodont, heterodont teeth. Near
the anterior end of the mandible there is a very long fang-like tooth, longitudinally
striated and slightly recurved, which arises from a broad base and attains to con-
siderable prominence. The other teeth are smaller, though the next succeeding one
is still of considerable size. All of the teeth preserved are longitudinally striated,
but only the two anterior ones are recurved to any extent.

Measurements of the Type.

mm. mm.

Length of the skull in median line 75 Width of jaw, maximum 16

Width of skull at posterior border, estimated 80 Width of jaw, minimum 5

Width of skull across orbits 60 Length of largest tooth 11

Width of orbit 10.6 Width of longest tooth at base 4.5

Length of orbit 12 Length of shortest tooth 3

Interorbital space 16 Width of shortest tooth at base 1

Length of jaw, as preserved 48

Sauropleura foveata Cope.

Cope, Trans. Am. Phil. Soc., xiv, p. 24, 1869.

Cope, Geol. Surv. Ohio, 11, pt. 11, p. 406, pi. xxxvi, fig. 1, 1875.

Type : Specimen No. 8676 G and obverse No. 8675 G, American Museum of
Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

This species is represented by an interclavicle and its impression, which show a
beautiful sculpturing entirely distinct from that of the other species of this genus.
In size it is intermediate between the largest of the interclavicles of S. scutellata and
S. pauciradiata. The pattern of the sculpturing is, however, its main distinction.
The plate is finely pitted and there are few evidences of grooves. Near the posterior
border of the plate the pits become somewhat defined by ridges which take on a
radiating pattern with the center of the plate as the center. The beveled margins
are rugose, except at the edges.

Measurements of the Type Specimen of Sauropleura foveata Cope.

mm.

Median length of interclavicle 43

Width of interclavicle, maximum 23

1 62 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

Sauropleura enchodus Cope.
Cope, Proc. Am. Phil. Soc, p. 406, 1885 (Anisodexis).

Type: Specimen No. (51) 2558, American Museum of Natural History, Newberry
Collection.

Horizon and locality: Discovered by Sam Huston at the Linton, Ohio, Coal
Measures.

An examination of the type specimen (plate 16, fig. 4) of this form resulted in no
new facts. The reference of the species to the Permian genus Anisodexis by Cope
is probably incorrect. Our knowledge of the two known species of this genus is not
sufficient to separate them, but for the sake of convenience the Linton species is
placed in the genus Sauropleura. It is located here principally on account of the
form and structure of the teeth. Cope's original description is given below:

"The generic characters are apparent in the very unequally sized teeth with round
section. The portion upon which the species is based is a part of the right ramus of the
mandible, which is in the specimen viewed from the inner side. The jaw is obliquely and
smoothly truncated from below, for the symphysis and surface of the bone is smooth.
There is a very large tooth near the extremity of the dentary bone. Behind it is an interval
equal to three times the diameter of its base, which is followed by a tooth of about one-
third the length of the first tooth. Posterior to this one are two teeth of the same size as
the second, all being separated from each other by about a tooth's diameter. These are
followed by three subequal teeth of about two-thirds the length of the first tooth, and
separated by about their own diameter from each other. They are all perfectly straight,
very acute, and without the trace of a cutting-edge. The inflection-grooves extend to or
a little beyond the middle of the length."

The present species is smaller than the type of Anisodexis imbrecarius from the
Texas Permian, to which genus Cope originally referred the present species, and the
apices of the teeth do not display the opposite cutting-edges seen in the Texas form.

Measurements of the Type Specimen.

mm. mm.

Length of jaw, including 7 teeth 31 Length of third tooth 3.5

Depth of ramus at second tooth 10 Length of sixth tooth 7.5

Length of first tooth 10.5

SKIN OF SAUROPLEURA SP.
Moodie, Bull. Am. Mus. Nat. Hist., xxvi, p. 355, pi. lx, fig. 1, 1909.

The present specimen is interesting on account of the presence of what I take to
be a portion of the skin, which is preserved as a smooth mold over the ribs and ven-
tral scutellas. The skin was undoubtedly that of the back, since the creature is
preserved on its belly, and is interesting in not showing the slightest trace of scales
or other hard plates. The ventral scutellae are characteristic of the species of the
genus Sauropleura. With one species of this genus, Sauropleura scutellata Newberry,
the writer has found associated scutes of some size, and the same fact has been
noted by Cope.

Genus SAURERPETON Moodie, 1909.
Moodie, Jour. Geol., xvn, No. 1, p. 80, fig. 23, 1909.

Type: Saurerpeton latithorax Cope.

This generic name is erected for the reception of a single species described by
Cope in 1897. The name is made necessary by the wide divergence of the charac-
ters exhibited by the present species from those of the species of the genus {Sauro-

THE MICROSAURIAN FAMILY SAUROPLEURID^E. 163

pleura) to which Cope (176) referred this species. The form is not a member of the
genus Sauropleura, for reasons given below.

The species of the genus Sauropleura have a lanceolate head with homodont den-
tition or nearly so. The orbits are located well back in the skull. The form of the
body is elongate and slender and the limbs where known are long and attenuated.
The ventral scutellation consists of oat-shaped scutes arranged in a chevron series.
The form here described as Saurerpeton latithorax Cope has nearly the opposite of
all of these characters, and it is incongruous to locate the form under the former
genus. The skull of Saurerpeton latithorax Cope is broad and heavy. The teeth
are heterodont. The body is broad and stout and the limbs are of unusually
strong proportions. The character of the ventral armature is also of a very differ-
ent type. In Saurerpeton it consists of very broad imbricating scutes which form a
single piece across the abdomen and are angulated to form the chevron pattern
which is so common among the Stegocephalia.

Saurerpeton latithorax Cope.

Cope, Proc. Am. Phil. Soc, xxxvi, p. 86, pi. iii, fig. 4, 1897.
Moodie, Jour. Geol., xvn, No. 1, p. 80, fig. 23, 1909.

Type: Specimen No. 4471, U. S. National Museum.

Horizon and locality: Linton, Ohio, Coal Measures. Collected by R. D. Lacoe.

This species is indicated by remains of the anterior half of a large amphibian
(plate 17) preserved on a block of bituminous coal from the Linton mine. The form
is unusual in the proportions of the head and the width of the thoracic region. In
these characters it stands alone among the Amphibia from this locality, where the
forms are for the most part of rather slender build and tapering, pointed head.

The skull is represented in a fairly complete condition and shows the usual
stegocephalian arrangement of the skull elements, as well as the sculpturing of the
bones, which is similar to that found in other members of the Microsauria. The
skull is broadly rounded, with the posterior border incised, the broad tympanic
notches thus rendering the shape of the skull somewhat like that of the branchio-
saurs. The orbits are broad ovals and lie well forward in the skull. They are sepa-
rated by a space which is greater than the greatest diameter of the orbit. The
pineal foramen lies well back and is clearly indicated as a circular opening which
lies in the median suture in the posterior half of the parietals. The nostrils seem to
be elongate and have an oblique position, as is represented in the diagram (fig. 35),
but this character is not ascertained definitely.

The borders of the premaxillae and the anterior suture of the nasal can not be
determined, though they may have had some such arrangement as suggested. The
nasal is represented, so far as is determinable, by an oblong element lying between the
frontal and the anterior border of the skull. The frontals are very large and form a
portion of the inner border of the orbit. The parietal is probably the largest bone
in the cranium and together the two elements form a large quadrangular space
in the posterior half of the skull. They inclose the circular pineal foramen. The
postparietal is a small element lying on the posterior border of the skull and with
the tabulare and a part of the supratemporal forms the projection. The prefrontal

1 64

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

is probably a small element, especially if the lacrimal is present. There seems to
be an indication of a suture separating the lacrimal from the prefrontal, but this
character is not assured. The boundaries of the maxillae are clear anteriorly. They
indicate that this element was elongate, as is usual, and impressions of teeth borne
on the mandible would indicate the probability that the teeth were heterodont.
In the premaxillary region there is a long, strong tooth preserved, and on the max-
illary near the posterior ex-
tremity of this bone there are
impressions of teeth which are
no more than one-fourth as
large as the premaxillary one.
The borders of the jugal can
not be ascertained, since the
skull is injured on both sides
in this region. Likewise the
quadratojugal is conjectural.
The boundaries of the post-
frontal and the postorbital are
clearly ascertained. They to-
gether form the entire poste-
rior border of the orbit and
send prolongations along the
lateral borders of these open-
ings. They are both acumi-
nate posteriorly and these
points are inclosed by the tab-
ulare and supratemporal for
the postfrontal and by the
squamosal and prosquamosal „

f ,, , . . _,. riG. 35. — Outline drawing of the skull and skeleton, as preserved, of

Space IOr tne pOStOrbltal. I he Saurerpelon latithnrax (Cope), from the Linton, Ohio, Coal Measures,

hrainrlormo ^f +U^ 4-„u 1 Original in U. S. National Museum. X 0.7s.

boundaries Ot the tabulare Skull: /,, frontal; j jugal; mx, maxilla; n, nasal; no, nostril; or, orbit;

Show this element to be Quite pnr'c Paneta': P°f, postfrontal; po, postorbital; pp, postparietal; pf,

" prefrontal; pmx, premaxilla; qj, quadratojugal; m, mandible; spo,

large and extending forward supraorbital lateral-line canal; spt, supratemporal; sq, squamosal;

• , ... .... tub, tabulare; v, pineal foramen.

mtO an aCUmmatlOn Which IS Skeleton: d, clavicle; c, carpus; rb, ribs; ft, humerus; it, interclavicle;

inclosed bv the Oarietal nnrl »ie^metacarpals;/w, pectoral scutelte;r, radius; w, ventral scutclUe;

the postfrontal. The sutures bounding the squamosal have been obscured by injury
in removing the specimen and are indeterminable.

The vertebral column is represented by a ridge showing along the median plane
of the specimen. Its characters can not be ascertained. There are a few evidences
of ribs. They represent long, slender, non-alate elements, their mode of articu-
lation with the vertebras being obscured by the plate-like ventral armature which
covered the abdomen of the animal.

The pectoral girdle is represented by the remains of three elements which are
interpreted as being the interclavicle and the two clavicles. The interclavicle is
broad and is rounded posteriorly. There is no evidence of the usual acumination

THE MICROSAURIAN FAMILY SAUROPLEURID/E. 1 65

posteriorly. The element is nearly as wide as long. There is a prominent longi-
tudinal keel on the ventral surface of the interclavicle and radiating lines which
may indicate the courses of blood-vessels or nerves or may be the ornamentations
of the element, probably the latter. The clavicle has the usual microsaurian form.
It has three points and is truncate exteriorly. It is ornamented with radiating
grooves of a shallow and not strongly pronounced character. There is no evidence
of the coarse sculpture of the later forms. If the scapula is represented it is merely
by an indeterminate fragment insufficient for description.

The pectoral limbs are preserved nearly entire. The left fore limb lacks only a
few phalangeal bones, and these were preserved with the remainder of the skeleton
but were lost in the mining process. The humerus is an extraordinary element on
account of its robust dimensions. It is very stoutly built and represents a power-
ful limb. It is expanded at each extremity and the width of its shaft is about equal
to one-fourth of its length. The ulna and radius present the same characters as
the humerus, i.e., in being robust, with stout shaft and expanded ends. The ulna is
slightly longer than the radius and has an expanded upper end. The radius is short
and does not have the proximal expansion. The carpus was cartilaginous. Its posi-
tion is represented by a blank space on the coal. There are 4 digits preserved and
in all probability this was the entire number. The metacarpals are elongate and
expanded at the extremities. The first and second digits are represented nearly com-
plete. The first digit is extremely interesting in the possession of a claw-like terminal
phalanx which much resembles that of some lizards. There are 3 phalanges in the
first digit and 4 in the second. The phalangeal formula may have been 3-4-4?.

The ventral scutellation of this species is of an unusual character. It consists
of broad, imbricated scutes which are in a single piece and which are arranged in
the usual chevron pattern. The scutes were, apparently, broadest in the middle
and tapered somewhat at the extremities. This character alone is sufficient for
separating the genus from that of any other known form.

The genus finds its nearest allies in the forms of the species of the genus Sauro-
plcitra, in which Cope formerly located the present species. The skull of the form
described as Tuditanus radial us Cope is quite similar to the present form, both in
the sculpturing and arrangement of the elements. The characters wherein the pres-
ent form resembles the species of Sauropleura are the possession of broad pectoral
plates and strong, digitate limbs. The general form of the body and skull is different
in the two groups. It is slender in Sauropleura and decidedly stout, short, and
heavy in Saurerpeton.

M KASI KEMENTS OF THE TYPE.

mm. mm.

Length of specimen 130 Width of clavicle 18

Median length of skull 51 Length of abdominal scutes 28

Width of skull at posterior border 62 Length of humerus 19

Width of skull across orbits 48 Width of humerus at upper end 6.5

Length of orbit 14 Length of ulna II

Width of orbit 10 Length of radius 9

Interorbital space 16 Length of metacarpal 3

Length of longest tooth preserved 4 Length of first digit 12

Length of shortest tooth preserved I Length of terminal phalanx 3.5

Length of the interclavicle 26 Length of lower jaw on the curve 70

Width of interclavicle, maximum 23 Width of lower jaw, maximum 8

Length of clavicle 22 Length of rib 30

1 66 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

Genus CTENERPETON Cope, 18Q7.
Cope, Proc. Am. Phil. Soc, xxxvi, p. 83, pi. iii, fig. 1, 1897.

Type : Ctenerpeton alveolatum Cope.

The genus Ctenerpeton was founded by Cope for the reception of a single species
which presents some very unusual characters. The form shows close relationships
to the genera Urocordylus (334), CEstocephalus, and Ptyonius. These genera agree
with the present one in the possession of very characteristic vertebrae which are
signalized by the elongate and ornamented characters of the neural and haemal
spines. These project prominently from the body of the vertebra and have the
ends of the projections truncate and divided into fine points, thus causing the spine
to have much the appearance of a comb. The surface of the neural spine is some-
times marked with a shallow groove. The spines are longer and more slender in
the genera CEstocephalus, Urocordylus, and Ctenerpeton than they are in the species
of the genus Ptyonius, where they are short, although the usual pectinations are
present.

The character on which this genus rests is the shelf -like extension (plate 23, fig. 2)
of the abdominal plates. This is of a very unusual character and entirely unknown
in any other species of Carboniferous Amphibia. The term Ctenerpeton has reference
to the fact that the ends of these shelf-like plates project in a pectination along the
side of the abdomen.

Ctenerpeton alveolatum Cope.

Cope, Proc. Am. Phil. Soc, xxxvi, p. 83, pi. iii, fig. 1, 1897.
Moodie, Proc. U. S. Nat. Mus., 37, p. 24, pi. 10, 1909.

Type: Specimen No. 4475, U. S. National Museum, Lacoe Collection.

Horizon and locality: Linton, Ohio, Coal Measures.

The species rests on a single specimen (plate 19) from Linton, Ohio, and is pre-
served on a block of bituminous coal. It is in a very good state of preservation.
There are present on the block of coal a part of the right fore limb, the greater part
of the dorsal portion of the animal, and the anterior part of the tail. There are no
evidences of hind limbs, although this may not be taken as evidence that they were
not present on the animal. No thoracic plates have been observed. Thexhevron
armature is present beyond the cloacal region, but there are no evidences of the
specialized clasping organs which are apparently developed from the abdominal
armature in some forms (251).

Each dermosseous rod of the abdominal scutellation consists of three pieces —
a median angulated portion and the two lateral parts which form the shelf-like
projection along the side of the abdomen (plate 23, fig. 2). The marginal chevron
differs in form from the other plate, aside from the fact that it is not angulated.
The lateral shelf is composed of flattened plates which articulate with the median
piece, and at the place of articulation there is a ridge present in the specimen. The ex-
terior plates are curved backwards and are somewhat attenuated distally. They are
broader than the median piece and differ also in the absence of the characteristic
alveoli. The median plate is angulated and is of more slender proportions than the lat-
eral pieces. Its ventral surface is ornamented with a single row of closely placed

MOOOli

PiATE 23

2.

Left leg and pelvis of Ichlhynviihus platypus Cope, from the Coal Measures of Ohio.

X 1.7. Original the property of the Department of Geology of Columbia University.

r=centrale; /c"=fibula;yf=fibulare: /v— -femur; t=intermedium; //=ilium; 7~=tibia;

/, /-5=distal tarsalia; /;'=tibiale: F=caudal vertebra; /-K=digits.
Ventral scutellas of Ctenerpeton alveolatum Cope, from the Coal Measures of Ohio. X 3.5.

Original in U. S. National Museum.

THE MICROSAURIAN FAMILY SAUROPLEURID.E. 1 67

alveoli which resemble in a great degree the alveoli of the jaw of some small animal.
The ventral scutellation is broad anteriorly, but becomes more slender posteriorly
and shortly posterior to the cloacal region disappears.

The fore limb is represented by the upper portions of the ulna and radius and
2 digits of the right hand. The digits are long and slender and seem to represent
digits I and II, since they show evidences of 3 and 4 phalanges respectively. The
portions of the fore arm preserved are too meager for description.

The vertebrae have already been characterized as of the type first described in
Urocordylus. The neural fans are not much, if any, wider than the haemal fans.
They are both situated on an elongate spine with a slender base. The edges of the
two fans are pectinated and the dorsal spine is distinguished by the presence of a
longitudinal groove in the center of the spine. The length of the tail may have
been considerable, judging from the character of the vertebrae preserved

Measurements.

mm. mm.

Length of specimen 150 Width of the same vertebra with spines 20

Width of belly, maximum 28 Height of neural spine 8

Length of lateral chevron plate 7 Height of haemal spine 8

Width of lateral chevron plate 1.75 Width at distal end of hsemal spine 3

Length from tip of lateral chevron to median Width at distal end of neural spine 3.5

angle 10 Length of foot, as preserved 12

Length of a caudal vertebra 10

Genus LEPTOPHRACTUS Cope, 1873.

Cope, Proc. Phila. Acad. Nat. Sci., p. 340, 1873.
Cope, Geol. Surv. Ohio, 11, pt. 11, p. 399, 1875.
Cope, Proc. Am. Phil. Soc, xx, p. 461, 1882.

Type: Leptophractus obsoletus Cope.

The genus was established on various parts of the cranium of a large amphibian.
The only parts which can with certainty be referred to the genus are the upper and
lower jaws of 3 specimens. These bear large teeth, round in section at the base,
but with acute compressed apex, with a cutting -edge on the anterior face; the
enamel is delicately grooved, as an external indication of the labyrinthic structure.
A characteristic feature is seen in the presence of a large elongate tooth in the upper
jaw, in the position of a canine which much exceeds in length any of the others.
The sculpture of the cranium is but little marked in the known specimens. In the
type the lower jaw is marked with inosculating grooves. Three species are known,
which are among the largest of the Linton Amphibia.

Leptophractus obsoletus Cope.

Cope, Proc. Phila. Acad. Nat. Sci., p. 340, 1873.
Cope, Geol. Surv. Ohio, 11, pt. 11, p. 399, 1875.
Cope, Proc. Am. Phil. Soc, xx, p. 461, 1882.

Type: Specimen Nos. 55 G and 57 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures.

The species was about as large as an adult Florida alligator, and probably
exceeded or at least equaled in size any of the Carboniferous Amphibia. The fol-
lowing account is taken directly from Professor Cope's " Batrachia of the Ohio Coal
Measures" (123). The description has been verified from an examination of the
type material.

j68 the coal measures AMPHIBIA OF NORTH AMERICA.

" The teeth are rather distantly grooved for some distance above the base. They are
of different sizes; the smaller are compressed, and with fore and aft cutting edges. The
external surface of the dentary bone is marked with short oblique grooves along its middle
region; above these are grooves which inosculate, forming a figure like an open net dragged
in the long direction.

"Excepting the grooves the teeth are smooth. The smaller ones are close together
and their crowns are curved backwards; the larger ones are at more remote intervals; both
have enlarged bases. Whether both forms are in the same series I can not determine.
There are from four to five of the smaller to an inch.

Measurements of Type of Leptophractus obsoletus Cope.

mm. mm.

Deplh of fragment of jaw 75 Width of paired median scuta 56

Length of smaller teeth 19 Width of single scute 36

Length of longer tooth 23 Length of single scute 48

Width of vertex at middle scuta 176

' ' Some vertebrae were found at the same locality, but there is no evidence as to the
species to which they may have pertained. They are short, concave on one end, and
probably so on the other. The centrum of one is 12 mm. in diameter ; neural spines injured."
(Geol. Surv. Ohio, 11, pt. 11, pi. 39, fig. 3.)

"A third and larger specimen was found by Professor Newberry during the field season
of 1874. It includes an oblique view of one side, and the top of the cranium from the
posterior part of the orbits to the end of the muzzle, with the corresponding part of the
alveolar region of the dentary bone, with teeth. The bones of the skull appear to have
been rather light, and though the surface is irregular, the sculpture consists only of shallow
impressions of varying size and intervals. The orbits are also badly defined, but appear
to have been large, and separated by a narrow frontal bone. The premaxillary bone is
preserved, and shows clearly the sutures that separate it from its fellow and from the
maxillary. A large foramen — perhaps the nostril — separates it from the maxillary, so that
it forms an irregular crescent. It supports two teeth, of which the anterior is the larger,
but there were perhaps others in advance, as the alveolar border is imperfect towards the
end of the muzzle. The anterior two teeth of the maxillary bone are followed by a strong
groove which rises towards the sides of the muzzle. At first sight this gives the impression
of the maxillo-premaxillary suture, and makes it appear that both the premaxillary bones
are preserved, and that the foramen above described separates the premaxillary spines,
instead of representing the nostril. The cutting edges of the teeth of these bones have,
however, one direction, whence they represent one side of the cranium only; were both
sides represented, the directions of the tooth axes would be reversed.

' ' The premaxillary and maxillary teeth exhibit a cutting edge on the outer posterior
margin of the distal half; the base of the crown is subround in section. The line-like
grooves are distinct but not numerous, their intervals measuring 75 mm. Beyond them
the enamel is smooth. The second maxillary tooth is larger than the first, which is equal
to the last premaxillary. The third and fourth maxillaries are equal to the second, but
the fifth is larger and longer, exceeding all the others. The teeth of the dentary bone
differ from those of the upper jaw in having the cutting edge of the crown on the anterior
aspect, while the posterior border is obtuse. There is an obtuse cutting edge on the pos-
terior margin of the anterior mandibular teeth.

"This description is derived from an adult animal, as the maxillary teeth in some
instances are partially worn away by friction on their anterior and outer faces."

Measurements.

mm. mm.

Length of maxillary bone preserved 146 Diameter of same at base 6

Length of same supporting live teeth 73 Diameter of sccondat base 8

Length of first maxillary tooth 15 Length of basis of fifteen teeth of the dentary 145

THE MICROSAURIAN FAMILY SAUROPLEURID.4i.

169

i mjt< j a A ■ *,•**._

Fig. 36. — Mandible of Leptophraclus denlatus new species, from the
Linton, Ohio, Coal Measures. X I. Original in American Museum
of Natural History.

, Leptophractus dentatus new species.

Type: Specimen No. 1085 G, American Museum of Natural History. Col-
lected by Dr. J. S. Newberry.

Horizon and locality: Linton, Ohio, Coal Measures.

The type is a single right mandible, nearly entire, of a rather large animal. The
specific characters for the separation of the new form from the previously described
L. obsoletus and L. lineolatus are
the smaller size and slenderness
of the mandible, associated with
uniform teeth, which are slender
and delicately fluted.

There are 1 7 teeth preserved,
the largest of which is 8 mm.
in length. From the posterior
tooth the series gradually descends to half this length 0.5 inch from the anterior
end of the mandible.

The exact form of the mandible can not be determined, but so far as can be
seen it is very slender, coming almost to a point at the anterior end. The poste-
rior portion is wide, but apparently not very heavy. There is a fragment associated
with the specimen which discloses a few teeth, but its position in the cranium can
not be determined.

Measurements of the Type of Leptophractus dentatus Moodie.

mm. mm.

Length of mandible, as preserved 80 Length of most anterior tooth 4 .

Anterior width of mandible 3 Length of tenth posterior tooth 7

Posterior width 28 Width of this tooth at base 3

Leptophractus lineolatus Cope.
Cope, Proc. Am. Phil. Soc, xvi, p. 576, 1877.

Type: Specimen No. 1088 G, American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures. (Plate 22, fig. I.)

Cope's description of this species, to which I have nothing to add, is as follows :

" This species is based on portions of the skull of two individuals of large size. Both
upper and lower jaw with teeth are represented and the teeth are of very large size. The
deepest element preserved has been provisionally referred to the mandible and will be so
described. This element bears two types of teeth in very heterodont fashion. The teeth in
the back portion of the jaw are rather short and slender. The teeth more anteriorly are,
some of them, very long, rather stout, with their bases longitudinally fluted, as are they all.
The longest tooth in the jaw measures slightly less than an inch. The bone is not well
preserved, but seems to have been ornamented with grooves of no great depth. The most
anterior teeth of the jaw are smaller than the posterior ones.

' The upper jaw is set with teeth which are more uniform in size and there is but little
tendency to heterodonty. These teeth are also striated at their base and all end in a sharp
point. They are all, apparently, straight. There is no tendency to curve as there is in the
genus Macrerpeton. The upper teeth are more closely set than are those of the lower jaw,
which are rather distantly placed.

"Another specimen of a smaller individual presents the same portions of the skeleton
and the same characters. It is possible that this skull will be found to belong to Ichthycan-
llius ohiensis Cope, which is based on very large vertebras and limb bones. The remains

170

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

described as Leptophractus lineolatus are, however, unlike any other skull remains which are
thus far known.

' ' This species represents one of the largest types of the Carboniferous Amphibia of
Ohio. It probably attained a length of several feet. It was also the most carnivorous of
any of the forms."

Measurements of the Type Specimen of Leptophractus lineolatus Cope.

mm. mm.

Length of longest tooth in jaw 22

Anteroposterior diameter of the same at base 6

Length of small maxillary tooth 7

Anteroposterior diameter of same at base 2

Length of specimen, as preserved 98

Depth of dentary bone at the middle 30

Anteroposterior diameter of mandibular tooth at

base 3-5

Length of a posterior mandibular tooth 9

Collected by Dr. J. S. Newberry. Other specimens of the species are Nos.
1086 G and 1087 G, American Museum.

Eurythorax sublsevis Cope.*

Cope, Proc. Am. Phil. Soc, p. 177, 1871.

Cope, Geol. Surv. Ohio, 11, pt. 11, p. 401, pi. xl, fig. 4, 1875.

Type: Specimen No. 8605 G, American Museum of Natural History. Collec-
tion of Dr. J. S. Newberry.

Horizon and locality: Linton, Ohio, Coal Measures.

The genus and species were established on a single, large, almost perfect inter-
clavicle of peculiar form. It may or may not be distinct, but it can not be asso-
ciated definitely with any known genus, so that
it will be designated as Cope described it. It
exhibits on its outer or lateral borders broad,
smooth surfaces for the contact of the overlapping
margins of the lateral plates. The form is sub-
round, with a large excavation from the posterior
margin on each side. The narrowed portion left
in the middle behind has a convex outline. Some
delicate radiating grooves occur on the exposed
surface, but they are very shallow.

Measurements of Type.

mm.

Length of interclavicle 71.5

Greatest width 78 Fig. 37. — So-called interclavicle of Eurythorax

Width of lateral concavity 39 sublcevis Cope. (Sagenodus.) X 0.75. (After

Cope.)

* Dr. Hussakof now regards (Bull. Amar. Mus. Nat. Hist., vol. 35, p. 128, 1916) the type specimen of this species
as an opercular element of one of the dipnoan fishes [Sagenodus sublavis (Cope)]. The discussion is left here, however,
as a matter of historical interest.

CHAPTER XXII.

THE M1CROSAURIAN FAMILY ICHTHYCANTHIDyE, FROM THE COAL MEASURES

OF OHIO.

Family ICHTHYCANTHIDjE new family.

This family is the most recently recognized group of the Linton fauna. Its
members, of which there are two known species, are distinct from all other Coal
Measures Amphibia in the possession of an osseous tarsus (483, 484), with its asso-
ciated reptile-like limb bones. There are preserved fine scutellae in a large patch
near the vertebral column. The vertebral spines are broad and heavy, with the
vertebral centra amphiccelous.

Genus ICHTHYCANTHUS Cope, 1877.

Cope, Proc. Am. Phil. Soc, p. 573, Feb. 3, 1877 (Pal. Bull. 24).

Baur, Beitrage zur Morphogenie des Carpus und Tarsus der Vertebraten, I Theil, p. 16, 1888.

Cope, Trans. Am. Phil. Soc, xvi, p. 289, fig. I, 1888.

Moodie, Science, n.s., xli, No. 1044, p. 34, 1915.

Moodie, Am. Jour. Sci., xxxix, pp. 509-512, fig. 2, May, 1915.

Type: Ichthycanthus ohiensis Cope.

The generic characters are derived from the characters presented by the pos-
terior dorsal and caudal vertebrae, with adjacent parts. The posterior limbs are
well developed, with distinct tibia and fibula, osseous tarsus, and 5 digits. Ribs
elongate, simple, curved. Abdominal armature consisting of bristle-like rods in
anteriorly directed chevrons. Dorsal vertebrae not elongate, with simple neural
spines. Tail large, its vertebrae ossified, and furnished with slender chevron bones
which terminate in a haemal spine. Neural spines broad and directed backwards ; the
caudal series somewhat resembling that of a fish. All the centra are amphiccelous.

This genus differs from all those with enlarged and sculptured neural spines,
and from those with abdominal scutes. It is equally distinct from those without
ribs, abdominal rods, or limbs. It is possible that some of the species referred to
Tuditanus, in which these parts are unknown, may belong to it, or that it may be
established on a small species of Leptophr actus, a genus known only as yet from the
skull. With our present imperfect knowledge of the Linton forms it seems best to
refer /. ohiensis and I. platypus to this distinct genus, Ichthycanthus.

Ichthycanthus ohiensis Cope.

Cope, Proc. Am. Phil. Soc, 1877, p. 573 (Pal. Bull. 24).

Type : Specimen in the American Museum of Natural History.

Horizon and locality: Linton, Ohio, Coal Measures. Collected by Dr. J. S.
Newberry, in the summer of 1876.

The centra of the dorsal vertebrae are about as long as deep, and their sides are
deeply concave; there are 4 anterior to the pelvis which are without ribs. The
caudal vertebrae are robust, and 7, from the first, support a small tubercle-like
diapophysis. The chevron bones are short and acuminate; the neural spines are a
little shorter, narrow, and truncate, and directed backwards at the same angle as

171

172 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

the chevron bones. They are much reduced on the eighteenth caudal vertebra,
where the chevron bones are considerably longer.

The abdominal rods are quite slender. The hind limb is quite stout for this
order. The femur is regularly expanded at both extremities, but the distal is deeply
and openly grooved, distinguishing the condyles, while the proximal end is plane.
There is no trochanter visible. The ulna and radius are well separated, and are
three-fifths the length of the femur. There is a large fibular tarsal bone of a sub-
quadrate outline. In immediate contact with it is probably the external digit with
5 phalanges or segments ; the ungual is simply conic. The femur is as long as 5 dorsal
vertebrae. The ribs have expanded, undivided heads, and extend to the abdominal
armature.

Measurements of the Type of Ichthycanthus ohiensis Cope.

mm. mm.

Length of last 10 dorsal vertebrae 47 Proximal diameter of femur 8

Length of first 23 caudal vertebrae 117 Width of lower leg 9

Length of a posterior rib 29 Length of fibula 15

Length of a posterior dorsal vertebra 5 Length of tarsal bone 6

Length of twenty-second caudal vertebra 5 Length of digit 27

Length of femur 25

Ichthycanthus platypus Cope.

Cope, Proc. Am. Phil. Soc., pp. 574, 575, 1877.

Cope, Trans. Am. Phil. Soc, xvi, p. 289, fig. 1, 1888.

Baur, Beitrage zur Morphogenie des Carpus und Tarsus der Vertebratcn, 1 Theil, p. 16, 1888.

Moodie, Science, n.s., xli, No. 1044, p. 34, 1915.

Moodie, Am. Jour. Sci., xxxix, pp. 509-512, fig. 2, 1915.

Type: Specimen No. 7954 G, and obverse, Department of Geology, Columbia
University. (Plate 23, fig. 1.)

Horizon and locality: Linton, Ohio, Coal Measures.

This amphibian is represented by the same portions of the skeleton as the preced-
ing species, furnishing a good basis for comparison. It is very well preserved, display-
ing the characters, especially of the hind foot, which is almost entirely represented.

Several features distinguish it from the /. ohiensis, one of which is of more than
usual value if correctly indicated by the fossil. There are 10 vertebras from the
anterior end to the sacrum preserved in place, and none of them supports a rib,
nor are there any ribs visible anywhere on the block of shale. I suspect that they
exist on more anterior vertebrae, or may have been displaced to a more anterior
position than they normally occupy. The abdominal chevrons are more anterior in
position than are those of the /. ohiensis. The hind legs are longer than in that
species; in this one the femur equals 7.5 vertebral centra in length. The external
digit, on the other hand, while bearing 5 phalanges, is distinctly shorter. The
fibular tarsal is of a transverse oval, not quadrate, form.

The dorsal centra are shorter and deeper than long; the neural arches are ele-
vated, with short but distinct zygapophyses, and a flat, subquadrate, superiorly
truncate, neural spine. They bear short, vertically compressed diapophyses near the
base of the arches . The neural spines of the caudal vertebrae become rapidly more slen-
der, and also diminish in length, while the zygapophyses are continued to the fifteenth
vertebra. The chevron bones are slender, and inclose a moderate haemal opening.

The femur is gradually expanded to the extremities. Proximally there is a tro-
chanteric ala, besides the obtuse head. Distally the condyles are well distinguished,

PIATE 2«

Diplodua
tooth

Photograph of the type specimen of I'elion lyelli Wyman, from the Ohio Coal Measures. Supposed to
represent the ancestral form of the Salientia. X 2. Original in American Museum of Natural History.

Scales of Cercariomorphus parvisquamis Cope, a microsaur from the Ohio Coal Measures. X 10. Original
in American Museum of Natural History.

Type specimen of Cercariomorphus pannsquamis Cope, from the Ohio Coal Measures. Original in
American Museum of Natural History. X 0.75.

THE MKROSAURIAN FAMILY ICHTHYCANTHIDjE. 173

the external or fibular being truncate. The fibula is less than three-fifths the length
of the femur, and is expanded at both extremities. Two proximal tarsals are dis-
tinct; the one next the fibula is larger than the other and transverse suboval in
form. It has a median dividing ridge as though composed of fibulare and inter-
medium coossified. The tibiale is subtriangular. There are five distinct phalan-
geal tarsals. The toes are, in the order of their lengths, beginning with the shortest,
1-2-5-3-4. Their phalanges (including metatarsals) are, in the proper order,
commencing with the hallux, 3-3-4-5 ?~5, the distal end of the fourth finger being
lost . These bones are rather stout, and the unguals are simply conic. The form of
the foot is short and wide. The number of the phalanges is nearly similar to that
found in Amphibamus grandiceps, excepting that in that species the fifth digit has
but 4. They are more numerous on most of the digits in Sauropleura digitata. Cope
(Trans. Am. Phil. Soc, xvi, 289, fig. 1, 1888) contributed the following note on
Ichthycanthus platypus:

"A reexamination of the type specimen of this species from the Coal Measures of Ohio,

preserved in the Museum of Columbia College, New York, enables me to refer this species

to the Rachitomi. The neural spines are distinct, showing that it belongs, probably, to the

Eryopidae. As the skull is not preserved, I can not determine the genus positively, but

refer it for the present to Eryops. I append a figure of the posterior foot, which displays

the characters of the tarsus of this group for the first time. The number of tarsals is as

in a Theromorph reptile, except that two elements represent the cuboid bone as in the

reptile, Stercosternum tumidum Cope; giving five elements in the distal tarsal row. There

is but one centrale and no intermedium. Two fragments of caudal vertebra; adhere to

the specimen."

Measurements of Ichthycanthus platypus Cope.

mm. mm.

Length of 10 dorsal vertebrae 45 Length of fibula 18

Length of 15 caudal vertebrae 55 Diameter of fibula proximally 7

Length of centrum of a dorsal 3.8 Width of sole at second row of tarsal bones 17

Total elevation of a posterior dorsal 14 Length of foot to end of third digit 31

Length of femur 32 Length of first digit 10

Diameter of femur medially 4.5 Length of third digit 22

Diameter of femur distally 8.3 Length of fifth digit 20

The writer has had the privilege of restudying this interesting specimen and has
already (484) described the foot and tarsus, as follows:

The only known specimen of this anomalous amphibian is incomplete, repre-
senting the posterior half of the skeleton, and an abundance of ventral scutellas or cal-
cified myocommata. The block of coal containing these interesting remains is from
Linton, Ohio, and is preserved in the geological collections of Columbia University,
from which institution Professor Grabau very courteously forwarded it for study.

Ichthycanthus platypus was described by Cope from the Linton, Ohio, Coal
Measures, locating it doubtfully in the Permian genus Eryops on account of the
unusual condition of the tarsus and reconsidering a former decision in favor of a
Coal Measures genus Ichthycanthus. In this disposition of the species into the Per-
mian genus he is followed by Hay (317) ; but Baur (28) regarded the form as a mem-
ber of the Coal Measures genus Ichthycanthus, after commenting on the later defi-
nition by Cope. The type of the genus, Ichthycanthus, to which Cope first allied the
species under consideration, is /. ohiensis, a supposed amphibian from the Coal
Measures of Linton, Ohio, founded on incomplete material.

174 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

The form combines in an unusual and remarkable degree reptilian and amphibian
characteristics. The leg bones, pelvis, and tarsus are all strikingly reptilian, but the
phalanges in the arrangement of elements are so typically amphibian that if we had no
other means of diagnosis we would incline to locate this Coal Measures species among
the Amphibia. The leg (plate 23, fig. 1) recalls in its structure that of another Coal
Measures species, Eosauravus copei Williston, which is, however, clearly a reptile.
While there is a general degree of similarity between the foot structure of Eosauravus
copei and Ichthycanthus platypus, yet there are very great differences in the phalan-
geal formula and the arrangement of the tarsal elements. These differences are clear
and indicate a separation of the two species into distinct classes. The phalangeal
formula in the Eosauravus, 2-3-4-5-4, is typically reptilian; while in the Ichthy-
canthus, 2-2-3-3-3, ^ is amphibian. The tarsus of the Ichthycanthus is amphibian
in the presence of an intermedium, but this is very small and the remaining tarsal
structures have- nothing which might not be found in an early reptile. There may
be a single or even two centralia in the reptilian tarsus among the early forms.

The amphibian nature of the species having thus been established, it remains to
give a detailed account of its skeletal anatomy, with comparative references to such
other ancient forms as are available. Little can be said of the vertebral column,
since only the molds of a few vertebrae remain, and these are so obscured by a
closely adherent pellicle of carbonaceous material that their form can not be dis-
tinctly discerned. They are high, with relatively broad neural spines. There are no
ribs preserved. The pelvis is obscured, but it is possible to determine the presence
of an elongate ilium and an ischium. The leg of the left side is the best preserved
of all the elements, and it is to this that our attention will be confined. The oppo-
site leg is not so complete, yet all the long bones and a part of the tarsus are pre-
served with sufficient clearness to corroborate the findings of the left side.

The femur, as has been stated, is reptilian in appearance. This is due to the
well-rounded articular surfaces, as though the endochondrium were well developed,
and to the large development of the greater and lesser trochanters, which are quite
prominent, though these are distorted and depressed in fossilization. The bone is
stout and well built and its form suggests an active habit of life. The tibia and
fibula are separate, and do not otherwise have sufficiently noteworthy characters
to call for a special description in this place, except to note an unusual anterior
crest on the tibia. To the lower ends of these bones articulate the first row of tarsal
elements, the tibiale, intermedium, and fibulare. The tarsus is composed of 9
elements arranged in 3 rows. The proximal row is composed of the tibiale, the
intermedium, and the fibulare. On the edge of the tibiale there lies a portion of
one of the caudal vertebrae, so that the form of this tarsal element is slightly ob-
scured. The intermedium is a small, rounded element lying between the larger
elements. The fibulare is rectangular and projects a considerable distance out from
the tibia, but articulates directly with the large lateral distal tarsal. The centrale
is triangular in form and is opposed directly by the tibiale and tarsalia 1 to 3.
The phalanges are robust in appearance. The entire foot gives one the impression
of a very broad structure. The ungual phalanges were apparently bluntly clawed.

CHAPTER XXIII.

SUPPOSED MICROSAURIAN SPECIES OF UNCERTAIN RELATIONSHIP.

The following three species are so unusual and so incompletely known that they
can not be considered with any of the above families: Brachydectes newberryi Cope,
Linton, Ohio; Amblyodon problematicum Dawson, Nova Scotia; Proterpeton gurleyi
Moodie, Danville, Illinois.

Genus BRACHYDECTES Cope, 1868.

Cope, Proc. Phila. Acad. Nat. Sci., 1868, p. 214.

Cope, Trans. Am. Phil. Soc, 1868, p. 14.

Cope, Geol. Surv. Ohio, n, pt. 11, p. 388, 1875, pi. xxvii, fig. 2.

Type: Brachydectes newberryi Cope.

Cope (Geol. Surv. Ohio, vol. n, pt. n, p. 388, 1875), says:

" This genus is indicated by two rami of a mandible and a portion of a premaxillary only.
These, when compared with those of (Estocephalus and Tuditanus, from the same locality,
and with others described by authors, are so much stouter, i.e., shorter and more elevated,
that they evidently belong to a genus unlike either. The genus further differs from (Esto-
cephalus in having the teeth of equal size to the posterior part of the series ; that is, to the
base of the elevated coronoid process. The teeth are elongate cylindrical cones, with their
acute tips turned a little posteriorly. The fractured ones display a large pulp cavity. The
three premaxillaries preserved are similar, but without curvature at the tips. They do not
exhibit striae or any other sculpture.

"So far as the remains known go, the genus is nearer Hyierpeton than any other.
According to Dawson, that genus is provided with a large canine-like tooth, at the anterior
extremity of the maxillary, on the inner row, which is inserted into a distinct socket. No
such tooth appears among those of this genus. The latter does not give any indication of
the very elevated coronoid process of Brachydectes, though the external portion of the
dentary bone in that region being lost, little can be said about it."

Brachydectes newberryi Cope.

Cope, Proc. Phila. Acad. Nat. Sci., p. 214, 1868.

Cope, Trans. Am. Phil. Soc, 1868, p. 14.

Cope, Geol. Surv. Ohio, 11, pt. 11, p. 388, pi. xxvii, fig. 2, 1875.

Type : Specimen No. 8604 G, American Museum of Natural History.

Horizon and locality: Linton Ohio, Coal Measures.

Cope (Geol. Surv. Ohio, vol. n, pt II, p. 388, 1875) says of this form:

" The species is represented by one nearly perfect ramus mandibuli, one dentary bone
and one premaxillary, probably not complete.

' ' The dentary bone appears to have been attached by suture to the articular and angular,
as its free margin has very much of the outline of that suture in Amphiuma and lizards. The
coronoid process would also seem to be a part of the same bone as in Amphiuma and Meno-
pcma, and not composed of the coronoid bone as in lizards. It rises immediately behind the
last tooth, and displays no suture.

175

I76 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

"The lower portion of the dentary is prolonged into an acute angle. This is separated
by a deep and wide concavity from the superior posterior prolongation, which is obtuse,
and rises at once into the coronoid process. Teeth on this dentary, seven ; the same number
is on the preserved ramus ; this number is suspected to be complete, or nearly so. The teeth
terminate at the obvious termination of each ramus, which is, it is true, slightly obscured.
These teeth are the longest in the Microsauria in relation to the depth of the ramus, equal-
ing the largest in (Estocephalus. They are doubtless exposed, as are some of those of the
last-named genus, by the splitting away of the outer parapet of the dentary bone. As no
traces of alveoli have been thus rendered visible I suspect the dentition to have been
acrodont, as in some existing Batrachia.

' ' No external surface of the mandible remains, but there are no impressions of sculpture
on the matrix. A little external face of the premaxillary displays none.

" The species is dedicated to Professor John S. Newberry, the able director of the Geo-
logical Survey of Ohio, and discoverer of most of the Batrachia herein described."

Measurements of the Type.

mm. mm.

Length of ramus of mandible (imperfect) 22 Length of dentary 16

Depth at last tooth 5 Depth at coronoid process 7.5

Length of exposed tooth 3.5 Depth at first tooth 3

Genus PROTERPETON new genus.

Type: Proterpeton gurleyi Moodie.

Known from a single vertebra. Spine very high and heavy, the neural canal
large.

Proterpeton gurleyi new species.

Type: Specimen No. 13,296, Walker Museum, University of Chicago.

Horizon and locality: Coal Measures near Danville, Illinois.

The vertebra, as preserved, is well characterized by the figure (plate 22, fig. 2).
The spine is high and heavy, the neural canal is large, and the centrum reduced.
The form is very unusual. It is apparently from the cervical region, as there are no
indications of zygapophyses, transverse processes, or haemal arches, although they
may have been abraded; apparently not, however. The type specimen was dis-
covered near Danville, Illinois, about the horizon of the Danville coal, so that it is
quite high in the Allegheny series of the Pennsylvanian and of about the same
horizon as the phalangeal bone from Breeze, Illinois, which may be provisionally
associated with this form. There is no assurance that Proterpeton gurleyi is an
amphibian. The vertebra may have belonged to a fish.

Measurements of the Type of Proterpeton gurleyi Moodie.

mm. mm.

Entire height of vertebra 24 Height of neural canal (crushed ?) 6.5

Width at side of neural canal 21.5 Width of vertebral centrum anteroposteriorly 5.5

Width of neural canal 13 Height of neural spine from top of neural canal ... 9

Genus AMBLYODON Dawson, 1882.
Dawson, Phil. Trans. Roy. Soc. London, pt. 11, p. 644, pi. 40, figs. 57-61, 1882.

Type: Amblyodon problematicum Dawson.

This genus was described by Dawson in 1882 from very imperfect remains. He
says that it is "characterized by stout cylindrical teeth, blunt at the apices; but
otherwise imperfectly known."

SUPPOSED MICROSAURIAN SPECIES OF UNCERTAIN RELATIONSHIP.

177

Amblyodon problematicura Dawson.
Dawson, Phil. Trans. Roy. Soc. London, pt. u, p. 644, pi. 40, figs. 57 to 61, 1882.

TyPe: Specimen No. 3061-10, Peter Redpath Museum, McGill University-
Horizon and locality: Coal Measures of Nova

Scotia.

A fragment of a jaw 1 cm. in length has 10

cylindrical teeth, simple and smooth, with large

pulp cavities and rounded regularly at the apices.

With these are 4 vertebrae of the usual type,

measuring together 1 cm. Fragments of cranial

bones also occur and are obscurely pitted. There

is also what seems to be the shaft of a limb

bone and a few oval scales. A flat and some-
what rhombic bone, with a style at one side,

may possibly be a thoracic plate or possibly a

parasphenoid.

The material is too scanty for any satisfactory description of this animal, but it

is provisionally named Amblyodon problematic urn.

Fig. 38. — Skeletal elements of Amblyodon sp.
from the Coal Measures of Nova Scotia,
a, tooth, X 25; b, section of tooth, X 25;
d, fragment of thoracic plate; /, shaft of
limb bone; e, rib.

CHAPTER XXIV.

THE TEMNOSPONDYLOUS AMPHIBIA OF THE COAL MEASURES
OF NORTH AMERICA.

DEFINITION OF THE ORDER TEMNOSPONDYLIA, ZITTEL, 1887.

Zittel, Handbuch der Paleontologie, Abth. I, Bd. 3, p. 384, 1887.

Terrestrial or semi-aquatic vertebrata; skull bones pitted and grooved ; lateral-
line canals present in well-developed form; pineal foramen sometimes absent;
sclerotic plates present ; vertebras rachitomous or embolomerous ; notochord partly
persistent; one or two sacral vertebrae; tail present, long or short; limbs and
girdles well developed; limb bones well ossified and bones of arm and leg sepa-
rate; pectoral and pelvic girdles composed of the usual stegocephalian elements;
an osseous pubis present; a cleithrum present on the scapula; carpus and tarsus
ossified, carpals 11 and tarsals 12 in one form; phalangeal formula, 2, 3, 3, 4, 2
for the hand and 2, 3, 3, 3, 2 for the foot; fore and hind limbs pentadactyl in a
few forms; venter covered with an armature of osseous scutes, sometimes over-
lapping; skin of back bare or armored with heavy plates; ribs heavy, double-
headed, curved and moderately long, or short ; body short and heavy, as compared
to skull about 2 to 1 .

Range: Coal Measures to upper Permian.

Distribution: North America: Illinois, Kansas, Oklahoma, Texas, and Penn-
sylvania; Europe: Germany, Bohemia; France; Asia: India.

Family CRICOTID^ Cope, 1884.

Cope, Am. Nat,, xvm, p. 38, 1884.

General form of the body elongate, with triangular skull and short, stout
limbs. Snout narrow, orbits large, elongated oval, situated near the middle of
the skull. External bones faintly sculptured, sensor}- canals conspicuous, parietal
foramen large. Teeth conical, of unequal size. Presacral vertebras composed of
horseshoe-shaped pleurocentra and hypocentra, the former alone supporting the
neural arch. In the caudals the pleurocentra and hypocentra form complete
rings, and both elements take part in the support of the neural arch, but the
haemal arch is borne exclusively by the hypocentra. A close abdominal armor of
imbricate scales, arranged in a chevron pattern. Caudal vertebrae numerous.
Chevrons cobssified with the intercentra.

Genus SPONDYLERPETON Moodie.

Moodie, Kans. Univ. Sci. Bull., vi, No. 2, p. 355, 1912.

Type: Spondylerpeton spinatum Moodie.

The genus is based on a specimen consisting of 9 imperfect vertebrae, from the
caudal region of a relatively large amphibian. The present genus exceeds Diplo-
spondylus from the Gaskohle of Bohemia (251) by twice its size and is about two-
thirds the size of Cricotus heteroclitus Cope (98) from the Permian of Kansas. The
vertebras are twice as high as wide, differing thus from Cricotus, in which the ver-
tebras are nearly circular. A character which is of great importance is the large size
of the intercentrum, which almost equals the pleurocentrum in size. It is similar
to the pleurocentrum in shape, except for the attached neurocentrum and chevron on
the latter. The present genus differs from Diplospondylus in the greater length of

178

THE TEMNOSPONDYLOUS AMPHIBIA.

179

the intercentrum and pleurocentrum, in the greater size, in the larger proportions
of the neurocentrum, and the greater proportionate size of the intercentra.

Spondylerpeton spinatum Moodie.
Moodie, Kans. Univ. Sei. Bull., vi, No. 2, pp. 355-357, pi. 8, figs. 1 and 2; pi. 9, fig. 1, 1912.

Type : Specimen No. 793 (26) and obverse, Yale University Museum.

Horizon and locality: Mazon Creek, near Morris, Illinois.

The species is very imperfectly known. Sufficient is present, however, to show
its wide generic differences from other forms of the Cricotidae. These characters
are of a phylogenetic nature and indicate the more primitive nature of the present
form, as we would expect from its geological position. The sutures separating the
four vertebral elements are clearly ap-
parent. The pleurocentral-neurocentral
suture is apparent in 4 vertebras.

There is but a single pleurocentrum
preserved complete. This shows the
form of the attached neurocentrum and
chevron, which corresponds to the hypo-
centrum pleurale according to Fritsch.
The pleurocentrum is flattened laterally,
with a rather large canal for the noto-
chord. Its sides are marked with 4
longitudinal grooves. Surfaces for the
attachment of the ribs are not present,
and for this reason, as well as the pres-
ence of chevrons, the vertebras are sup-
posed to be caudals. As such they rep-
resent an animal of some 3 or 4 feet in
length. It was the giant of the Mazon
Creek Amphibia. (Plate 4, figs. 1, 2.)

Attached to the upper side of the
pleurocentrum by a sutural union occurs
the neurocentrum. The neural arch is quite large and is oval in outline, although
somewhat constricted at the tip. The spine of the neurocentrum is rather long and
broad at the base, measuring 12 mm. across the anterior zygapophysis. The neuro-
centrum is laterally flattened and ends in a rather acute and somewhat rugose
point. It was probably tipped with cartilage. The anterior zygapophysis occurs
well down on the neurocentrum, its lower edge being 5 mm. from the suture sepa-
rating the pleurocentrum and the neurocentrum. The posterior zygapophysis occurs
quite high up on the neurocentrum and lies at a distance of 15 mm. from the pleuro-
neurocentral suture, thus indicating an extreme posterior inclination of the neural
spine. The posterior zygapophysis of the best preserved vertebra is separated
from its mate, the anterior zygapophysis, in the next succeeding vertebra by a
space of 5 mm.

Fig. 39. — The vertebrae of Spondylerpeton spinatum
Moodie, the only known temnospondyle from the
Mazon Creek shales. X I. hy, hypocentrum; inc,
intercentrum; pc, pleurocentrum; nc, neurocentrum.

180 THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

The ventral surface of the pleurocentrum bears a structure, which is, without
doubt, a chevron, although the character of its opening can not be determined. It
is elongated and is united by a broad base to the pleurocentrum. Its union is by a
clearly defined suture, which is apparent in 3 vertebrae. The condition represented
by the specimen duplicates almost exactly the condition figured by Cope for the
caudal region of Cricotus Cope.*

The intercentrum of the present form is fully as large as the pleurocentrum.
The significance of this has already been mentioned. The body of the centrum is
pierced by a large notochordal canal.

Measurements of the Type of Spon'dyi.erpeton spinatum Moodie.

mm. mm.

Length of specimen 60 Height of intercentrum 10.5

Length of pleurocentrum 1 1 .5 Height of chevron 3

Height of pleurocentrum to base of neurocentrum . 20 Length of chevron 18

Length of neurocentrum 33 Width of notochordal opening in centrum 5

Width of neurocentrum at base 9 Height of same 4.5

Width across anterior zygapophysis 12 Height of neural canal 12

Width across posterior zygapophysis 10 Greatest width of neural canal 6

Length of intercentrum 10

Family ERYOPIDiE Cope, 1882.

Cope, Am. Nat., xvi, p. 334, 1882.

Large, terrestrial or amphibious vertebrata; skull bones deeply marked with
pits and grooves which take the form of lateral-line canals; infra- and supra-
orbital canals, antorbital commissure, jugal canal, and occipital cross-commissure
of the lateral-line system present in Eryops megacephalus Cope ; carpus and tarsus
osseous; pubis an osseous plate, surrounded in life by a large amount of cartilage;
fore and hind limbs pentadactyl ; orbits, in the typical genus, located far back on
the skull and near the median line ; cleithrum present on the scapula ; vertebrae
rachitomous, the intercentrum supporting the arch in the dorsal region ; para-
sphenoid well-developed or reduced; teeth on pterygoids, palatines, prevomers,
and parasphenoid.

Range: Upper Pennsylvanian to Permian.
Distribution: America, Europe, Asia.

Genus ERYOPS Cope, 1877.

Type: Eryops megacephalus Cope.

Skull long, comparatively narrow; proportion of length to breadth about 9
to 7. Roof bones coarsely sculptured posteriorly, finely sculptured anteriorly.
Nasals and premaxillae very large; frontals excluded from the orbits by junction
of pre- and post-frontals. Pterygoids not meeting in the median line ; parasphe-
noid dagger-shaped, tapering gradually to a point just in front of the palatine
foramen; prevomers large. Orbits subcircular, situated in the posterior half of the
skull; nares subovate, remote, at a considerable distance from the tip of the skull.
Many minute denticles, on pterygoids, palatines, prevomers, and parasphenoid.
Teeth circular in cross-section, strongly ribbed near the base, dentine strongly
infolded. Three large teeth on each palatine. Mandible without postcotyloid
process. Vertebras rachitomous. Ribs double-headed. Pubis osseous. Three
species (Permian), E. megacephalus, E. lotus, and E. willistoni, are assigned to
the genus.

Eryops is represented in the Carboniferous deposits of North America by incom-
plete vertebral remains described by Case (94) from near Pittsburgh, Pennsylvania.

*Cope, E. D., Trans. Am. Phil. Soc, xvi, p. 246, 1890.

MOODIE

PLATE 25

■\>,

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. Nat., xxx, 936, pi. xxi.

-. 1896. The ancestry of the Testudinata.

Amer. Nat., xxx, 399.
. 1896. Appendix on a species of Trimero-

rhachis. Proc. Amer. Phil. Soc, xxxv, 137.
. 1896. The reptilian order Cotylosauria.

Supplement: Some new Batrachia from the

Permian beds of Texas. Proc. Amer. Phil. Soc,

xxxiv, 455.

1897. On new Paleozoic Vertebrata from

183.

184.

185.

t86.

187.

Illinois, Ohio, and Pennsylvania. Proc. Amer.
Phil. Soc, xxxvi, 71-90, three plates.

. 1898. Syllabus of Lectures on the Verte-
brata. Batrachia, 43-51. Published for Uni-
versity of Pennsylvania.

Cox, E. T. 1874. Colettosaurus indianaensis. 5th
Ann. Rept. Geol. Surv. Indiana, 247, pi. 1.

Credner, Hermann. 1881. Ueber Branchiosaurus
amblystomus, einen neuen Stegocephalen aus den
Rothliegenden-Kalke von Niederhasslich im
Plauen'schen Griinde. Sitzungsb. naturf. Gesell.
zu Leipzig, 43.

. 1 88 1. Die Stegocephalen und Saurier aus

dem Rothliegenden des Plauen'schen Grundes bei
Dresden. I Theil: Einleitung und Branchio-
saurus gracilis. Zeit. d. deutsch. geol. Gesell.,
xxxm, 298-329, taf. xv-xviii.

. 1 88 1. Die Stegocephalen und Saurier, etc.

II Theil: Branchiosaurus amblystomus. Zeit.
d. deutsch. geol. Gesell., xxxm, 574-603, taf.
xxii-xxiv.

. 1881. Ueber einige Stegocephalen (Laby-

rinthodonten) aus dem sachsischen Rothliegen-
den. Sitzungsb. naturf. Gesell. Leipzig, 1-11.
(Note on Branchiosaurus and Melanerpeton, 43,
45-)

. 1882. Ueber Melanerpeton Fritsch aus dem

Rothliegenden-Kalk von Niederhasslich im Plau-
en'schen Grundes. Sitzungsb. naturf. Gesell.
Leipzig, 45-47-

. 1882. Die Stegocephalen und Saurier, etc.

III Theil: Pelosaurus laticeps, Archegosaurus
decheni, A. latirostris. Zeit. d. deutsch. geol.
Gesell., xxxi v, 213-237, tafeln xii-xiii.

. 1883. Die Stegocephalen und Saurier, etc.

IV Theil: Branchiosaurus gracilis, Acanthos-
toma vorax, Melanerpeton, Discosaurus permi-
anus. Zeit. d. deutsch. geol. Gesell., xxxv, 275-
300, tafeln xi-xii.

. 1885. Die Stegocephalen aus dem Roth-
liegenden des Plauen'schen Grundes bei Dresden.

V Theil: Melanerpeton pulcherrimum, Pelo-
saurus laticeps, Archegosaurus; Sparagmites
arciger, Hylonomus fritschii, Hyloplesion. Zeit.
d. deutsch. geol. Gesell., xxxvn, 694-736, tafeln
xxvii-xxix.

1886. Die Stegocephalen aus dem Roth-

liegenden des Plauen'schen Grundes bei Dresden.
VI Theil: Die Entwickelungsgeschichte von
Branchiosaurus amblystomus Cred. Zeit. d.
deutsch. geol. Gesell., xxxvm, 576-633, tafeln
xvi-xix.

188. Credner, Hermann. 1886. Archegosaurus Rcste
aus dem Rothliegenden v. Offenbach. Zeit. d.
deutsch. geol. Gesell., xxxvm, 696.

1890. Die Stegocephalen und Saurier aus

189

190.

191.

192.

193-

194.

dem Rothliegenden des Plauen'schen Grundes bei
Dresden. IX Theil: Hylonomus geinitzi, Petro-
bates truncatus, Discosaurus permianus. Zeit.
d. deutsch. geol. Gesell., xlh, 240-277, tafeln
ix-xi, figs.

1887. Stegocephalen des Rothliegenden.

Wandtafeln mit Erlauterungen, Leipzig. Folio.
Tafel i, Branchiosaurus amblystomus; Tafel ii,
Pelosaurus laticeps and Melanerpeton pulcher-
rimum.

. 1888. Wandtafeln mit Stegocephalen des

Rothliegenden. Neues Jahrb. f. Mineral. Geol.
u. Paleon., Bd. I, 67-69.

. 1 89 1. Die Urvierfussler (Eotetrapoda) des

Sachsischen Rothliegenden. Allgemein-verstand-
liche naturwiss. Abhandl., Heft 15. Sonder-
Abdruck aus der " Naturwisser.schaftlichen Woch-
enschrift," 4-52, 53 figs.

. 1893. Die Stegocephalen und Saurier aus

dem Rothliegenden des Plauen'schen Grundes bei
Dresden. X Theil: Sclerocephalus labyrinthicus
H. B. Geinitz. Zeit. d. deutsch. geol. Gesell.,
xlv, 639-704, tafeln xxx-xxxii.

— - — . 1893. Zur Histologie der Faltenzahne
l'aleozoischer Stegocephalen. Abhandl. Sachs.
Gesell. Wiss., xx, 545-551.

195. . 1897. Elemente der Geologic Amphibia,

489-494.

196. Cummins, W. F. 1908. The localities and horizons

of Permian vertebrate fossils in Texas. Jour.
Geol., xvi, 737-745-

197. Cuvier, Georg. 1825. Des Batrachiens fossiles.

Recherches sur les Ossemens Fossiles, 3d ed., 431-
444, v, pt. u. (Homo diluvii testis et theoskopos.)

198. Davison, Alvin. 1895. A contribution to the

anatomy and phylogeny of Amphiuma means.
Jour. Morphol., xi, 375-410, 2 pis. (See Kings-
ley, Tufts College Studies, No. 7.)

199. Davis, James W. 1883. On the occurrence of

remains of Labyrinthodonts in the Yorkdale
rocks of Wensleydale (Carboniferous). Rept.
53d Meeting Brit. Assn. Adv. Sci., Southport,
Sept., 492; see also Nature, xxvm, 578.

200. Dawson, J. W. 1853. On the Coal Measures of the

South Joggins, Nova Scotia. Quart. Jour. Geol.
Soc, x, 1-41, with bibliography of papers on the
coal fields of Nova Scotia.

201. •. 1855. Acadian geology. Edinburgh, 1855,

Amphibia, 160.

. 1887. Carboniferous quadrupeds. The

Story of the Earth and Man, 143-150. 9th ed.
1855. Notice of the discovery of a reptilian

202.

203.

skull in the Coal Measures of Pictou (Nova

Scotia). Quart. Jour. Geol. Soc, xi, 8-9.
204. . 1859. 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. [A descriptive
account of the remains of land animals found in
the Coal Formation of Nova Scotia, with remarks
on their bearing on the theories of the formation
of coal and of the origin of species.) See
also Jour. Geol. Soc., xvi, 273; xvm, 5-19,
470; also Neues Jahrb. Mineral., Geol. u. Paleon.,
1864, 507.

. 1 863. Notice of a new species of Dendrerpe-

ton and of the dermal covering of certain Carbo-
niferous reptiles. Quart. Jour. Geol. Soc., xix,
469-473-

. 1863. Air-breathers of the Coal Period.

Canadian Naturalist and Geologist, vm, 1-12,
81-92, 161-175, 268-295, with pis. 1-6.

1 863. The Air-breathers of the Coal Period.

Amer. Jour. Sci. (2), xxxvi, 430-432.
— - — . 1868. Land animals of the Coal Period.

Acadian Geology, 1868, 2d ed., chap, xvm, 353-
382.

1870. Note on some new animal remains

from the Carboniferous and Devonian of Canada.
Geol. Mag., vil, 87; Quart. Jour. Geol. Soc.,
xxvi, 166-167; Canadian Naturalist (2), v, 98-99.
(Same paper in all: Baphetes minor.)
. 1876. 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. The Oldest Air-breathers, chap, x,
259-307, with restorations and figures.

. 1894. Preliminary note on the recent dis-
coveries of Batrachians and other air-breathers in
the Coal Formation of Nova Scotia. Canadian
Record of Science, vi, 1-7, I fig.

. 1895. Synopsis of the air-breathing ani-
mals of the Paleozoic of Canada (up to 1894).
Trans. Roy. Soc. Canada, XII, sec. IV, 71-78.

224.
225.

226.

I 227.
228.

229.

230.

231-
232.

233-
234-
235-
236.

237-
238.

239-
240.

Deichmieli.er, J. V. 1882. Ueber "Die Stego-
cephalen aus dem Rothliegenden des Plauen'schcn
Grundes bei Dresden. Ill Theil: von H. Cred-
ner." Sitzungsb. Isis, Dresden, 71.

. 1884. Ueber Branchiosaurus petrolei,

Nachtrage zur Dyas III. Mittheilungen des kgl.
mineral, geol. u. praehist. Museum in Dresden,
Heft 6, Paleontol. Abhandl. aus dem Dresdener
Museum, 5-17, Taf. 1.

Dollo, Louis. 1884. Note sur le Batracien de
Bernissart. Bull. Mus. Roy. Hist. Nat. Belg ,
m. 85~93. pl- i»- (Hylasobatrachus croyii from
the Wealden.)

. 1885. Les Labyrinthodontes. Rev. Quest.

Scientif. Bruxelles, xvn, 305-312.

■ '895. Sur la Phylogenie des Dipneustes.

Bull. Soc. Beige de Geol. Paleont. Hydrol., ix,
79-178, pis. v-x. (Discusses relations to the
Amphibia.)

Dunlop, Robert. 1910. The fossil Amphibia in the
Kilmarnock Museum previous to the fire of
November 26, 1909. Annals Kilmarnock Glen-
field Ramblers Society, No. 6, 10, 2 pis.

Eastman, Chas. R. 1902. The Carboniferous fish
fauna of Mazon Creek, Illinois. Jour. Geol., x,
535-541. with list of fossil vertebrates.

. 1902. Translation of Zittel's Text-book of

Paleontology, vol. II, 114-139, figs. 193-235.

-. 1907. Devonic fishes of the New York

Formations. Memoir No. 10, New York State

Mus., 180-183.
Eichwald, E. 1848. On Zygosaurus lucius from the

Permian of Russia. Bull, de la Soc. imp. des

Natl, de Moscou, xxi, 159.
. i860. Letha?a rossica ou Paleontologie de

la Russie, I, Stuttgart. 1620-1633, tafeln 57-59,

Labyrinthodonten.
Embleton and Atthey. 1874. On the skull and

other bones of Loxomma. Ann. and Mag. Nat.

Hist., ser. 4, xiv, 38-63, pis. iv-vii.
Emery, C. 1897. Die fossilen Reste von Archego-

saurus und Eryops und ihre Bedeutung fur die

Morphologie des Gliedmassenskelets. Anat. Anz.,

xiv, No. 8, 201-208, figs. 1-7.
Emmons, E. 1857. Perm und Trias System in Nord

Carolina. Edinb. n. Phil. Jour., v, 370.
. 1857. Dictyocephalus elegans Leidy.

American Geology, pt. vi, Labyrinthodonts, 59,

figs- 31-32-

-. 1863. Dictyocephalus elegans Leidy. Man-

242.

243-

ual of Geology, New York. Labyrinthodonts,

180, 184.
Emmrich, A. 1852. Bericht ueber die Naturfor-

scherversammlung in Gotha im Jahre 1851. Jahrb.

d. k. k. geol. Reichsanstalt, p. 155, Wien.
Etheridge, Robert. 1866. On the discovery of

several new Labyrinthodont reptiles in the Coal

Measures of Ireland. Geol. Mag., m, 4. (See

Huxley, 1867.)
Fraas, E. 1S89. Die Labyrinthodonten der Schwtb-

iscben Trias. Paleontographica, XXXVI, 1-158,

with bibliography.
. 1896. Die Schwabischen Trias-Saurier.

Fcstgabe des koniglichen naturalien Cabinets

in Stuttgart zur 42 Versammlung dcr deutsch.

geol. Gcscll., Stuttgart, 1-8, pis. i-ii.

208

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

244. Fraas.E. 1901. Labyrinthodon aus dem Buntsand-
stein von Teinach. Jahreshefte des Vereins f.
vaterlandische Naturkunde in Wurtemberg, Jahr.,

LVH, 318-320.

-. 1903. Die geognostische Sammlung Wur-

245-

246.

247.

tembergs im Parterre-Saal zugleich ein Leitfaden
fur die geologischen Verhaltnisse und die vor-
weltlichen Bewohner unseres Landes. 2d Auflage.
1903. Rana danubina H. von Meyer var.

rara, 0. Fraas aus dem Obermiocan von Stcin-
heim. Jahreshefte des Vereins f. vaterl. Natur-
kunde in Wurtemberg, Jahr. 1903, 105-110,
with fig.

-. 1909. Rana haufiftana n. sp. aus den Dyso-

dilschiefern des Randecker Maares. Jahreshefte
des Vereins f. vaterl. Naturk. in Wurtemberg,
Jahr. 1909, 7, with 1 fig.

247a. . 1913. Neue Labyrinthodonten aus der

schwabischen Trias. Paleontographica, Bd. lx,
275-294, pis. xvi-xxii, with text-figs. 1-5.

248. Frech, Fritz. 1901. Lethaea geognostica oder

Beschreibung und Abbildung der fur die Gebirgs-
formation bezeichnendsten Versteinerungen. I
Theil, Lethaea palaaozoica, 2 band. Die Dyas,
459-471. 22 figs., pi. 56.

249. Fritsch, Anton. 1870. Sitzungsberichte der k.

Gesell. d. Wissen., April 27, 1870. (Describes 2
new species of Amphibia.)

1875. Ueber die Fauna des Pilsner und

250.

251

252.

253-

Rakonitzer Beckens. Sitzungsb. d. k. Boehm-
ischen Gesell. d. Wissenschft., March.

1883-1899. Fauna der Gaskohle und der

Kalksteine der Permformation Boehmens. Bd. 1,
1-182, 116 figs. Tafeln i-xlviii; Bd. II, 1-64, figs.
1 17-142, Tafeln xlix-lxx; Bd. IV, supplement 85-
98, figs. 383-394, Tafeln clxiii-clxiv. This is the
chief work on extinct Amphibia.

189,5. Ueber neue Wirbelthiere aus der

Permformation Boehmens, nebst einer Ueber-
sicht der aus derselben bekannt gewordenen
Arten. Sitzungsber. d. k. Boehmisch. Gesell. der
Wissenschaften, Prag, 1.

-. 1905. Vorlaufige Notiz ueber .Miscellanea

Paleontologica aus Boehmen und America.
Sitzungsber. d. k. Boehm. Gesell. dej Wissen-
schaften, Prag, Kammplatten, 3.

254. Fritsch, Karl Freiherr. 1879. Referat ueber

Gaudry's Sur les Reptiles des Temps primier;
Nachweis von Protriton petrolei bei Oberhof in
Thueringen. Neues Jahrb. f. Mineral. Geol. u.
Paleon., 720.

255. Fuchs, Hugo. 1909. Betrachtungen ueber die

Schlafengegend am Schadel der Quadrupeda.
Anat. Anz., xxxv, No. 5/7, 1 13-167, 25 figs.

256. Gadow, Hans. 1896. On the evolution of the

vertebral column of Amphibia and Amnipta.
Phil. Trans. Roy. Soc. London, ser. B, clxxxvii,
I_57» 5° figs-. and bibliography.

257. . 1901. Amphibia and reptiles. Cambridge

Nat. Hist., vm, Stegocephala, 80. London.

258. Gaudry, Albert. 1866. Note sur le Reptile dS-

couvert a Muse. Comptes Rendus de l'Academie
de Science, August, 341.

259. . 1867. M6moire sur le Reptile decouvert

par M. Frossard a Muse. Nouv. Archiv du
Musee d'Hist. Nat., m, 22-40, pi. iii.

260. - — . 1868. Note sur L'Actinodon frossardi.

261.
262.

263.

264.

265.

266.

267.
268.

269.

270.

271.
272.
273-

274.

275-
276.

277-

278.
279.

280.

281.
282.

283.

Gaudry, Albert. 1874. Les Ettcs de Temps prim-
aire. Revue Scientifique, ser. 2, XIII, 993.

. 1875. Les Reptiles des Schistes bitumi-

neaux dAutun. Bull. Soc. Geol. de France, 3d
ser., iv, 720.

-. 1875. Sur la Decouverte de Batraciens

proprement dits dans le Terrain primaire. Comp-
tes Rendus, lxxx, 441-443.

. 1875. Sur la Decouverte de Batraciens

dans le Terrain primaire. Bull, de la Soc. Geol.
de France, ser. 3, m, 300-306, pis. vii-viii. (Sec
also 263 above; Jour. Zool., iv, 38-41, 342.)

. 1876. Les Reptiles des Schistes bitumi-

neux d'Autun. Bull. Soc. Geol. de France, ser.
Ill, IV, 720-723, pi. xxii; 1879, VII, 62; 1885, 3d
ser., xm, 44.

. 1878. Sur les Reptiles de Temps primaires.

Comptes Rendus, lxxxvii, 956.

1878. Les Vertebres fossiles des environs

dAutun. Bull. Soc. Geol. France, vn, 62.

-. 1878. Les Reptiles d. l'Epoque Permienne

aux Environs d'Autun. Bull. Soc. Geol. de France,
ser. 3, vii, 62-76, pis. iii-iv.

. 1880. Sur un Reptile tres perfectionne'

trouve' dans le terraine Permien. Comptes
Rendus, xci, 669-671.

— . 1 88 1. Sur les plus anciens Reptiles trouves

en France. Comptes Rendus Acad. Sci. Paris
xcn, 1143-1145.

. 1 88 1. On a highly organized reptile from

the Permian Formation. Amer. Nat., vn, 67-71.
1883. Les Reptiles primaires. Arch. Zool.

Exper., 2d ser., I, 5-30, pis. 1-7. with figs.

1 883. Les Enchainements du Monde animal

dans les Temps geblogiques. Fossiles Primaires,

with 4 figs. Paris.
. 1884. Nouvelle Note sur !es Reptiles

Permiens. Bull. Soc. Geol. de France, 3 ser.,

xm, 44-51, pis. iv-v. Paris.
. 1884. Nouvelle Note sur les Reptiles

Permiens. Comptes Rendus Acad. Sci. Paris,

xcix, 737-738.
— . 1885. Paleontology in Germany and

America. Revue Sci., 7, Paris; Geol. Mag. (3),
11, London.

1886. Sur un nouveau Genre de Reptile du

Permien d'Autun. Bull. Soc. Geol. de France, ser.
in, xiv, 430-433, taf. 23.
. 1887. L'Actinodon. Nouvelles Archives

Mus. Hist. Nat., Paris, 2d ser., x, 30 pp., 3 pis.
. 1888. Les Vertebres fossiles des environs

d'Autun. Mem. Soc. d'Hist. nat. d'Autun, 90

pp. et 6 planches.
. 1885. Sur les Similitudes que plusiers

Reptiles ont eues dans divers Pays de Monde ver

la Fin des Temps primaires. Compte Rendus de

la 3d session Congres gcologique international a

Berlin, 1-4.
. 1888. Les Vertebras fossiles des environs

d'Autun. Autun.

1 890. Les Enchainement du Monde Animal

Bull. Soc. Geol. de France, Ser. 2, xxv, 576.

dans les Temps Geologiques, Fossiles Secondaires.
Amphibia, 169.
Gaupp, Ernst. 1895. Beitrage zur Morphologie
des Schadels. Ill: Zur vergleichenden Anatomie
der Schlafengegend am knochernen Wirbelthier-
Schadel. Morphologische Arbeiten herausge-
geben von G. Schwalbe, iv, 1, 77-128.

BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA.

209

284. Gegenbaur, Carl. 1862. Untersuchungen zur ver-

gleichen. Anatomic dcr Wirbelsaule bei Amphi-
bien und Reptilien. Leipzig.

285. . 1895. Clavicula und Cleithrum. Morphol.

Jahrb., xxm, 1-20.

286. Geikie, ARCHinALD. 1905. Scheuchzer's "Fossil

Man." Founders of Geology, 2d ed., 98-99.
London.

287. Geinitz, H. B. 1864. Palaeosiren beincrtii Gein.

ein neues Reptile aus der unteren Dyas von
Oelberg bei Braman. Neues Jahrb. f. Mineral.,
Geol. u. Paleon., 513-516.

288. . 1885. Ueber Thierfahrten in der Steinkohl-

enformation von Zwickau. Festschrift Natur.
Gesell. Isis, 63-66, pi. 2.

und Deichmueller. 1882. Ueber die fos-

289.

290.

silen Saurier in dem Kalke des Rothliegenden bei
Dresden. Sitzungsber. Isis, 6-9.

1882. Die Saurier des unterer Dyas von

Sachsen. Paleontographica, xxix, 9-46, pis. 1-9.

291. Gergens, Dr. 1844. Letter to Professor Bronn

announcing the discovery of the Apatheon de-
scribed by von Meyer as an amphibian. The
first announcement of Amphibia from Carbonifer-
ous. Neues Jahrb. f. Mineral., Geol. u. Paleon., 49.

292. Giebel, C. G. 1847. Fauna der Vorwelt mit steter

Berucksichtigung der lcbenden Thiere. Am-
phibia in Bd. I, Abth. 2, 165-213.

293. . 1854. Odontography, Leipsic.

294. Goldenburg, Fr. 1873. Fauna Saraepontana fos-

silis — Die fossilen Thiere aus der Steinkohlenfor-
mation von Saarbrucken. Heft I, p. 4, Taf. 1,
fig. 1. (Anthracosaurus raniceps.)

295. Goldfuss, Dr. A. 1831. Beitrage zur Kenntniss

verschiedener Reptilien der Vorwelt. Reptilien
aus der schieferigen Braunkohle. Verhandlungen
der k. leop. Carol. Akad. d. Naturf., xv, Abth. 1,
1 17-126, pis. xii-xiii.

-. 1847. Ueber das alteste der mit Bestimmt-

296.

297.

heit erkannten Reptilien . Neues Jahrb. f . Mineral. ,
Geol. u. Paleon., 400, taf. 6. (Archegosaurus.)
1847. Beitrage zur vorweltlichen Fauna

des Steinkohlengebirgen. Bonn, quarto, taf. 1-3.

298. Goodrich, E. S. 191 i. On the segmentation of the

occipital region of the head in the Batrachia
Urodela. Proc. Zool. Soc. London, 101-120, figs.

299. Gregory, W. K. 191 1. The limbs of Eryops and

the origin of paired limbs from fins. Science,
n.s., xxxin, No. 848, 508-509.

1913. Crossopterygian ancestry of the

300.

3000.

Amphibia. Science, n. s., xxxvn, No. 960, 806-
808.

-. 1915. Present status of the problem of

the origin of the Tetrapoda, with special reference
to the skull and paired limbs. Ann. New York
Academy of Science, XXVI, 3I/-383. P'- iv.
(Stegocephali, p. 365.)

301. Gunther, Albert. 1889. On sexual differences

found in bones of some recent and fossil species
of frogs and fishes. Ann. and Mag. Nat. Hist.,
ser. 3, iv, 377.

302. Gurich, G. 1884. Ueber einige Saurier des ober-

schles. Muschelkalkes. Zeit. d. deutsch. geol.
Gesell., xxxvi, 141.

303. Hancock, A. 1869. Note on Anthracosaurus. Ann.

and Mag. Nat. Hist., 4th ser., iv, 270.

304. Hancock and Atthey. 1868. Notes on the remains
of some reptiles and fishes from the shales of the
Northumberland Coal Field. Ann. and Mag.
Nat. Hist., 4th Ser., I, 266-278, pis. xiv-xvi.

3°5- • 1869. On a new Labyrinthodont Amphib-
ian from the Northumberland Coal Field and on
the occurrence in the same locality of Anthraco-
saurus. Ann. and Mag. Nat. Hist., ser. 4, iv,
182-189; also Nat. Hist. Trans. Northumberland
and Durham, ill, 1869, 330. (Urocordyhts
reticulatus.)

. 1870. On Urocordylus, Pteroplax. Nat.

Hist. Trans. Northumberland and Durham, in,

306.
307.
308.

309.

310.
3"-

312-

313.
314

315-
316.

317-

318.
319-
320.

321.

322.

30. 79-

. 1870. On the occurrence of Loxomma all-

manni in the Northumberland Coal Field. Ann.
and Mag. Nat. Hist., ser. 4, v, 374-379.

. 1870. Description of a Labyrinthodont

Amphibian; a new generic form obtained in the
Coal Shales at Newsham. Ann. and Mag. Nat.
Hist., ser. iv, vi, 56-65, pi. i. (Batrachiderpeton
lineatum.)

. 1 87 1. Description of a considerable por-
tion of a mandibular ramus of Anthracosaurus
russelli, with notes on Loxomma and Archichthys.
Ann. and Mag. Nat. Hist., 4th ser., vn, 73, pi. vi.

. 1871. Batrachiderpeton. Nat. Hist. Trans.

Northumberland and Durham, rv, 208.

Hancock and Howse. 1870. On a new Labyrintho-
dont Amphibian from the magnesium limestone
of Midderidge, Durham. Quart. Jonr. Geol.
Soc., xxvi, 556-564, pi. 38. Permian of Eng-
land, Lepidotosaurus diiffii, a Labyrinthodont of
some 3 to 4 feet in length.

Haeckel, Ernst. 1895. Systematische Phylogenie
der Wirbelthiere (Vertebrata). Amphibia, 266-
283. Also History of Creation, Trans., 5th ed.,
1909, 11, 295-300.

Hatch and Corstorphine. 1905. The geology of
South Africa. The Karoo System. 194.

Hatcher, J. B. 1905. The vertebrate fauna of the
Judith River Beds. Bull. U. S. Geol. Surv., No.
257, 67-103. (Scapherpeton and Hemitrypus.)

Hay, O. P. 1890. The skeletal anatomy of Amphi-
uma during its early stages. Jour. Morphol., iv,
1 i_34i with pis.

. 1900. Description of some vertebrates of

Carboniferous Age. Proc. Amer. Phil. Soc.,
xxxix, No. 161, 96-123. (Amphibamus grandi-
ceps.)

. 1902. Catalogue and bibliography of fossil

Vertebrata of North America. Bull. U. S. Geol.
Surv., No. 179, Batrachia, 409-425.

Hickling, G. 1909. British Permian footprints.
Mem. and Proc. Manchester Lit. and Phil. Soc,

53. 3°. Pi- 3-
Hilton, Wm. A. 1902. A structural feature con-
nected with the mating of Diemyctylus virides-

cens. Amer. Nat., xxxvi, 643.
Hoepen, E. C. N. van. 1915. Stegoccphalia of

Senekal. Ann. of the Transvaal Mus., v, No. 2,

125-149, pis. xvi-xxiv.
Hoernes, Rud. 1884. Elcmente der Paleontologic,

445-453- Leipzig.
Hoffman, A. H. 1873-78. Amphibien. Bd. Vi, 2d

Abth., Bronn's Classen und Ordnungen des

Thierreiches.

210

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

323. Huene, Friedrich von. 1902. Uebersicht ueber i
die Reptilien der Trias. Geol. u. Paleon. Ab-
handl., N. F., vi, H. i, 20, 21 , 81.

igio. Neubeschreibung des Permischen

324

325-
326.

327-

Stegocephalen-Dasyceps bucklandi aus Kenil-
worth. Geol. u. Paleon. Abhandl. herausgegeben
v. E. Koken, N. F., vm, H. 6, 33-46, fig. pi. i, ii.

. 1912. Beitrage zur Kenntniss des Schadels

von Eryops. Anat. Anz., Bd. 41, pp. 98-104, figs.
-. 1913. Ueber Lysorophus aus dem Perm

von Texas. Anat. Anz., Bd. 43, 389-396.
— . 19 13. The skull elements of the Permian

Tetrapoda in the American Museum of Natural
History, New York. Bull. Am. Mus. Nat. Hist.,
xxxii, 315-386, 57 figs.
328. Huxley, Thomas Henry. 1859. On Dasyceps
bucklandi (Labyrinthodon bucklandi Lloyd).
Mem. Geol. Surv. United Kingdom, 52-56, figs.
1-2; Scientific Memoirs, 11, 263, figs. 1-2.

1859. On a fragment of a lower jaw of a

329-

33°-

331-

332.

333-

334-

335-

336.

large Labyrinthodont from Cubbington. Mem.
Geol. Surv. United Kingdom, 56-57; Scientific
Memoirs, 11, 269.

1859. On some Amphibian and reptilian

remains from South Africa and Australia. Quart.
Jour. Geol. Soc., xv, 642-649, pis. xxi-xxii;
Scientific Memoirs, 11, 120-129, p's. 7-8.
. 1862. On new Labvrinthodonts from the

Edinburgh Coal Field. Quart. Jour. Geol. Soc.,
xvm, 291-296, pi. xi; Scientific Memoirs, 11,

53""535. Pi- 37-
. 1863. Description of Anthracosaurus rus-

selli, a new Labyrinthodont from the Lanark-
shire Coal-Field. Quart. Jour. Geol. Soc, xix,
56-68, figs.; Scientific Memoirs, 11, 558-572, figs.
1-2.

. 1865. On a collection of vertebrate fossils

from the Panchet Rocks, Ranigunj. Bengal.
Mem. Geol. Surv. of India; Paleontologia Indica,
ser. iv; Indian Pretertiary Vertebrata, t, Am-
phibia. 3-8, pis. vi, and figs.; Scientific Memoirs,
Hi, 90-120, figs., pi. 16.

1867. On a collection of fossil Vertebrata

337-

from the Jarrow Colliery, County of Kilkenny,
Ireland. Trans. Roy. Irish Acad., 1871, xxiv,
351-369 (Read Jan. 6, 1866), pis. xix-xxiii; Scien-
tific Memoirs, in, 180-197, pis. 17-21.

. 1869. On a new Labyrinthodont from

Bradford. Quart. Jour. Geol. Soc. London, xxv,
309-311, pi. xi; Scientific Memoirs, III, 391-393,
pi. 26.

. 1875. Amphibia. Art. in "Encyclopedia

Britannica," 9th ed., I, 750-771.

. 1879. On the characters of the pelvis in

the Mammalia and the conclusions respecting the
origin of mammals which may be based on them.
Proc. Roy. Soc. Lond., xxvm, 395-405, figs. ;
Scientific Memoirs, iv, 345-356, figs.
338. Jaeger, Dr. Georg Friedrick. 1824. De Ichthyo-
sauri sive Proteosauri Fossilis Specimenibus in
Agro Bollensi in Wurtembergia. Folio, 11. (First
mention of the discovery of Labyrinthodonts, a
tooth and occipital region of a cranium, which in
1 833 Jaeger designated Mastodonsaurus giganteus.
This was and still is the largest known amphibian.)

339- — ; • 1828. Ueber die fossilen Reptilien welche

im Wurtemburg aufgefunden worden sind.
Quarto, 35-38, pis. iv-v.

340. Jaeger, Dr. G. F. 1833. Mastodonsaurus gigan-
teus. Bull, de la Soc. Geol. de France, HI, 86.

341. . 1850. Ueber die Uebereinstimmung des

Pygopterus lucius Agassiz mit dem Arehego-
saurus decheni. Abh. Math-phys. Classe d. Bayr.
Akad., v, 877, pi.

342. Jaekel, Otto. 1894. Ueber die sogennante Falten-

zahne und complicirte Zahnbildung ueberhaupt.
Sitzb. d. Gesell. naturf. Freunde zu Berlin, No. 5,
146.

343. . 1896. Ueber die Korperform und Haut

Bedeckung von Stegocephalen. Sitzungsber. d.
Gesell. naturf. Freunde zu Berlin, 1-8, 1 fig.

344. . 1896. Die Organization von Archegosaurus.

Zeit. d. deutsch. geol. Gesell., xlviii, 505-521,

figs. 1-10.
345. . 1902. Gephyrostegus bohemicus. Zeit. d.

346.

347-

348.

349-

35o.

351-

deutsch. geol. Gesell., liv, Heft 3, 127-132.
. 1903. Ueber die Epiphyse und Hypophyse.

Sitzungsb. d. Gesell. naturf. Freunde, No. 2,

27-58.
. 1903. Ueber Ceraterpeton, Diceratosaurus

und Diplocaulus. Neues Jahrb. f. Mineral., Geol.

u. Paleon., Bd. I, 109-134, pis. ii-v.
. 1904. Ueber die Bildung der ersten Hals-

wirbel und die Wirbelbildung im Allgemeinen.

Zeit. d. deutsch. geol. Gesell., lvi, 109-119, figs.
. 1905. Ueber die primare Gliederung des

Unterkiefers. Sitzungsber. d. Gesell. naturf.

Freunde, No. 4, 134.
. 1909. Ueber die Klassen der Tetrapoden.

Zool. Anz., xxxiv, Nr. 7/8, Apr. 20, 193-212,

figs. 1-15.

1909. Ueber die Beurtheilung der paarigen

352-

Extremitaten. Sitzungsb. d. konig. Preuss. Akad.
d. Wissen., xxvi, 707-724.

. 1913. Ueber den Bau des Schaedels. Anat.

Anz.. Erganzungsheft, Bd. 44, 77-94. mit figuren.

353. Jordan. H. 1849. Erganzende Bemerkungen zu der

Abhandlung von Goldfuss ueber die Gattung
Archegosaurus. Verhandl. d. naturhist. Vereines
f. Rheinlande u. Westphalen, vi, 76-81, taf. 4.

354. Kayser, Friedrich. 1902. Geologie von Boehmen.

Verlag von I. Taussig, 1074-1210. (Formations
of Bohemia where Amphibia have been found.)

355. Kinahan, G. H. 1878. Geology of Ireland, London;

The Jarrow Colliery. 116-119.

356. King, Alired T. 1844. Description of fossil foot-

marks supposed to be referable to the classes
Birds, Reptilia. and Mammalia found in the
Carboniferous series in Westmoreland County,
Pennsylvania. Proc. Acad. Nat. Sci., Phila.,

175-179-

357. . 1845. Description of a fossil footmark of

the quadruped Thenaropus. Amer. Jour. Sci.,
xlviii, 343-352, 12 figs.; also 1 (n. s.), 268.

358. Kingsley, J. S. Systematic position of Coecilians.

Tufts College Studies No. 7 (xxx); also No. 7
(xxvm), Amphiuma.

359. Ki.aatsch, Hermann. 1896. Die Brustflosse der

Crossopterygier. Festschrift fur Gegenbaur, Bd.
1, 259-391, Taf. i-iv.

360. Knipe, Henry R. 1912. Amphibians of the Car-

boniferous. Evolution in the Past, 59, and plate.

361. Kunisch, Hermann. 1885. Ueber die Unterkiefer von

Mastodonsaurus silesiacus Hun. Zeit. d. deutsch.
geol. Gesell., xxxvn, 528-533, with figs. 1-2.

BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA.

211

362. Kunisch, Hermann. 1890. Labyrimhodonten,

Reste d. oberschles. Musehelkalkes. Zeit d.
deutsch. geol. GcmD., xi.ii, 377-385. Taf. xx.

363. Kutorga, Stefh. 1838. Bcitrag zur Kenntniss der

organ. Ueberreste des Kupfersandsteines am west-
lichen Abhange des Urals. St. Petersburg, rait
tafeln.

364. Lake, J. J. 1878. Reptiles of the primary period.

English Mechanic, xxvm, 438-439.

365. Lambe, L. M. 1902. Scapherpeton tectum — a

batrachian from the Cretaceous. Contr. Cana-
dian Paleontology, ill, pt. 11, 31.

366. . 1904. Progress of vertebrate paleontology

in Canada. Trans. Roy. Soc. Canada, x, sec. iv,

I3-56-

367. Laube, Gustav C. 1898. Andriasreste aus der

Boehmischen Braunkohlenformation. Abhandl.
d. deutsch. naturw.-med. Vereines f. Boehmen
"Lotos," 1, heft 2, 25.

1898. Amphibienreste aus dem Diatoma-

368.

369-

ceenschiefer von Sulloditz. Abhandl. d. deutsch.
naturw.-med. Vereines f. Boehmen "Lotos," 1,
heft 3, 10, taf. i, fig. 6.

1 901. Synopsis der Wirbelthierfauna der

bohm-Braunlcohlenformation und Beschreibung
neuer oder bisher unvollstandig bekannter Arten.
Abhandl. d. deutsch. naturw.-med. Vereines f.
Boehmen "Lotos," 11, heft 4, 52-58.

370. ■ 1909. Neue Andriasreste aus dem Tonen

von Preschen bei Bilin. "Lotos," lvii, 6.

371. Lea, Isaac. 1849. On traces of a fossil reptile

(Sauropus primaevus) found in the old Red
Sandstone (Pennsylvania). Rep. Brit. Assn. Adv.
Sci., 56, 2d part; Proc. Amer. Phil. Soc, 91-94.

372. . 1855. Fossil footmarks in the Red Sand-
stone of Pottsville. 1-16, with a large plate.
Large folio; Trans. Amer. Phil. Soc, x, 1852, Apr.

1-13. P's. 31-33-

373. Leidy, Joseph. 1856. Notices of remains of extinct

vertebrated animals discovered by Professor E.
Emmons. Proc. Acad. Nat. Sci. Phila., 256.
(Dictyocephalus elegans.)

374. . 1878. Fossil foot-tracks of the Anthracite

Coal Measures. Proc. Acad. Nat. Sci. Phila.,
xxxi, 164-165.

375. Le Roy, J. J. 1874. Notiz ueber Archegosaurus

decheni Goldfuss und A. latirostris Jordan.
Niederl. Archiv. f. Zoologie. n, 89-98.

376. Lesley, J. P. 1889. A dictionary of the fossils of

Pennsylvania and neighboring states. In three
volumes. Published by the Penn. Geol. Sun'.

377. Leunis, Johannes. 1883. Synopsis der Thierkunde.

3d Auflage, Bd. I, 599-631. (Amphibia.)

378. Lillie, Frank R. 1908. The development of the

chick. The Vertebrce, 412-424, figs. 246, 249,
250.

379. Lloyd, G. 1849. On a new species of Labyrintho-

don from the New Red Sandstone of Warwick-
shire. Rept. Brit. Assn. Adv. Science, 56. (See
Huxley (328) and F. von Huene (324).)

380. Logan, William. 1842. Proc. Geol. Soc, London

hi, pt. 11, 707.

381. Lohest, Maximin. 1888. Decouverte de plus an-

cien Amphibien connu et de quelques Fossiles
remarquables de Famennien supeneur de Modave.
Notice preliminaire. Ann. de la Soc Geol.
Belgique, XV, p. cxx, fig. r.

382.
383.

384.
385.

386.

387.
388.

389.

390

391-
392-

393-

394-

395-

396.

397-

398-
399-

400.

Lortet, Dr. L 1892. Les Reptiles fossiles du Bassin
du Rhone. Ordre des Amphibiens, 125-127, pi.
xii; Archives du Museum d'Hist. Natur. de
Lyon, v.

Lucas, F. A. 1904. A new Batrachian and a new
reptile from the Trias of Arizona. Proc. U. S-
Nat. Mus., xxvii, 193-195, pi. (Heterodonto-
suchus, a belodont; Metoposaurus fraasii, a
labyrinthodont.)

I. ydkkkkr, Richard. 1879. Reptilia and Batrachia
(Amphibia). Paleontologia Fndica, ser. iv, 1,
1-36, pis. i-vi; Amphibia, 17-20, pis. iii-vi.
(Pachygonia, Gonioglyptus, Mastodonsaurus.)

. 1880. A sketch of the history of the fossil

Vertebrataof India. Jour. Asiatic Society Bengal,
xlix, pt. 11, 8-40; Fossil Batrachians, 16, list of
species, 38-39.

1 88 1. Note on some Gondwana verte-

brates. Rec Geol. Surv. India, xiv, 174. (Gonio-
glyptus, Pachygonia, Glyptognathus.)

-. 1883. Note on the Bijori Labyrinthodont.

Rec. Geol. Surv. India, xvi, 93.

1883. Synopsis of the fossil Vertebrata of

India. Rec. Geol. Surv. India, xvi, 61 ; Amphibia,
64, 88.

1885. The Reptilia and Amphibia of the

Maleri and Denwa Groups. Paleontologia Indica,
ser. iv, 1, pt. 5, 6 pis.

-. 1885. The Bijori Labyrinthodont. Paleon-

tologia Indica, ser. iv, I, 1-14, pt. 4; also, Cope,
Amer. Nat., xix, 592.

-. 1887. The fossil Vertebrata of India. Rec.

Geol. Surv. India, xx, p. 68.

-. 1887. Dr. Hermann Credner on the de-

velopment of Branchiosaurus. Geol. Mag., n. s.,
Dec ill, iv, 276-278.

-. 1890. Catalogue of fossil Amphibia and

Reptilia in the British Museum (Natural History).
Amphibia, part iv, 121-225.

1891. On a Labyrinthodont skull from the

Kilkenny Coal Measures (Ireland). Quart. Jour.
Geol. Soc, xlvii, 343-347. 1 fig-

. The new natural history, v, Amphibia,

257-310; Labyrinthodontia, 311. (See also
Nicholson and Lydekker (499).)

Lyell, Chas., and J. W. Dawson. 1853. On the
remains of the reptile (Dendrerpeton acadianum)
and a land shell (Coal Measures). Quart. Jour.
Geol. Soc, ix, 58-63, with pis.

Maggi, Leopoldo. 1898. Placche osteodermiche
interparietali Stegocephali e rispondenti Centri
Ossificazione interparietali dell 'Uomo. Reale
Inst. Lombard, di Sci. Lett. (2), xxxi, 3X1, 228,
pi. 1.

. 1897. Resultats de Recherches morpho-

logiques sur des Os et des Fontanelles du Crane
humain. Arch. Ital. d. Biol., T. 27, 230-238.

Makowsky, Alex. 1876. Ueber einen neuen Laby-
rimhodonten — Archegosaurus austricus. Sitz-
ungsb. d. lb Akad. wiss. Wien. Math. Xatur.
Classe, Abth. 1. lxxiii, Heft 3, 155-166.

und A. Rzehak. 1884. Die geologischen

Verhaltnisse der Umgebung von Brunen. Ver-
handlgn. d. Nat. Verein zu Brunn, xxn. (Die
fruhere Archegosaurus austricus wird zu Melaner-
peton gestellt.)

212

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

401. Malbranc, M. 1875. Von der Seitenlinie und ihre

Sinnesorganen bei Amphibien. Leipzig. Zeit.
f. wiss. Zool., xxvi, 24, pis. 1-4.

402. Mantell, G. A. 1 85 1. Petrifactions and their

teachings. Fossil Batrachians, 183.

403. . 1866. Wonders of geology. Labyrintho-

donts, 551.

404. Marsh, O. C. 1862. Discovery of new Enalio-

saurian remains in the Coal Formation of Nova
Scotia. On the saurian vertebra? from Nova
Scotia. Amer. Jour. Sci. and Arts, xxxiv, Mar. I,
pi. 1, figs. 1-2. See also Neues Jahrb. Mineral.,
Geol. u. Paleon., 1863, 247; Canad. Nat. and
Geol., vii, 205-213; Quart. Jour. Geol. Soc.
London, xix, 52-56; Huxley, Scientific Memoirs,
n, 566.

405. . 1887. Eobatrachus agilis in the (Como

Beds) Jura of North America. Amer. Jour. Sci.,
xxxm, 328; Vertebrate fossils of the Denver
Basin, Monograph U. S. Geol. Surv., xxvii, 508,
1897.

406. . 1894. Footprints of vertebrates in the Coal

Measures of Kansas. Amer. Jour. Sci., xlviii,
2d ser., July-Dec, 81-84, pis. ii-iii.

407. . 1896. Amphibian footprints from the De-
vonian. Amer. Jour. Sci. (4), 11, 374-375, with
fig-

408. Matthew, G. F. 1903. New genera of Batrachian

footprints of the Carboniferous system of eastern
Canada. Canadian Record of Science, ix, No. 2,
99-1 11, with figs.

409- • I903- An attempt to classify Paleozoic

Batrachian footprints. Trans. Roy. Soc. Canada
(2), ix (m), 1903, 109-121.

4io- • I9°3- On Batrachian and other footprints

from the Coal Measures of Joggins, N. S. Bull.
Nat. Hist. New Brunswick, v, No. xxi, 103-108,
with plate of excellent photos; No. xxn, 247.

4ii- • I905- Note in reference to Batrachian

footprints. Bull. Soc. New Brunswick, xxi, 102.

412- • '9°5- Note on the genus Hylopus of

Dawson. Bull. Soc. New Brunswick, xxn, 247-
252, figs.

4'3- • 1904- New species and a new genus of

Batrachian footprints of the Carboniferous sys-
tem in Eastern Canada. Trans. Roy. Soc.
Canada, x (iv), 77-110, pis. 1-6.

414. Matthew, W. D. 1909. The oldest land reptiles of
North America. Amer. Mus. Jour., ix, No. 4,
91-95. photos of Eryops and Diplocaulus.

4'5- • 1911- The Amphibians of the great Coal

Swamp. Amer. Mus. Jour., xi, No. 6, 197-200.
(Restoration of Eryops.)

416. Merrill, Geo. P. 1907. Catalogue of the type
and figured specimens of fossils, minerals, rocks
and ores. Part II: Fossil Vertebrates, 9-12'
Bull. U. S. Nat. Mus., No. 53, pt. 11. Two species
of Amphibia are listed as fishes, 74, 77

417- Metcalfe, A. T. 1884. On further discoveries of
vertebrate remains in the Triassic strata of the
South Coast of Devonshire, between Budleigh
Salterton and Lidmouth. Quart. Jour. Geol'
Soc, XL, 257.

418. Meyer, Hermann von. 1838. Recherches sur les
Ossem. foss. du gres bigarre de Soultz les Bains.

4 ''■ • l834- Odontosaurus voltzii. Mem. de la

Soc. d'hist. nat. de Strasbourg.

420. Meyer, Hermann von. 1842. Labyrinthodontia.

Neues Jahrb. f. Mineral., Geol. u. Paleon., 301-
304. (Proposes system of classification.)

421. . 1844. Beitrage zur Paleontologie Wurtem-

bergs, 1-41, mit 12 tafeln. Enthaltend die fos-
silen Wirbelthierereste aus den Triasgebilden mit
besonderer Rucksicht auf die Labyrinthodonten
des Keupers.

422. . 1844. Apatheon pedestris. Neues Jahrb.

f. Mineral., Geol. u. Paleon., 336. (He regarded
Apatheon as a salamander-like form, a Branchio-
saurian.)

423- • 1845. System der fossilen Saurier. Neues

Jahrb. f. Mineral., Geol. u. Paleon., 278-285.

424- • 1847. Die Steinbrucke von Oeningen. Fossile

Batrachier. Fauna der Vorwelt, Bd. 1, 18-40,
pis. 9-10.

425- • 1847-1855. Die Saurier des Muschelkalkes

mit Rucksicht auf die Saurier aus Bunten Sand-
stein und Keuper. Fauna der Vorwelt, Bd. 11,
1-164, pis. 58-64. Large folio. (Mastodon-
saurus, Metopias, Capitosaurus.)

426. . 1848. Apateon pedestris aus der Stein-

kohlenformation von Mitnsterappel. Paleonto-
graphica, Bd. 1, 153-154, pi. xx, fig. 1.

427- • 1848. Mittheilung an Professor Bronn.

Labyrinthodontia. Neues. Jahrb. f. Mineral.,
Geol. u. Paleon., 469. (Labyrinthodon ocella.)

428. — . 1849. Ueber den Archegosaurus der Stein-

kohlenformation. Paleontographica, Bd. 1, 209,
Dec. (1851), pi. 33.

429- . 1851. Mittheilung an Professor Bronn,

Rana jaegeri. Neues Jahrb. f. Mineral., Geol. u.
Paleon., 78.

43°- ' ■ 1 85 1. Beschreibung der fossilen Deca-

poden, Fische, Batrachier, und Saugethiere aus
dem Tertiaren Susswassergebilden des nordlichen
Boehmens. Paleontographica, Bd. 11, 43.

43 •■ • 1852. Mittheilungen an Professor Bronn.

Neues Jahrb. f. Mineral, Geol. u. Paleon.: Pateo-
batrachus gigas, 465; Rana noggerathi, 58; R.
troscheli, 466; R. salzhauserensis, 467.

432. . 1853. Mittheilungen an Professor Bronn.

Neues Jahrb. f. Mineral., Geol. u. Paleon.: Rana
meriani, 163, 185; Pakeobatrachus gigas, 162.

433- • 1854. Mittheilung ueber Archegosaurus.

Neues Jahrb. f. Mineral., Geol. u. Paleon., 424.

434- • 1856. Ueber Osteophone roemeri, Briefl.

Mittheil. an Professor Bronn. Neues Jahrb. f.
Mineral., Geol. u. Paleon., 824.

435- • 1857- Mittheilung an Professor Bronn.

Neues Jahrb. f. Mineral., Geol. u. Paleon.: Pakeo-
batrachus gracilis, 555.

436. ■ — . 1857. Reptilien aus der Steinkohlenforma-

tion in Deutschland. Paleontographica, Bd. vi,
219-220, text-fig. (Archegosaurus.)

437- ■ 1857. Osteophorus roemeri, Beschreibung

derselben in Roemer: Ueber Fisch- und Pflanzen-
fuhrende Mergelschiefer des Rothliegenden bei
Neundorf. Zeit. d. deutsch. geol. Gesell., ix, 61.

438. . 1858. Reptilien aus der Kohlsteinforma-

tion in Deutschland. Cassel, Folio; also Pale-
ontographica, Bd. VI, 59-218, Tafeln viii-xxiii.
(Archegosaurus.)

439- • 1858. Labyrinthodonten aus dem bunten

Sandstein von Bernburg. Paleontographica, Bd.
vi. 221-245, Tafeln xxiv-xxvii.

BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA.

213

440. Mkyer, Hermann von. 1858. Mittheilung an
Professor Bronn. Neues Jahrb. f. Mineral. Geol.
u. Paleon.: Rana danubina, 203.

44'- ' — ■ 1859- Mittheilung an Professor Bronn.

Neues Jahrb. f. Mineral. Geol. u. Paleon.: Triton
basal ticus, 430; Andrias tschudii, 723.

442. ■ i860. Ueber Archegosaurus. Erwiderung

auf die Ausfuhrungen von Quenstedt in " Epochen
der Natur," p. 410. Brierl. Mittheil. an Professor
Bronn. Neues Jahrb. f. Mineral. Geol. u. Paleon.,
71-72.

i860. Osteophorus roemeri aus dem Roth-

443

444

445

446

447

448

449

450

45i

452

453

454-

liegenden von klein Neundorf in Schliesen.

Paleontographica, Bd. Vll, 99-104, pi. xi.
. i860. Frocsche aus Tertiar Gebilden

Deutschlands. Paleontographica, Bd. vn, 123-

182, pis. 16-22.
. i860. Melosaurus uralensis aus der Per-

mische System des westlichen Urals. Paleonto-
graphica, Bd. vh, 90-98, pi. x.
. i860. Salamandrinen aus der Braunkohle

am Rhein und in Boehmen. Paleontographica,

Bd. vn, 47-73, pis. viii, ix.
. 1863. Heliarchon furcillatus, ein Batra-

chier aus der Braunkohle von Rott in Siebenge-

birge. Paleontographica, Bd. x, 292-298, Taf.

50, figs. 5-6.

1866. Reptilien aus dem Kupfersandstein

Russlands. Paleontographica, Bd. xv, 97-130.
Miai.l, L. C. 1873. Report on the Labyrinthodonts

of the Coal Measures. Rept. Brit. Assn. Adv.

Sci., 225, pis. 1-3.
. 1874. On the classification of the Lahy-

rinthodontia. Rept. Brit. Assn. Adv. Sci., 149-

192, with 2 plates.

1874. The classification of the Labyrintho-

donts. Geol. Mag., 2d ser., 1, 513.

1874. On the occurrence of a Labyrintho-

dont in the Yoredale Rocks of Wensleydale.
Quart. Jour. Geol. Soc, xxx, 775.

-. 1874. On the remains of Labyrinthodontia

from the Keupcr Sandstone of Warwick, pre-
served in the Warwick Museum. Quart. Jour.
Geol. Soc, xxx, 417-435, pis. 26-28. (Masto-
donsaurus pachygnathus; Labyrinthodon lepto-
gnathus; Diadetognathus varvicensis.)

1875. Sur les Labyrinthodontes du Ter-

rain houiller. Deuxieme partie. Jour, de Zool.,
iv, I9-37-

455. Miller, S. A. 1889. North American geology and

paleontology. Amphibia, 614.

456. Mivart, Geo. 1870. On the axial skeleton of the

Urodela. Proc. Zool. Soc. London, 260-278.

457. Moodie, Roy L. 1908. Dissorophus — A correction.

Science, n.s., xxvn, No. 679, 30.

-. 1908. The lateral line system in extinct

458.

459-

460.

461.

Amphibia. Jour. Morphol., xix, No. 2, 511-540,
17 figs.
. 1908. The ancestry of the caudate Am-
phibia. Amer. Nat., 42, 361-373, figs. 1-10.
-. 1908. The dawn of quadrupeds in North

America. Pop. Sci. Monthly, lxxii, June, 558-
566, 5 figs.

1908. The clasping organs of extinct and

recent Amphibia. Biol. Bull., xiv, No. 4, Mar.,
249-259, with 5 figs.

462.
463.

464.
465-

466.
467.

468.
469.

470.
47i-

472.
473-

474-

475-

476.

477-

478.

479-

480.
481.
482.

483-
484.

Moodie, Roy L. 1909. New forms (extinct Amphi-
bia) from the Carboniferous. Jour. Geol., xvn,
No. 1, Jan.-Peb., 38-82, 24 figs.

. 1909. The Carboniferous quadrupeds of

Kansas, Ohio, Illinois, and Pennsylvania in their
relation to the classification of the so-called
Amphibia and Stegocephalia. Trans. Kans.
Acad. Sci., xxii, 239-243, pis. 1-3.

. 1909. Carboniferous air-breathing verte-
brates of the United States National Museum.
Proc. U. S. Nat. Mus., 37, ri-28, pis. 4-10.

1909. New or little-known forms of Car-

boniferous Amphibia in the American Museum

of Natural History. Bull. Amer. Mus. Nat.

Hist., xxvi, art. xxv, 347-357, pis. lviii-lxv.
. 1909. The Lysorophidie. Amer. Nat.,

XLiii, 116. (Review.)
. 1909. The Stegocephala. Amer. Nat.,

XLiii, 119. (A review of paper by Hugo

Schwarz.)

-. 1909. The morphology of the vertebrate

sacral rib. Anat. Anz., xxxiv, No. 15, 361-364.
-. 1909. The Microsauria, ancestors of the

Reptilia. Geol. Mag., Dec. v, vi, No. 539, 216-220,
1 fig.

1910. The Amphibia of the Mazon Creek

Shales. Science, n. s., xxxi, No. 789, p. 233.

1910. The alimentary canal of a Carbonif-

erous salamander. Amer. Nat., XLIV, No. 522,

367-375. 4 figs-

-. 1910. A new Labyrinthodont from Kansas.

Science, n. s.. xxxii, No. 829, Nov. 18.

-. 191 1. A new Labyrinthodont from the

Kansas Coal Measures. Proc. U. S. Nat. Mus.,

xxxix, 489-495, 4 figs.
. 191 1. Two Amphibians, one of them new,

from the Carboniferous of Illinois. Proc. U. S.

Nat. Mus., XL, 429-433, figs. 1-2.
-. 191 1. Recent contributions to a knowledge

of the extinct Amphibia. Amer. Nat., xi.v, June,

375-384-

-. 191 1. The temnospondylous Amphibia,

and a new species of Eryops from the Permian of
Oklahoma. Kans. Univ. Sci. Bull., v, No. 13,
235-253. pis. xlix-liv.

-. 1912. The skull structure of Diplocaulus

magnicornis and the amphibian order, Diplo-
caulia. Jour. Morphol., xxm, No. 1, 31-39, pis.
1-2.

-. 1912. The Pennsylvanic Amphibia of the

Mazon Creek, Illinois, Shales. Kans. Univ. Sci.
Bull., vi, No. 2, 323-359. Pis. 1-14-

-. 1912. The Mazon Creek, Illinois, Shales

and their amphibian fauna. Amer. Jour. Sci.,
xxxiv, Sept., 277-285, figs. 1-2.

1912. An American Jurassic frog. Amer.

Jour. Sci., xxxiv, Sept., 286-288.

1914. The fossil frogs of North America.

Amer. Jour. Sci., xxxvm, Dec, 53i-53<>. figs- 1-2.
-. 1914. A list of the described species of

fossil Amphibia. Kans. Univ. Sci. Bull., 9, No. 2,
13-28.

1915. A remarkable Microsaur from the

Coal Measures of Ohio. Science, n. s., xli. No.
1044, 34-35-

. 1915. Coal Measures Amphibian with an

osseous tarsus. Amer. Jour. Sci., xxxix. May,
509-512, figs. 1, 2.

214

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

485. Moodie.RoyL. igij.ThescaledAmphibiaoftheCoal

Measures. Science, n. s., xli, No. 1056, 463-464.

486. — . 19 1 5. Some recent studies on fossil

Amphibia. Amer. Nat., xux, June, 369-376.

487- • 1915- A new fish brain from the Coal

Measures of Kansas, with a review of other fossil
brains. Jour. Comp. Neurol., xxv, No. 2, 164.

488. . 1915. A further contribution to a knowledge

of the lateral line system in extinct Amphibia. Jour.
Comp. Neurol., xxv, No. 4, 317-328, figs. 1-9.

. 1915. Some methods of studying fossils

embedded in coal. Kans. Univ. Sci. Bull., IX,
No. 16, 187-193, figs. 1-3, pi. l.

. The migrations and geographic distribu-
tion of the fossil Amphibia. Amer. Jour. Sci.,
xl, 186-190, 1 map.

■ The Coal Measures Amphibia and the

Crossopterygia. Amer. Nat., xlix, 637-644.

489.

489a.

4896.

490. Mudge, B.F. 1873. Recent discoveries of fossil foot-

prints in Kansas. Trans. Kans. Acad. Sci. , 11, 7-9.

491. Munster, Georg, Graf zu. 1834. Vorlaufige

Nachricht ueber einige neuen Reptilien im
Muschelkalk von Baiern. Neues Jahrb. f. Min-
eral., Geol. u. Paleon., 527. (Mastodonsaurus.)

492- • 1836. Capitosaurus arenaceus. Neues

Jahrb. f. Mineral., Geol. u. Paleon., 580.

493- • 1839- Mastodonsaurus adriani. Beitrage

zur Petrefactionkunde, 102.

494. Murchison, Roderick I. 1872. Siluria— A history

of the oldest rocks in the British Isles and other
countries. 5th ed., 302, 343.

495. Newberry, J. S. 1856. Description of several new

genera and species of fossil fishes from the Car-
boniferous strata of Ohio. Proc. Acad. Nat. Sci.
Phila., viii, 98.

496- — ■ • 1867. On some fossil reptiles and fishes

from the coal strata of Ohio, Kentucky, and
Illinois. Proc. Amer. Assn. Adv. Sci.; 144-146.

497- • 1878. Geology of Jefferson County, Ohio.

Geol. Surv. Ohio, in, 736-737.

498- • 1889. Palaeozoic fishes of North America.

Monograph U. S. Geol. Surv., xvi, 228. (Dis-
cusses Kammplatten; Amphibia of Linton, Ohio,
211-214; Fauna of Mazon Creek, 214. See also
Hodge, 1878, Geol. Surv. Ohio, in, 592.)

499. Nicholson and Lydekker. 1889. Manual of pale-

ontology. Amphibia, II, 1018-1045.

500. Oldham, Dr. R. D. 1875. On a salamandriform

Labyrinthodont from the Coal Measures near
Castlecomer, Ireland. Rept. Rugby School Nat.
Hist. Soc, 74-76, pi. vii.

501. — - — . 1893. Geology of India. 2d ed. The

Gondwana System, 149.

502. Owen, Sir Richard. 1 840-1 845. Batrachians.

Odontography, 187-218, pis. 62, 63, 64.

503- • 1842. On the teeth of the species of the

genus Labyrinthodon of Jaeger from the German
Keuper and the sandstone of Warwick and Leam-
ington. Ann. and Mag. Nat. Hist., vm, 58; also,
Geol. Trans. 11, ser. vi, 503.

504- • 1842. Description of parts of the skeleton

and teeth of the genus Labyrinthodon. with re-
marks on Cheirotherium. Trans. Geol. Soc.
London, vi, pt. 2, 515-543, pis. xliii-xlvii.

5»5- • 1842. On the teeth of species of the genus

Labyrinthodon. Trans. Geol. Soc. London, vi,
pt. 2, 503-513, pi. 43, fig. j, (Discusses validity
of term Mastodonsaurus. Read Jan. 11, 1841.)

506

507
508.

5°9-
510.

5"-
512.

513-
514-

. Owen, Sir Richard. 1847. On the Batracholites in-
dicative of a small species of frog (Rana pusilla).
Quart. Jour. Geol. Soc., m, 224. (See Clark, 103,
p. 221.)

. 1853. Notice of a batrachoid fossil in the

British Coal Shale. Quart. Jour. Geol. Soc.
London, ix, 67-70, pi. iii.

-. 1853. Notice of a batrachoid fossil — Para-

515-

516.
517-

518-

519-
520.

batrachus colei — in the British Coal Shale. Quart.
Jour. Geol. Soc. London, x, 207.

• 1853. Baphetes planiceps — a fossil em-
bedded in a mass of coal from Nova Scotia.
Quart. Jour. Geol. Soc. London, x, 207-208, pi. ix.

. 1854. Description of the cranium of a

Labyrinthodont reptile (Brachyops) from the
Triassic of Central India. Quart. Jour. Geol.
Soc. London, x, 473. (Abstract.)

• 1855. Description of the cranium of a

Labyrinthodont reptile from Mangala, Central
India. Quart. Jour. Geol. Soc. London, xi, 37, pi.

. 1859. On the orders of fossil and recent

Reptilia and their distribution in time. Rept.
Brit. Assn. Adv. Sci., 153-166.

. 1861. Paleontology, 2d ed., Amphibia, 193.

. 1862. Descriptions of specimens of fossil

Reptilia discovered in the Coal Measures of the
South Joggins, Nova Scotia. Quart. Jour. Geol.
Soc. London, xvm, 238-244, pis. 9-10.

-. 1865. Description of some remains of an

522,
523.
524-

525-
526.

527-

air-breathing vertebrate (Anthrakerpeton cras-

sosteum) from the Coal Shale of Glamorganshire.

Geol. Mag., 11, 6-8, pis. i-ii.
. 1866. On the anatomy of the Vertebrata.

Amphibia, I, 8.
■ 1876. On Petrophryne granulata Ow., a

Labyrinthodont reptile from the Trias of South

Africa. Bull, de la Soc. Imp. de Nat. de Moscou.

(Review.)
. 1876. Catalogue of the fossil Reptilia of

South Africa in the collection of the British

Museum. Labyrinthodontia, 67-70, pi. xx, figs.

13-20; pi. lxx, fig. 3.
. 1884. On a Labyrinthodont Amphibian

from the Cape of Good Hope. Quart. Jour. Geol.

Soc. London, XL, 333, 2 pis.
. 1884. On Rhytidosteus capensis Ow., a

Labyrinthodont from the Trias of Cape of Good

Hope. Ann. and Mag. Nat. Hist. (5), xin, 481.
Pabst, Wm. 1896-1905. Beitrage zur Kenntniss der

Thierfahrten in dem Rothliegenden Deutschlands.

Zeit. der deutsch. geol. Gesell., 1896, xlviii, 638,

808; 1900, lii, 48; lvii, 1 -14, 361-379, with pis.

(Some of the tracks described may be amphibian.)
Pander, Chr. H. 1858. Ueber die Ctenodipterinen

des devonischen Systems.
Pictet, F. J. 1853. Traite de paleontologie, 2d ed.,

tome premier.
Poli.ard.H.B. 1891. On the anatomy and phylogen-

etic position of Polypterus. Anat. Anz. , VL338 ; Zool.

Jahrb.(Morphol.Abth.),v, 1893, 387-428', with pis.
Portis, A. 1885. Resti di Batraci fossili Italiani:

appunti paleontologi, pt. 2, p. 30.
Pusch, Bergrath. 1842. Fossile Batrachier und

Ophidier Rcste aus Podolien. Neues Jahrb. f.

Mineral., Geol. u. Paleon., 179-180.
Quenstedt, F. A. von. 1850. Die Mastodonsaurier
im gruenen Keupersandstein Wiirtembergs sind
Batrachier, 4to, 1-34, 4 Tafeln.

BIBLIOGRAPHY OF THE FOSSIL AMPHIBIA.

31!

528. Qualen, Wangenheim von. 1S45 and 1852. Two

papers on the skull of Zygosaurus lucius. Bull.
Soc. Imp. Nat. Moscou, xvm, 389 (1852),
47-!-

529. . 1 861. Bemcrkungen zum Archegosaurus.

Neues Jahrb. f. Mineral. Geol. u. Paleon., 294-300,
with pi. iii. (See also Baur, 1886, Amer. Nat., 173,
and Zool. Anz., 1886, Nr. 216.)

53«- • 1885. Atlas zum Handbuch der Petrifac-

tionskunde. Amphibia, pi. 18.

531. Raymond, Percy E. 1907. On the discovery of

reptilian remains in the Pennsylvanian near
Pittsburgh, Pa. Science, n. s., xxvi, 676, 835.

532. Rjabinin, M. A. 1911. Debris de Stegocephales

trouves aux Mines de Kargala Gouvernement
d'Orembourg. Bull, du Comm. Geol. St. Peters-
burg, xxx, No. 185, pis. 1-2.

533. . 1912. Ueber die Ueberreste der Stego-

cephalen aus den Kargalinokischen Bergwerken
des Gouvernments Orenberg, St. Petersburg,
37 pages, pis. 1-2.

534. Roemer, Ferdinand. 1857. Ueber Fische und Pfian-

zenfiihrende Mergelschiefer des Rothliegenden
bei Klein-Neundorf und ueber Acanthodes
gracilis. Zeit. d. deutsch. geol. Gesell., ix, 61.

535. Scheuchzer, J. J. 1726. Part of two human skele-

tons petrified. Phil. Trans. Roy. Soc. I-ondon,
1726, abridged edition, vn (1724-1734), 129.
(Translated from the Latin of a letter to Sir Hans
Sloan, a member of the Royal Society.)

536. Schmid, E. 1853. Die organischen Reste des

Muschelkalks. Neues Jahrb. f. Mineral., Geol. u.
Paleon., 15.

537. Schmidt, M. 1907. Labyrinthodontenreste aus dem

Hauptkonglomerat von Altensteig im wurtem-
bergischen Schwarzwald. Stuttgart. Mittheil. d.
Geol. Abth., d. K. Wurtemb. Statest. Landes-
amts, Jahrg. 63, No. 2, 1-10, with plate.

538. Schonfeld, G. 1911. Branchiosaurus tener Schonf.

Ein neuer Stegocephal aus dem rothl. des Nord-
west Sachsens. Isis, 19-43, with plate.

539. Schroder, Henry. 1913. Ein Stegocephalen

Schadel von Helgoland. Jahresb. d. k. preuss.
geol. Landesanstalt fur 1912, 33, Theil II, Heft 2,
Taf. 15-21.

540. Schwarz, Hugo. 1908. Ueber die Morphologie der

Wirbelsaule der Tetrapoden. Sitzungsb. d.
Gesell. naturf. Freunde zu Berlin, 315-329.

541. . 1908. Ueber die Wirbelsaule und die

Rippen holospondyler Stegocephalen (Lepo-
spondyli, Zittel). Beitrage zur Geol. u. Paleon.
Oesterreichs-Ungarns u. des Orients, Bd. xxi,
Heft 1-2, 63-105, 36 figs, and Bibliography.

542. Scott, W. B. 1908. An introduction to geology. 2d

revised edition. Amphibia, 608, 636, 652, 674,
692.

543. Sedgwick, Adam. 1905. Stegocephali in "Student's

Text-book of Zoology," 11, 313.

544. Seeley, H. G. 1875. On the posterior portion of a

lower jaw of Labyrinthodon (L. lavisi) from the
Trias of Lidmouth. Quart. Jour. Geol. Soc.,
xxxil, 278-284, pi. xix.

545. . 1879. Amphibians from the Permian Rocks

of Bohemia. Geol. Mag., Dec. 11, vi, 521-528, 4
woodcuts; 1885, 80-87. (A review of Fritsch's

work.)

54&-

1892. Further observation on Pareiasaurus.

547-
548.
549-

55<>-
551.

552-
553-

554-

555-

556.

557-

558.
559-

560.
561.

562.

563
564-
565-

566.
567-

Seeley, H.G. 1907. AnewLabyrinthodontfromthe
Karoo Beds, Cape Colony. Geol. Mag., Dec. v,
iv, No. x, 433-434 pi. 1.

. 1908. A Labyrinthodonl tooth from the

upper Karoo Beds, South Africa. Geol. Mag.,
Dec. v, v, No. vi, 241-243, pi. x.

Smith, John. Footprints in red calciferous sand-
stones in a railway cutting 2% miles N. of
West Kilbride. Trans. Geol. Soc. of Glasgow,
ix, 201.

Stejneger, Lkonhakd. 1904. Amphibia versus
Batrachia. Science, n.s., xx, 924.

. 1907. Herpetology of Japan and adjacent

territory. Bull. No. 58, U. S. Nat. Mus., 1-576,
pis. i-xxxv, and figs.

Steinmann-Doderlein. 1890. Elemente der Pale-
ontologie. Amphibia, 601.

Steinmann, Gustav. 1907. Einfuhrung in die
Paleontologie. Zweite Auflage. 2 Klasse: Am-
phibia, Lurche, 421-427, figs.

Stephens, A. T. 1886. On the Biloela Labyrinthc-
dont. Proc. Linn. Soc. New South Wales, u
ser., 1, 1 1 13.

1888. On some additional Labyrinthodont

fossils from the Hawksbury sandstone of New
South Wales. Proc. Linn. Soc. New South Wales,
1 (2). 1175-1192.

-. 1888. Note on a Labyrinthodont fossil

from Cockatoo Island (Mastodonsaurus). Proc.
Linn. Soc. New South Wales (2), I, 931-940.

1888. Second note on Platyceps wilkin-

soni. Proc. Linn. Soc. New South Wales, 11 (2),

156-158, pi. 1.
Sternberg, C. H. 1901. The Permian life of Texas.

Trans. Kans. Acad. Sci., 94-98.
Stevenson, J. J. 1907. Carboniferous of the Appa-
lachian basin. Bull. Geol. Soc. Amer., xvm, 29-

178.
Storrie, J. 1893-4. Notes on a tooth of a species

of Mastodonsaurus. Trans. Cardif. Nat. Soc.,

xxvi, pt. 11, 105, pi.
Stock, Thomas. 1881. On some British specimens

of the "Kammplatten" or " Kammleisten " of

Professor Fritsch. Ann. and Mag. Nat. Hist.,

5th ser., vm, 90-95, pi. vi.
. 1882. Notice of some discoveries recently

made in Carboniferous vertebrate paleontology.

Nature, xxvn, Nov. 2d, 22. (Describes new

species of Ophiderpeton (nanum) and Pteroplax

(cornuta).)
. 1882. Further observations on Kamm-
platten and Note on Ctenopthychius pectinatus.

Ann. and Mag. Nat. Hist. (5), ix, 253-257.

pi. viii.
Strickler, Ludwig. 1899. Ueber den microscop-

ischen Bau der Faltenzahne von Eryops mega-

cephalus. Paleontographica, xlvi, 85-94, 1 fig.,

Taf ein xi-xii.
Thevenin, Armand. 1905. Sur lc Decouvcrte

d'Amphibiens dans le Terrain houiller de Com-

mentry. Comptes Rendus Acad. Sci., cxli (26),

1268-1269.
. 1906. Amphibiens et Reptile du Terrain

Houiller de France. Annales de Paleontologie,

Tome 1, fas. in, 1-19, 2 pis.

1909. Les Stades d'Evolution des plus

Phil. Trans. Roy. Soc. London, clxxxiii, 367.

anciens Quadruples trouves en France. Comptes
rendus des Acad. Sci., cxlix, 1222, Dec. 20.

216

THE COAL MEASURES AMPHIBIA OF NORTH AMERICA.

568. Thevenin, Armand. 1910. Les plus anciens Quad-
ruples de France. Annales de Paleontogie, Tome
V, 1-64, pis. i-ix; V, fas. 1, 1-40, pis. 1-6.

569 Thompson, James, and Professor Young. 1869.
On new forms of Pteroplax and other Carbonifer-
ous Labyrinthodonts and other Megalichthys
with notes. Rept. Brit. Assn. Adv. Sci., pt. II,
101.

570. Thyng, F. W. 1906. The squamosal bone in Tet-

rapodous Vertebra ta. Tufts College Studies, n,
No. 2, scientific series, 35-74.

571. Tomes, Chas. S. 1878. A manual of dental Anat-

tomy, human and comparative. Sth ed., 65.

572. Tomes, John. 1850. On the structure of the den-

tal tissues of the order Rodentia. Phil. Trans.
Roy. Soc. London, 532. (Remarks on the struc-
ture of the teeth of Labyrinthodon jaegeri.)

573. Trautschold, H. 1884. Die Reste permischen

Reptilien des Paleontologischen Cabinets der
Universitat Kasan. Nouv. Mem. de la Soc. Imp.
Nat. Moscou, xv, 5-39, Taf. 1-8.

574. Tschudi, Johannes J. von. 1839. Classification der

Batrachier mit Berucksichtigung der fossilen
Thiere dieser Abtheilung der Reptilien. Mem.
de la Soc. Scientif. Nat. de Neuchatel, 4to, 11.
Review: Neues Jahrb. f. Mineral. Geol. Paleon.,
1841,835-840.

575. . 1837. Ueber den Homo diluvii Testis A.

Scheuchzer. Neucs Jahrb. f. Mineral., Geol. u.
Paleon., 545~547-

576. Twelvetrees, W. H. 1880. On a Labyrinthodont

skull (Platyops ricardi Twel.) from the upper
Permian cupriferous strata of Kargalinsk, near
Orenburg. Bull, de la Soc. Imp. des Natl, de
Moscou, lv, No. 1, 117-122, with figs.

577. Udden, J. A. 1901. Geology of Jefferson Co., Iowa.

Iowa Geol. Surv., xn, Ann. Rept., 406.

578. Versluys, J. 1909. Die Salamander und die

ursprunglichsten vierbeinigen Land wirbel thiere.
Naturwissenschaftliche Wochenschrift, Neue
Folge, Bd. vih, No. 3, 1-28, figs. 1-5.

579. Vidal, Louis Mariano. 1902. Sobre la presencia

del tramo Kimeridgense en el Montseih y hallazgo
de un Batracio en sus hiladas. Memorias Real
Academ. Cienc. y Artes de Barcelona, iv, No. 18.

580. . 1902. Sur le presence de l'etage Kime-

ridgien an Montseih (Province de Lerida, Es-
pagne) et decouverte d'un Batracien dans ces
assises. Memorias Real Academ. Cienc. y
Artes de Barcelona, iv, No. 18.

581 . Vogt, Carl. 1854. Archegosaurus und all Labyrintho-

donten sind Amphibien nicht Reptilien. Neues
Jahrb. f. Mineral., Geol. u. Paleon., 676-677.

582. Wagner, A. 1861. Neue Beitrage zur Kenntniss

der vorweltlichen Fauna des lithograph. — Schie-
fers. Abhandl. der k. Bayer Akad. der Wissen.
Classe, Bd. 11, Abth. ix, 3-60 and 415-528.

583. Waldheim, Fischer von. 1840. Nachtrag zu

Herrn Major von Qualen's geognostischen Bei-
tragen zur Kenntniss des westlicher Urals. Bull,
de la Soc. imp. des Natl, de Moscou, 488.

584. . 1847. Notice sur quelques Sauriens de

l'Oolithe du Government de Simborsk. Bull.
Soc. Nat. de Moscou, xx, pt. 1, 364, pi. v.

585. Walterstorff, W. 1887. Ueber fossile Froesche

insbesondere das Gattung Pakeobatrachus. Theil
i.Theil 11, 8vd. Jahresbericht d. naturw. Vereins
zu Magdeburg, 1-8 1.

586. Walterstorff, W. 1901. Ueber ein Exemplar von

Rana meriani von Meyer in Senkenbergischen
Museum zu Frankfurt. Bericht Senkenb. naturf.
Gesell., 39-51, pi.

587. Ward, J. 1890. On the geological features of the

North Staffordshire Coal-Field. Trans. North
Staffordshire Inst, of Mining and Mechanical
Engineers, x.pl.ix.

588. . 1900. On the occurrence of Labyrinthodont

remains in the Keuper sandstone of Stanton.
Trans. North Staffordshire Field Club, xxxiv,
108-112, pis. iv-v.

589. . Paleontology of the Patty Coal Field. Mem.

Geol Surv. United Kingdom, pt. in, 285-357.

590. Watson, D. M. S. 1914. The Cheirotherium.

Geol. Mag., vi, 1, No. 603, 395-398.

591. . 1912. The larger Coal Measure Amphibia.

Mem. and Proc. Manchester Lit. and Philos.
Soc, lvii, pt. 1, No. 1, 1-4, 1 pi.

592. — ■ — — . 1913. Batrachiderpeton lineatum, Hancock

and Atthey, a Coal-Measure Stegocephalian.
Proc. Lond. Zool. Soc, pt. iv, 949-962, with 2
plates and figures in text.

593. . 1913. The early evolution of the Amphibia.

Rept. 83d Meeting British Assn. Adv. Sci., 532.

1912. On some Reptilian lower jaws. Ann.

594-

595-

and Mag. Nat. Hist., 8th series, vol. 10, July-Dec,
573-587. with figures.

-. 1913. Micropholis Stowi Huxley, a temno-

spondylous amphibian from South Africa. Geol.
Mag., 5th Dec, io, 340-346, with figures.

596. Weiss, Ch. Ernst. 1871. Paleontologisch geog-

nostische Untersuchungen aus dem Gebirge auf
der Sudseite der rheinischen Devons. Bonn.
Sitzungsb. Niederrhein Gesell., 33-35. (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