Syllabus of lectures on the vertebrata

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Publications

OF THE

University of Pennsylvania.

Syllabus of Lectures

ON THE

VERTEBRATA

EDWARD D. COPE, Ph. D.,

Late Professor of Zoology and Comparative Anatomy in the

University of Pennsylvania.

With an Introduction by
HENRY FAIRFIELD OSBORN, Sc. D.,

Da Costa Professor of Zoology in Columbia University.

PHILADELPHIA:
PUBLISHED FOR THE UNIVERSITY OF PENNSYLVANIA.
1898.

ve

Press of Levytype Co., Engravers and Printers, Philadelphia.

THE LIFE AND WORKS OF COPE

ILLUSTRATING THE TRAINING OF A NATURALIST AND THE
ESSENTIAL CHARACTERISTICS OF A GREAT
COMPARATIVE ANATOMIST.

BY HENRY FAIRFIELD OSBORN.

The work of Professor Cope began in 1859, a most favor-
able year, when Comparative Anatomy first felt the impetus
of Darwin’s “ Origin of Species.” He was then only nine-
teen, and for thirty-eight years thereafter his active genius
hastened our progress in the knowledge and classification
of all the great divisions of the Vertebrata.

He passed away upon April 12th, 1897, at the age of fifty-
seven, in the full vigor of his intellectual powers, leaving
a large part of his work incomplete. Almost at the last
he contributed several reviews to the American Naturalist,
and upon the Tuesday preceding his death he sent to the
press this Syllabus of his Lectures before the University of
Pennsylvania, containing his latest opinions regarding the
arrangement and evolution of the Vertebrata. It seems
appropriate in this posthumous edition of the Syllabus, is-
sued by the University, to give a brief account of such inci-
dents in Professor Cope’s life as are worthy of imitation by
coming generations of students; also to set forth some of
the monumental features of his contributions to Compara-
tive Anatomy.

The most conspicuous feature of his character from
boyhood upwards was independence; this was partly the
secret of his venturesome and successful assaults upon all
traditional but defective systems of classification.

As a comparative anatomist he ranks both in the range

iv LIFE AND WORKS OF COPE.

and effectiveness of knowledge and, ideas with Cuvier and
Owen. When we consider the short life of some of the
favorite generalizations of these great men he may well
prove to be their superior as a philosophical anatomist.
His work, while inferior in style of presentation, has an-
other quality which distinguishes that of Huxley—namely,
its clear and immediate perception of the most essential
or distinctive feature in a group of animals. Asa natural
philosopher, while far less logical than Huxley, he was
more creative and constructive, his metaphysics ending in
theism rather than agnosticism.

Cope is not to be thought of merely as a specialist. After
Huxley he was the last representative of the old broad-
gauge school of anatomists and he is only to be compared
with members of that school. His life-work bears the
marks of great genius, of solid and accurate observation as
well as of inaccuracy due to bad logic or haste and over-
pressure of work. Although the greater number of his
Natural Orders and Natural Laws will remain as perma-
nent landmarks in our science a large part of his syste-
matic work will require laborious revision and is thus far
from standing as a model to the young zoologist.

In mere mass of production Cope’s work was extra-
ordinary. He leaves twenty octavo and three great quarto
volumes of collected researches. By his untimely death a
wide gap is left which can never be filled by one man.

BIoGRAPHICAL.

Edward Drinker Cope was born in Philadelphia, July
28th, 1840, of distinguished American ancestry. His great-
grandfather, Caleb Cope, was the staunch Quaker of Lan-
caster, Pa., who protected Major Andre from mob violence.
Thomas Pim Cope, his grandfather, founded the house of
Cope Brothers, famous in the early mercantile annals of
Philadelphia. His father, Alfred, the junior member of

LIFE AND WORKS OF COPE. Vv

this firm, was a man of very active intellect, and showed
rare judgment in Edward’s education.

Together the father and son became brisk investigators,
the father stimulating, by questions and by travel, the
strong love of Nature and of natural objects which the son
showed at an unusually early age. In August, 1857, they
took a sea voyage to Boston, and the son’s journal is full of
drawings of jellyfish, grampuses and other natural objects
seen by the way. When eight and a half years old he
made his first visit to the Museum of the Academy of
Natural Sciences, “on the 21st day of the 10th Mo., 1848,”
as entered in his journal; he brought away careful draw-
ings, measurements and descriptions of several larger birds,
but especially the figure of the entire skeleton of an Icthyo-
saur, With this quaint memorandum: ‘Two of the scler
otic plates look at the eye—thee will see these in it.” Avr
the age of ten he was taken upon a longer voyage to the
West Indies. It is not improbable that these voyages ex-
erted a lasting influence upon him.

The principal impression he gave in boyhood was of in-
cessant activity in mind and body, of quick and ingenious
thought, reaching in every direction for knowledge, and of
great independence in character and action. It is evident
that he owed far more to the direct study of Nature and to
his own impulses as a young investigator than to the five or
six years of formal education which he received at school.
He was especially fond of map drawing and of geographi-
cal studies. His natural talent for languages may have
been cultivated in some degree by his tutor, Dr. Joseph
Thomas, an excellent linguist, editor of a biographical
dictionary. Many of his spare winter hours were passed
at the Academy of Natural Sciences. After the age of
thirteen the summer intervals of boarding-school life and
later of tutoring were filled among the woods, fields and
streams of Chester County, Pa., where an intimate knowl-

vi LIFE AND WORKS OF COPE.

edge of birds was added to that of batrachians, reptiles, and
insects. He showed a particular fondness for snakes. One
of these excursions, taken at the age of nineteen, is de-
scribed in a letter to his cousin (dated June 24, 1859), in
which, at the close of a charming description of the botany
of the region, appears his discovery of a new. type:

“T traced the stream for a very considerable distance upon the
rocky hillside, my admiration never ceasing, but I finally turned
off into the woods towards some towering rocks. Here I actually
got to searching for Salamanders and was rewarded by capturing
two specimens of species which I never saw before alive. The first
(Spelerpes longicauda) is a great rarity here. Iam doubtful of its
having been previously noted in Chester County. Its length is 6
inches, of which its tail forms nearly four. The color is deep
brownish yellow, thickly spotted with black, which becomes con;
fluent on the tail, thus forming bands. To me a very interesting
animal—the type of the genus Spelerpes, and consequently of the
subfamily Spelerpinz, which I attempted to characterize in a paper
published in the Proceedings of the Academy of Natural Sciences.
I send thee a copy, with the request that thee will neither mention
nor show it,* for—however trifling—I would doubtless be miserably
annoyed by some if thee should. Nobody in this country (or in
Europe, of owrs) knows anything about Salamanders, but Professor
Baird and thy humble coz., that is, in some respects. Rusconi, the
only man who has observed their method of reproduction, has writ-
ten enough to excite greatly one’s curiosity and not fully satisfy it.
With suitable appliances of aquariums, etc., I should like to make
some observations. The other Salamander I caught was Plethodon
glutinosum—the young—remarkable for the great number of teeth
that lie together in two patches on the ‘basisphenoid’ bone; about
300 or more.”

Another passage gives an insight into his strong opinion,
so often expressed afterward, as to what constitute the real
pleasures of life :

“Pleasant it is, too, to find one whose admiration of nature and

_ *This passage probably indicates that he was sensitive to being laughed at for
his interest in these animals.

LIFE AND WORKS OF COPE. vii

detail is heightened, not chilled, by the necessary ‘investigation ’—
which, in my humble opinion is one of the most useful as well as
pleasing exercises of the intellect, in the circle of human study.
How many are there who are delighted with a ‘fine view,’ but who
seldom care to think of the mighty and mysterious agency that
reared the hills, of the wonderful structure and growth of the
forests that crown them, or of the complicated mechanism of the
myriads of higher organisms that abound everywhere ; who would
see but little interesting in a fungus, and who would shrink with
affected horror from a defenseless toad * * * Dr. Leidy is getting
up a great work on comparative anatomy which is to be the modern
standard. Such a work will be very useful to those who want to go
to the bottom of natural history ; it is an interesting study, too, to
notice the modification in form—the degradations,} substitutions,
etc., among the internal organs and bones. The structure, forms
and positions of teeth, too, are interesting to notice—so invariably
are they the index of the economy and the position in nature of the
animal.’’

This is the reflection of a lad of nineteen, an age at which
some modern educators would have us believe our young
men are just ready for the collegiate Freshman class. It is
obvious from other portions of the letter that by this time
young Cope’s career was fully determined in his own mind.
During the same year he went to Washington to study
and work in the Smithsonian Institution under Spen-
cer F. Baird, and it is amusing to observe him in the above
letter classing himself with Baird as the only Americans
who knew anything of the Batrachia.

Upon April 19, 1859, he contributed his first paper (al-
luded to above) to the Academy “On the primary divisions
of the Salamandridx, with a description of two new species.”
He followed this by a full description, in the same year, of
reptiles brought from West Africa by Du Chaillu, naming
several new forms; also by a catalogue of the venomous
snakes in the museum. In the succeeding three years he

+A word used by French writers of the time to express lines of evolutionary
descent.

viii LIFE AND WORKS OF COPE.

made twenty-four communications upon the Reptilia and
established himself at the age of twenty-one as one of the
leading herpetologists of the country.

Even in the papers he presented at this early age he
shows keen observation and powers of systematic diagnosis,
a wide range of self-acquired knowledge, and familiarity
with the personal and scientific characteristics of his dis-
tinguished seniors, Agassiz and Leidy. This period in-
cluded a year’s study (1858-59) of anatomy and clinical
instruction at the University of Pennsylvania. In 1863
he travelled abroad for several months, visiting especially
the museums of Leyden, Vienna, and Berlin, and greatly
extending his horizon as a comparative anatomist, for upon
his return he at once showed the impulse of a more philo-
sophical spirit, complete familiarity with the history of
opinion, and marked power of generalization. Thus his
papers, which begin to crowd the pages of the Proceedings
of the Academy of Natural Sciences, chiefly in recent herpe-
tology and ichthyology, display a new breadth and range
as seen in his division of the Anura into the Arcifera and
Raniformes (Firmisternia) and his demonstration of the
main evolution principles in these groups.

In 1864 Haverford College called him to a_professor-
ship of natural science. This position, however, he held
for only three years. Twenty-two years later he again re-
sumed teaching as a Professor of Geology and Paleontology
in the University of Pennsylvania, all the interval having
been devoted to exploration and research. In 1865 he first
began to extend his studies among the Mammalia, especially
the Cetacea, recent and extinct, of the Coastal Tertiary.
Early in 1866 a wider paleontological field opened in the
vertebrata of the Cretaceous marls of New Jersey, whence he
procured the remains of Dinosaurs, describing especially
the carnivorous Lelaps. In the same year appeared the
continuation of his tropical American and Sonoran herpe-

\

LIFE AND WORKS OF COPE. ix

tology and his third contribution to the history of the Ce-
tacea. Henceforward his papers become far too numerous
to consider together and we must endeavor to follow merely
the main outlines of his life-work.

This was a bright era in the history of the Philadelphia
Academy ; Leidy, Gill,and Harrison Allen being frequent
contributors. In 1868 Cope gave his first complete synop-
sis of the extinct Amphibia of the world. Between 1868
and 1870 he made his first six contributions upon the Plesio-
saurs and Mosasaurs of the Cretaceous of Kansas, and in
1871 began his first western explorations in these beds.
This led to his appointment as Vertebrate Paleontologist
of the U.S. Geological Survey, under Dr. Hayden, and to
further explorations in Wyoming (1872) and Colorado
(1873), which resulted in the discovery of many new types
of fishes, mosasaurs, chelonians, dinosaurs and other rep-
tiles, notably, Portheus, Platecarpus, Clidastes, Compsemys
(type of the Amphichelydia, Lydekker), Protostega, and
Agathaumus. These were described chiefly in the Annual
Reports of the U. S. Geological Survey and in the Proceedings
and Transactions of the American Philosophical Society, and
then culminated in his first large volume “ The Vertebrata
of the Cretaceous Formations of the West,’ No. II, of the
Hayden quartos, which was published in 1875.

He spent his summers in the Bad Lands, exploring the
Bridger and Washakie, Wasatch, New Mexican, and Judith
River (1887) formations. The latter exposures he visited
in 1874, in connection with the Wheeler Survey (Geological

‘Surveys West of the 100th Meridian) securing a collection
which is now preserved in the National Museum, and pub-
lishing a most vivid description of the geology of this in-
teresting region. His movements in the field are described
by one who was with him as so rapid and full of energy,
so regardless of food and rest, that he wore out the other
members of his parties and did not allow time for thorough

x LIFE AND WORKS OF COPE.

search ; yet he himself discovered a number of his most im-
portant types.

The fruits of the New Mexican journey appeared in
many bulletins and were finally collected in his second
great volume, “The Extinct Vertebrata obtained in New
Mexico by Parties of the Expedition of 1874,” Vol. IV., of the
Wheeler Survey. In 1874 appeared the first of his studies
upon the comparison of American and European horizons,
and of his contributions to the John Day fauna. His col-
‘lections were now accumulating so rapidly as to demand
more time for research and for many years he was fortunate
in securing the field services of Mr. C. H. Sternberg and
especially of Dr. J.'L. Wortman. He continued to make
brief expeditions, among the last being his trip into the
Laramie region..,

As early as 1868 it may be said that he had laid the
foundations for five great lines of research, which he pur-
sued concurrently to the end of his life; these must, how-
ever, be followed separately to be understood and ap-
preciated. Only for comparatively brief intervals would
he pursue one line exclusively in order to complete some
special memoir, because his marvelous memory apparently
held and resumed the details of all the others with perfect
ease.

CoMPARATIVE ANATOMY OF THE FISHES.

Cope’s work in Ichthyology would alone give him high
rank among zodlogists. His friend, Professor Theodore
Gill* who has largely contributed to this section of the
biography, observed that it was among the Fishes that Cope
had rendered his greatest contributions. The same obser-
vation, however, has been made by Professor Baur in rela-
tion to the Amphibians, and by Scott and Osborn in refer-
ence to the fossil Mammalia.

"* Professor Gill has kindly allowed the writer to use the advance sheets of his

memorial address before the American Philosophical Society upon Professor Cope’s.
contributions to Ichthyology and Herpetology.

LIFE AND WORKS OF COPE. xl

As early as December, 1861, Cope made what was called
a verbal communication to the Academy of Natural Sciences
of Philadelphia, in which he noticed several Cyprinoid
fishes, two of which he regarded as new, but which were
really identical with forms that had been previously de-
scribed in an imperfect manner. In his early papers (1864)
he appears as an enthusiastic systematist, studying especi-
ally the living forms of Teleosts, making careful diagnoses
of all types that came into his hands, critically considering
the problems of distribution, never casting aside those types
whose especial difficulties had been the stumbling block of
earlier writers. Thus he studied successively the fishes of
Michigan (1864-65), of Virginia (1868), of the Lesser An-
tilles (1870), again the fishes of South Carolina (1871), of
Alaska (1872), of Montana, those from South America col-
lected by Professor Orton (1872-78), those from the terri-
tories collected by the Wheeler Survey, and even not infre-
quently new forms from Africa and the East Indies.

Almost from the first Cope ventured upon ground which
had been trodden only by the greatest comparative anato-
mists and ichthyologists, such as Cuvier, Agassiz, Owen,
and Giinther, and set aside the superficial characters which
had been employed in the classification of the fishes. He
seems to have been moved by an instinct to seize upon the
less conspicuous structures which were none the less of
fundamental importance, and to disregard the more con-
spicuous features, such as the scales, which had formed the
chief guide of his predecessors, especially of Agassiz.

A fortunate step in his career was his purchase, while
abroad, of Professor Hyrtl’s private collection of fish skele-
tons, which gave him nearly a thousand admirable osteol-
ogical preparations for immediate study.

A brief glance at history suffices to make. clear at what
juncture Cope entered this field of science. The term Gan-
omDEs had been originally introduced into zodlogy by Agas-

xii LIFE AND WORKS OF COPE.

siz for all forms having enamel-covered scales, and as ap-
plied by him, covered a very heterogeneous combination of
fishes totally unrelated to each other. Johannes Miller
later retained this name for fishes which were separated
from other living forms by a chiasma of the optic nerves, a
multi-valvular and muscular conus arteriosus, and an in-
testinal spiral valve. In this shape the group, Ganoidei,
was long accepted as a subclass or order, but in 1866,
Owen broke away from the historic regard for external
characters, by uniting the Ganoids and Teleosts into the
TELEOSTOMI.*

Before the American Philosophical Society, in 1870,
and the American Association for the Advancement of
Science in 1871, Cope maintained that the primary divi-
sions of the Teleostomi are indicated by their fin structure,
enunciating a principle which now forms the accepted basis
of subordinal classification. Fin structure as a taxonomic
motive became uppermost in his mind among all the possi-
bie keys to the classification of the fishes. It undoubtedly
served to direct his attention later to the foot structure of
land vertebrates, especially of the great Dinosaurs in 1867
and the hoofed mammals in 1880, as of diagnostic value.

“The evolution of the fins, indeed, and especially of the paired
fins, is shown by Cope to be the most satisfactory and philosophical
clue to the arrangement of all the minor groups of fishes. Just as
the various modifications of the pentadacty] limb in the Ungulate
Mammals—the vertebrates which eventually become most com-
pletely adapted for progression on land—afford the principal means
of determining the natural subdivision of that order; so among the
greater groups of fishes—the vertebrates that become specially
adapted for progression in water—the successive modifications of the
primitive fin-folds form the most obvious clue to the phases through
which the various types have passed in the course of their spec-
ialzation.” (A. Smith Woodward, Catalogue of Fossil Fishes, Pt.
II. p. x.)

a ne dahes having the mouth surrounded by specialized membrane bones as in the
eleosts.

LIFE AND WORKS OF COPE. xiii

“The most interesting feature of the Crossopterygii consists in the
mode of specialization of their fins ; and this, as pointed out by
Cope, affords a satisfactory basis for the definition of the suborders.”
(Ibid, Pt. II, p. xxi.)

The objections of Kner and others were reinforced by
Cope with arguments derived from the osseous parts; and
he concluded that it was “evident that the subclass Ganoi-
dea cannot be maintained”. He then proceeded to the
consideration of the value of other characters and at length
resolved to recombine the scattered elements of the great
class of Pisces, limited by the exclusion of the Leptocar-
dians and Marsipobranchs, into five primary subdivisions
or subclasses.

In these papers he therefore boldly abandons the group
of Ganoids and redistributes the fishes into five great sub-
classes, namely, the Holocephali (Bonaparte, 1832-41), Se-
lachat (Cuvier, 1817), the Dipnoi (Miller), the Crossopterygu
(Huxley) with paired fins arranged so as to form a fringe
around a central lobe, and the Actinopteryi. He showed
that each of these forms a natural group, and the differen-
tiation between each and its nearest of kin is on the whole
well marked. Second to this emphasis upon fin structure
was that upon the jaw structure, or the union of the upper
jaw with the skull, which separates the Holocephali from the
Selachii. Third, he employs the modifications of the res-
piratory system for combinations into superfamily groups
of the Physoclysti, and the number of tail vertebre for the
combinations of Physostomt.*

Of these five great divisions he adopted Holocephali
from Bonaparte, the Crossopterygii from Huxley, propos-
ing on his own part the Actinopteryi or ray-finned fishes.
The latter great subdivision which comprises the majority

* “The greatest defect in this system,” says Professor Gill, ‘was Cope’s failure to
emphasize the important distinction between the cartilaginous fishes (Holocephali
aad Selachii) with no membraneous skeleton, and the Dipnoi, Crossopteri and
Actinopteri, which have a specialized system of membrane bones transmitted with
more or less modifications to all the higher Vertebrata,”

xiv LIFE AND WORKS OF COPE.

of living types he broke up into a number of groups by an
originality of analysis as marked asin his separation of the
five great primary divisions. He first divided the subclass
into three groups, named, Chondrostei, Physostomi; and Phy-
soclysti (he subsequently removed the Chondrostei from the
Actinopteri, and applied the name Malacopteri to the Phy-
sostomes, and Acanthopteri to the Physoclysts).

“No less than 24 orders were recognized in the Observa-
tions,” says Professor Gill,* ‘to receive the Acanthopterous
fishes and these were subsequently added to. Of the 24 or-
ders as many as 15 were endowed with new names. Eight of
the names have been adopted by most American naturalists
as the designations of orders or suborders. These are Sel-
achostomt, Scyphophort, Plectospondyli, Haplomi, Enchelyceph-
ali, Colocephali, Percesoces and Hemibranchit. The other
names have been sunk as synonyms. The most meritori-
ous of Cope’s generalizations expressed in ordinal terms, in
some respects at least, were those involving the constitu-
ents of the order Plectospondyla and the separation of the
eel-like fishes into different groups.

The order Plectospondyli (fishes with the anterior verte-
bree coalesced, as distinguished from Isospondyli (Cope) in
which all the vertebree are separated) was framed for the
numerous fresh-water fishes comprised in the families Ca-
tostomide (Suckers), Cyprinide, (Carps, etc.,), Cobitide
(Loaches), Sternopygide (Carapos, etc), and the numerous
South American and African fishes representing the Cyprin-
oideans, constituting the families Characinide and Ery-
thrinide. These had been arranged previously in three
groups widely separated and superficially quite unlike each
other and were considered to be related to forms with
which they are now known to have little in common ex-
cept some external characteristics. Their combination
fessor Gills Memorial Address hefore te American Philosepiical Society as far es

the middle of page xvi, Anumber of minor changes and insertions are made
elsewhere in order to make the matter clearer to students.

LIFE AND WORKS OF COPE. xv

in one great group or order (by Cope) was a most happy
one and even the diagnosis given was good. Nevertheless
it was long before the order was recognized by other icthy-
ologists. That it was not recognized earlier was partly due
to the inconsistencies in Cope’s own presentation, but still
more to doubts arising from certain unfortunate complica-
tions.

The main characters he assigned to the Plectospondyli
were the “anterior four vertebree much modified, and with
ossicula auditus,” also the presence of a preecoracoid arch.
But from this group he differentiated two large orders,*
distinguished by simple anterior vertebre, including the
form (Gymnotus, the electric eel) which had been pre-
viously placed in the same family with the Sternopygide,
and which appeared to all other investigators to be at least
very nearly related. With many contradictions con-
fronting other workers, it is not strange that they did not
at once accept Cope’s classification. But his intuition was
better than his logic or his application of diagnosis. Time
went on. It was ascertained that there was no distinction
between the electric eel and the Sternopygids, such as he
had alleged; it was ascertained that the opinion (which he
shared with distinguished predecessors) that the curious
cat-fishes known as Hypophthalmids had “vertebra wnmodi-
fied” was the extreme reverse of the truth; they were found,
in fact, to have the first four vertebrae not only co-ossified
but so modified and crowded that they were shoved into
the skull and could only be seen when the skull is bisected;
the anterior “unmodified vertebre ” of Cope and others be-
ing those which succeeded the modified ones.

Thus the objections against Cope’s classification were

*The electric eel was isolated as the representative of a peculiar order
(Glanencheli) because it was supposed to have ‘“‘no precoracoid.” Again, the com-
pination of “four vertebra co-ossified, and with ossicula auditus”’ was attributed
to another order (Nematognathi), but within the same order a group was admitted
with the attribute of ‘vertebre unmodified,” although it was remarked that the
fishes in question, ‘the Hypophthalmidae, are indeed scarcely to be reterred to this
order.”’

Xvi LIFE AND WORKS OF COPE.

successively dissipated, and the order Plectospondyli, modi-
fied by the addition to it of his Glanencheli, stands out as
one of the most important of his system.”

All fishes having an eel-like form, continues Professor
Gill—that is, with a long, snake-like aspect and without
ventral fins—had long been supposed to be nearly related
to each other and, if not forming one family (as the older
authors believed), to constitute at least a natural series of
families. Cope, however, demonstrated that there were
great diversities in internal structure among fishes charac-
terized by an external eel-like form. For example, the
electric eel and the Sternopygids are not at all related to
the typical eels, but really belong to the same great group
as the carp, dace, and roach. Although Cope was not the
first to recognize this absence of relationship to the eels, he
has the unquestioned merit of having first recognized to
what others the Sternopygids were to be approximated.
Moreover, he segregated the true eels into four ordinal
groups, two of which (Ichthyocephali and Holostomi) are now
generally combined in the order Symbranchia, while the
other two are recognized as suborders of the Linnean order
Apodes—an order that is Linnean, however, only in name
and in type.

Professor Gill closes with the following critique:

“J cannot consider his removal of the ‘subclass Dipnoi’ to a posi-
tion between the Holocephali and his Elasmobranchii (Plagiostomi, )
or his combination of his old subclass Crossopterygia with his sub-
class Actinopteri as improvements. But the subdivision of the
Crossoptergyia into Rhipidopterygia and Crossopterygia (or what
may be rather called Eucrossopterygia) and the new orders (or sub-
orders) which he established appear to express morphological details
as near as can be done with present material; at least his views,
with slight modifications, have been accepted by the best informed
living student of the extinct forms—I mean, of course, A. Smith
Woodward of the British Museum.”

LIFE AND WORKS OF COPE. xvii

Fossit FisHes.

In 1876 Huxley accepted Cope’s separation of the Holo-
cephali from the Selachii, and these five groups of living
fishes stand as the major classification of the present day,
which now rests upon the structure of the fins and of the
skull, owing chiefly to the labors of Owen, Huxley, and
Cope. Cope’s interest in the ancestry of the fishes was
naturally intensified as he became a more convinced evolu-
tionist, and by his increasing knowledge of the extinct
forms. Here his wide preliminary studies among living
types stood him in good stead, and stand as a model of the
close relations which should always subsist between zoologi-
cal and paleontological research. He was first brought into
contact with extinct fishes among the Cretaceous vertebrate
remains from the New Jersey green sand (1869), and con-
tinued among the rich yields of the Green River Tertiary
whales (1871-77) belonging to the Lower Eocene. Devonias
and Carboniferous or older fishes had long been in the
hands of Professor Newberry, the pioneer among the stu-
dents of Paleozoic fishes of North America. Cope’s dis-
coveries led him rather among the intermediate types of
the Permian period.

In his eager quest of phylogenetic relationships his sys-
tematic genius led him always to suggestive and often to
permanent results. In fact, the masterly part which Cope
has played in the major classification of the fossil, as well
as of the recent fishes, may well be gathered from a perusal
of the introduction of the three parts already published of
the “Catalogue of Fossil Fishes in the British Museum,”
by Woodward, from which many of the subjoined quota-
tions are taken.

In 1884 he proposed a new subclass of the Selachii,
namely the Ichthyotomi, founded upon the Permian genus
Diplodus, and subsequently enriched by his discovery of
Didymodus. Of these Woodward says: ‘In discuss-

Xvill LIFE AND WORKS OF COPE.

ing the bearing of the foregoing facts upon published
schemes of classification of the Klasmobranchii, the first
point to be considered is the validity of Professor Cope’s
division of the subclass into the two orders Ichthyotomi and
Selachii. If the characters of the dentition are of any sys-
tematic importance—and when genera of equivalent age
are under comparison we believe they are—there can be no
hesitation in associating the European and later Paleozoic
Pleuracanths with the skulls of the so-called Didymodus,
Cope, from the Permian of Texas. (Catalogue, Pt. I., p.
Xxiil.)

The order Ichthyotomi being firmly established in 1889,
Cope proposed another great suborder, Ostracodermi, to
include the peculiar armored fishes without lateral fins, but
with jointed appendages apparently articulated with the
head plates. Of these Smith Woodward speaks as follows:
‘A large number of these are still problematical, and it has
thus been deemed convenient to treat next in order the
great extinct group of Chordate animals to which Professor
Cope has applied the name of Ostracodermi. These pertain
either to the Class Pisces or to some lower denomination
yet to be determined. (Catalogue, Pt. II., p. xvii.) The
name Ostracodermi, is preferred for this subclass, because
Professor Cope seems to be the only naturalist who has
hitherto ventured to remove the Coccostean fishes far from
the order that comprises the Asterolepide.” (Catalogue, Pt.
II., p. xviii.)

For the most abstruse problems Cope had an invariable
resource of working hypotheses. Thus, the curious fish-
like Bothriolepis he compared to the armored Ascidian,
basing this surprising view upon the remarkable similarity
in the arrangement of plates, arguing that it was reason-
able to expect in the early period of Bothriolepis, that. back-
boned creatures should have been built upon the plan of
Ascidian tadpoles.

LIFE AND WORKS OF COPE. xix

Cope’s final opinions and additions to the arrangement
and phylogeny of the fishes appear in this Syllabus, and
may be considered as an extension of his earlier work upon
the fin and jaw structure. Of this Smith Woodward speaks
as follows: ‘Among the early families, the characters of
the median fins lead to the recognition of two or three
divisions. It is probable that one type in which the
median fin remains undivided and more or less in its
primitive condition will eventually be met with, even if it
be not already known. This group has received (from
Cope) the name of Haplistia, and we provisionally assign
to it the problematical Tarrasiidx. The second and third
types, though now clearly definable, are not satisfactorily
formulated in the somewhat fluctuating classifications of
Cope; and the terms Rhipidistia and Actinistia are
selected on the present occasion from those already pro-
posed by that author, as being most expressive and accu-
rate.” (Catalogue, Pt. II., p. xxii.)

STuDIES AMONG LivinG AND Extincr AMPHIBIANS.

“There never has been a naturalist.” writes Dr. Baur,
“who has published so many papers upon the taxonomy,
morphology and paleontology of the Amphibia and Rep-
tilia as Professor Cope.” The first of a series of more than
forty papers upon the former group is the one “On the Pri-
mary Divisions of the Salmandridxe, with descriptions of
two new species,” alluded to in his letter above, and pre-
sented at the age of 19 (April, 1859). It exhibited the pre-
cocious taxonomic instinct which soon afterwards prompted
him to attack and rearrange the major divisions of the
Amphibia. Rapidly following this first essay by others
upon the Anura, in 1865 and 1866 he outlined the larger
Ecaudate or Anurous divisions: I. Aglossa; Il. Bufoni-
formia; III. Areifera; IV. Raniformia.

At the age of 25 he described his first extinct Amphibian

xXx LIFE AND WORKS OF COPE.

Amphibamus, from the Carboniferous of Ohio, and at 28 he
published his first large quarto memoir, “ Synopsis of the
Extinct Batrachia, Reptilia and Aves of North America.” *
This contained, in addition to the above, the recent urode-
lous divisions, Trachystomata (Siren), Gymnophidia (Coci-
lia), Proteida (Necturus-Proteus); but of chief importance,
to include the Permian and Triassic forms of the world, he
proposed the great extinct order Stegocephali, which has
since been universally adopted. As a supplement to this
memoir appeared in 1874 his “ Catalogue of the Awr-Breath-
ing Vertebrata from the Coal Measures of Ohio,” including
results also published in the Paleontology of the Geologi-
cal Survey of Ohio of the same year. His researches and
collections in the typical coal measures and Permian ex-
tended to Iowa and ILlinois, leading to the determination
of Cricotus, which in 1880} he made the type of the sub-
order Embolomeri, or Stegocephalio with double vertebral
rings. In 1877 he received the first remains of Eryops and
Trimerorachis, from the supposed Triassic, but actually Per-
mian, beds of Texas, animals which in 1882 he made the
type of the Rachitomi, a second suborder of Stegocephalio.

This accession of material, as we shall see, ranks with that
from the Puerco among the chief events of Cope’s scientific
career, for the Permian of Texas yielded to him not only
these remarkable Batrachians with complex vertebra, but
also the great primitive representatives of the Reptilia.
The suborders Rachitomt and Embolomert have been
grouped as Temnospondyli in contrast with the specialized
Labyrinthodontia and simpler Microsauria of Europe, chiefly
made known through the labors of Fritsch, Credner, Gau-
dry, and Miall. Cope’s brief memoir of 1884 upon the
“ Batrachia of the Permian Period of North America” sum-
med up his previous contributions, but he anticipated that

* Trans. Amer. Phil. Soc., read 1868, pub. 1869. See also Proc. Phila. Acad. Nat.
Sci., 1868, p. 211.

{ American Naturalist, p. 610.

LIFE AND WORKS OF COPE. xxi

the more exhaustive monographic treatment of the rich
amphibian and reptilian fauna of this period, exclusively
collected and described by him, would constitute a volume
of the Hayden Survey memoirs and give him an oppor-
tunity of rounding up his prolonged studies. This volume
was never completed.

In the meantime his investigations upon the living
Batrachia extended to Central and South American species,
as well as to include his very original observations upon
the laws of geographical distribution of the Amphibia,
which were published by the Smithsonian Institution. In
1875 he prepared a “ Check List of the North American Ba-
trachia and Reptilia” for the U. S. National Museum; this
was followed by an essay “On the Zoological Position of
Texas” (1880). Soon afterwards, at the request of Spencer
F. Baird, Secretary of the Smithsonian Institution, he
began the preparation of a general work upon the Batra-
chia; this was facilitated by a manuscript prepared for a
work of the same character both by Baird and Girard, but
was not completed until 1889. Asa volume of 523 pages
and numerous plates this work,* while showing many
signs of haste and subject to considerable changes in the
larger systematic divisions, fortunately remains as a monu-
ment of the immense range of knowledge and observation
of its author upon the structure and habits of the living
representatives of this group. It must always be a matter
of regret that he could not have published his final views
upon the extinct forms. One of his most important gen-
eralizations from the latter, contained in a short memoir,
“The Intercentrum of the Terrestrial Vertebrata” (1881), is
that the vertebre of living amphibia are composed of inter-
centra and are, therefore, not homologous with the true
centra (pleurocentra) of reptiles, birds, and mammals.

* “Phe Batrachia of North Amerlca.”’ Bull. No. 34, U. S. Nat. Museum.

xxii LIFE AND WORKS OF COPE.

CLASSIFICATION OF THE REPTILIA BY CHARACTERS OF THE
FEET AND CRANIAL ARCHES.

We have already traced Cope’s initial work upon the
Reptilia. As in other groups, his researches rapidly
branched out in many directions, first, his treatment of the
reptiles of the Bridger and other fresh-water Tertiary lakes
in connection with the mammalian fauna; second, the con-
tinuation of his systematic description of the Kansas Creta-
ceous fauna; third, his brief papers upon the herbivorous
Dinosaurs of the Dakota (1877 and 1878) and the horned
Dinosaurs (Monoclonius) of the Laramie formations; fourth,
his numerous papers upon the Reptilia of the Triassic and
especially of the Permian.

The latter discoveries must be considered the most im-
portant and unique in their influence upon paleontology.
In 1875 he first announced the existence of reptiles in the
American Permian, and in 1877 he reported the first primi-
tive Crocodilia (Belodon) and Dinosauri (Clepsysaurus and
Zatomus) in the Triassic of North Carolina.

The detailed sequence of this reptilian work is clearly
stated by Professor Baur. Already in 1864 he published
a paper on the characters of the higher groups of the Squa-
mata,* a group proposed by Oppel to include the lizards
and snakes. ‘Two years later he made his first remarks
about the Dinosaur Lelaps,t and in 1867 he compared
the carnivorous Dinosaurs with the birds.{

“Prof. Cope gave an account of the extinct reptiles which ap-
proached the birds. He said that this approximation appeared to
be at two points. The first by the Pterosauria, to which the modi-
fied bird Archwopteryx presented points of affinity. The second,
and one not less striking, is by the Dinosauria of the orders Gonio-

* Proc. Acad. Phila., 1864, p. 224.
ae + Lelaps aquilunguis, Cope. Proc. Acad. Nat. Sci., Phila., July, 1866, pp. 275-
79.

{ Ibid., 1867, pp. 234-235.

LIFE AND WORKS OF COPE. Xxili

poda and Symphypoda. He showed the essential differences between
the ordinary Dinosauri« and the birds to consist in the distinct tar-
sal bones in two series, the anteriorly directed pubes, and the pres-
ence of teeth, of the first class. In the genus Lelaps, Cope, type
of the Goniopoda, the proximal series of tarsal bones was princi-
pally represented by one large astragaloid piece which had a very
extensive motion on those of the second series. This was immov-
ably bound to, and embraced, the tibia, and was perhaps continuous
with the fibula, much resembling the structure of the foot of the
chick of the ninth day, as given by Gegenbaur. The zygomatic
arch was of a very light description. He was convinced that the
most bird-like of the tracks of the Connecticut sandstone were made
by a nearly allied genus, the Bathygnathus, Leidy. These creatures,
no doubt, assumed a more or less erect position, and the weight of
the viscera, etc., was supported by the slender and dense pubic
bones, which were, to some extent, analogous to the marsupial bones
of implacental Mammalia, though probably not homologous with
them.

He said he was satisfied that the so-called clavicles of Iguanodon
and other Dinasauria were pubes, having a position similar to those
of the Crocodilia.

Also, that a species of Zelaps had been observed in France, by
Cuvier, which was different from the Lelaps aquilunguis, and which
he proposed should be called Lelaps gallicus.

Compsognathus, Wagner, type of the Symphypoda, expressed the
characters of the latter in the entire union of the tibia and fibula
with the first series of tarsal bones, a feature formerly supposed to
belong to the class Aves alone, until pointed out by Gegenbaur.
This genus also offered an approach to the birds in the transverse
direction of the pubes (unless this be due to distortion in the
specimen figured by Wagner), their position being intermediate be-
tween the position in most reptiles and in birds. Other bird-like
features were the great number and elongation of the vertebra of
the neck, and the very light construction of the arches and other
bones of the head.

He thought the penguin, with its separated metatarsals, formed
an approach on the side of the birds, but whether the closest ap-
proximation to the Symphypoda should be looked for here or among

XXIV LIFE AND WORKS OF COPE.

the long-tailed Ratite (ostrich, etc.), he was unable to indicate.”’
(Proc. Acad. Nat. Sci., Philadelphia, Vol. XIX, 1867, pp. 234-5.)

Huxley’s paper upon the same subject appeared in 1868.*

In 1870 Cope read an important paper before the Ameri-
can Association ‘“ On the homologies of some of the Cranial
Bones of the Reptilia and the systematic arrangement of the
class.” He discussed the following topics: 1. Homolo-
gies and Composition of the cranial arches. 2. The crani-
um of the Ichthyosauria. 3. The cranium of the Anomo-
dontia. 4. The homologies of the opisthotic. 5. The
squamosal bone. 6. The columella (epipterygoid). 7. The
systematic arrangement of the Reptilia. 8. Critical re-
marks on the system. 9. The Rhynchocephalia and sup-
posed Lacertilia of the Trias and Permian. 10. Strati-
graphic relation of the orders of Reptilia.

His classification is as follows:

A. Extremities beyond proximal segment not differentiated
as to form.

I. Ichthyopterygia: Order Ichthyopterygia.

B. Extremities differentiated.
Il. Streptostylica: Orders Lacertilia, Pythonomorpha,
Ophidia.
III. Synaptosauria: Orders Rhynchocephalia, Testudi-
nata, Sauropterygia.

IV. Archosauria: Orders Anomodontia, Dinosauria,
Crocodilia, Ornithosauria.

In 1875 the large volume “ The Vertebrata of the Cretace-
ous Formations of the West” appeared, as Vol. II of the
Rep. U.S. Geol. Surv. Territ. (802 pp., Pl. LVIL). This
work contains extensive descriptions, especially of the

*Popular Science Review, 1868, pp. 237-247.
+Proe. Assoc. Ady. Sci., XIX, pp. 194-247.

LIFE AND WORKS OF COPE. XXV

Mosasaurs, also of Testudines, Crocodilia, Plesiosaurs and
Dinosaurs (Iguanodontia): Agathauwmas, Cope (Triceratops,
Marsh); Hadrosaurus, Leidy; (Trachodon, Leidy; Diclonius,
Cope, Laosawrus, Marsh).

Cope’s most epoch-making contributions, however, are
his researches on the Permian Reptiles of Texas, which
commenced in 1878. In the Proceedings af the American
Philosophical Society of this year he established the sub-or-
der Pelycosauria of the Rhynchocephalia to contain Clepsy-
drops, Dimetrodon, Diadectes, Bolosaurus, Pariotichus, Empe-
dias. In the December American Naturalist of the same
year the order Theromorpha (reptiles having characters of
mammals) was created, with the sub-orders Anomodontia,
Owen, and Pelycosauria, Cope. : The Pelycosauria were con-
sidered as the ancestors of the Mammalia. In 1880* a new
division of the Theromorpha was established, with the
name of Cotylosauria, to contain the family Diadectide. In
a skull of Empedias he described two occipital condyles,
being misled by the missing basioccipital. In 1883+ he
placed his genera Pariotichus and Pantylus in a new family ,
Pariotichide, characterized by the over-roofing of the
temporal fossee and the presence of the supra-occipital and
par-occipital plates (intercalare, Cope). He now found the
basioccipital in position and the order Cotylosawria was
given up. In1890(March 12th), Cope, however, again em-
ployed the Cotylosauria as a suborder under the Theromora
distinguishing three families: Careiasauridx, Pariotichide,
Diadectide. In 1882t Seeley had established the order
Pareiasauria, which Lydekker (1889) and Zittel considered
as a suborder of the Theromora. In 1892§ the Cotylo-
sauria was raised to ordinal rank by Cope.

The last two papers published by Cope in the Proceed-

*American Naturalist, p. 304.
+Proc. Amer. Philos. Soc., p. 631.
{Proc. Roy. Sov., Vol. 44, p. 383.
§Trans. Am. Phil. Soc., Vol. X VII.

xxvi LIFE AND WORKS OF COPE.

ings of the American Philosophical Society give much new
evidence about this very interesting group. The titles of
these papers are: “ The Reptilian Order Cotylosauria” * and
Second Contribution to the History of the Cotylosauria.”t In
this paper a new family, Ofocelidx, was described with the
following characters: Posterior border of temporal roof ex-
cavated laterally by the meatus auditorius externus. Teeth
present in a single row, not transversely expanded. Ribs
immediately overlaid by parallel transverse derm-ossifica-
tions which form a carapace. This family he considered,
or at least suggested was, ancestral to the Chelonia.

Cope had in preparation for many years an extensive
work on the Lacertilia and Ophidia of the United States, to
be published, like his Batrachia, in a bulletin of the United
States National Museum.{ The MSS. forthis work cost him
much labor, especially during the last two years of his life
and for a while interrupted all his other work. It was
characteristic of him to turn aside for a laborious detailed
investigation upon the soft anatomy of the snakes in the
hopes of finding some satisfactory means of classifying this
puzzling group from the structure of the copulatory or-
gans. This investigation constituted his latest original
work and was barely completed before his death.

EVOLUTION AND CLASSIFICATION OF THE MAMMALIA.

Up to 1868 Leidy held the Western paleontological field
exclusively. In this year Marsh and Cope also entered the

*Proc. Am. Philos. Soc., Vol. XXXIV, 1896 (Feb. 2d), pp. 436-457, Pls. VII-X.

tIbid., Vol. XXXYV, pp. 112-139, Pls. VII-X, Aug. 15, 1896.

{Many preliminary papers have appeared for this publication, of which the
following are named:

_ ag analytical Table of the Genera of Snakes.” Proc. Am. Philos. Soc., 1886, pp,
“The Osteology of the Lacertilia.””. Proc. Am. Philos. Soc., Vol. XXX, 1892
185-221, Pls. II-VI ad

“On Degenerate Types of Scapular and Pelvic Arches in the Lacertilia.”” Journ.
Morphol hips VIT, 1892, pp. teeter ae XII

“The Classification of the hidia.’”?’ ‘trans. Amer. Philos. Soc.,
eas 1a, 1895, ep. 186-219, Pls. XIV-XXXI ote Yok aS viee

on a Le emipenes "of the Sauria.” P08. Acad. Nat. Sci., Phil., August. 1896,
pp- 7

LIFE AND WORKS OF COPE. xxvil

Western territory and began the simultaneous exploration
and description of a limited fauna in a somewhat limited
region, with the inevitable result of a struggle for priority
and a permanent rupture of friendly intercourse. It is
necessary to allude to the fact, because it greatly affected
the subsequent history of American paleontology. Fortu-
nately, the western fossil area proved to be a vast one, and
the remarkable discoveries by Wortman in the Big Horn
and Wasatch, beginning in 1878, also by Baldwin in the
Puerco of New Mexico, beginning in 1880, and the explora-
tions already described of Cummins in the Permian of
Texas, afforded Cope a noble field of research quite free
from the haste of rivalry.

As early as 1864 Cope had outlined a classification of
the Squamata, taking advantage of characters distinguish-
ing the snakes and lizards given by Miiller, also those
given by Stannius. Here, as in the fishes, Cope perceived
the superior value of the hard parts of the body to the
tongue and soft parts, which were then employed by the
greater number of anatomists; as well pointed out by Boul-
enger:

“Whilst engaged in a revision of the Lizard-collection in the
British Museum, I have felt the necessity of a thorough systematic
rearrangement of the order Lacertilia. The classifications proposed
by Dumeril and Bibron and Gray, and now still generally in use,
with slight modifications, are, on the whole, as unnatural as can be,
and founded, to a great extent, on characters of pholidosis and physi-
ognomy. Physiognomy is worth nothing asa guide in the formation of
higher groups; as to the characters afforded by the scales I have con-
vinced myself that they are very deceptive, and ought to be taken into
consideration in the definition of families only when accompanied by
other characters. Like Cope, whose lizard families I regard as the
most natural hitherto proposed, I shall lay greater stress on osteo-
logical characters and on the structure of the tongue. Special im-
portance must also be attached to the presence or absence, and the
structure, of dermal ossifications on the head and body, and these

XXVii LIFE AND WORKS OF COPE.

will be found to correspond with many other characters. Bocourt,
to whom is due the merit of having pointed out their systematic im-
portance, did not realize the very great progress made by means of
that character, the modifications of which he so ably illustrated, for
he still maintains the artificial group Scincoidiens, in spite of the
objections of Cope, whose views are evidently confirmed by the re-
searches of the French herpetologist.’’ (‘‘ Synopsis of the families
of existing Lacertilia.”” Annals and Magazine of Natural History,
Vol. XIV, Fifth Ser., 1884.) .

EvouutTion oF THE MAMMALS.

Cope’s most numerous and voluminous writings were
devoted to Mammals, and to appreciate the importance of
his contributions to this group it is necessary to cast a brief
glance over the history of mammalian paleontology.
Cuvier, the founder of this branch of science, had repre-
sented the ecole des faits in opposition to Geoffroy St.
Hilaire, and founded a school wholly opposed to general-
ization as to the origin and succession of animal life, and
firmly adherent to the Special Creation hypothesis. As a
master of comparative anatomy, Cuvier exerted an immense
influence upon the succeeding French paleontologists, such
as Jourdan, Croizet, Cristol, De Blainville, Aymard, Lartet,
and Pomel. It is true that De Blainville and Gervais
showed a wide range of knowledge, but Gaudry was the
first of the French paleontologists to grasp the spirit of evo-
lution. In Germany, Jager and Blumenbach ranked as
more or less voluminous descriptive writers, while Kaup
showed superior powers of analysis.

Cuvier’s unnatural classification of the hoofed animals
into the Solipédes, or horses, and Pachyderms, or rhinoce-
roses and hippopotami, prevailed and was adopted even by
Leidy in this country. Richard Owen, by far the greatest
man after Cuvier, made a decided advance, and, as in the
classification of the fishes and reptiles, was the direct pre-
decessor of Cope. He defined the new mammalian orders,

LIFE AND WORKS OF COPE. xxix

Marsupialia and Toxodontia, but he especially broke down
Cuvier’s classification of the Ungulates by distinguishing
the Perissodactyla from the Artiodactyla upon the basis of
foot structure, the importance of which Cuvier himself had
only dimly perceived.

In this country the earlier contributions of Jefferson,
Harlan, and Gibbes were overshadowed in the mid-century
by the numerous valuable works of Leidy, who became at
once the founder of American Vertebrate Paleontology,
although he entirely lacked the philosophical spirit either
in anatomy or in evolution. Thus, from all this long post-
Cuvierian period an immense number of facts issued, but
only two generalizations, the first of what may be regarded
as the great laws or principles in the evolution and classifi-
cation of the Mammalia. These laws are as follows:

I.—The Law of Brain-Growth.—This principle, that the
older Mammalia had smaller brains, and that in order of
succession there was a steady increase in brain size, was
enunciated by Lartet, and has been subsequently elaborated
and demonstrated by Marsh.

Il.—The Classification of the Hoofed Animals by Foot
Structure—This was discovered by Owen in his division
above alluded to, which first directed attention to the im-
portance of differences in the feet.

The three vertebrate paleontologists of the new period
who responded most fully to the Darwinian movement
were Huxley, who unwillingly entered the field, but soon
found an opportunity of overthrowing Cuvier’s Law of Cor-
relation. Huxley’s greatest generalization was the central
position of the order Insectivora. He had few opportunities
of working upon fossil mammals. He erroneously placed
Paloplotherium instead of Hyracotherium in the horse line,
and erroneously supported Reichert’s theory of the homol-
ogy of the quadrate. Cope and Marsh alike responded to

xxx LIFE AND WORKS OF COPE,

the Darwinian impulse. In Russia appeared Waldemar
Kowalevsky, who had a short but brilliant career in Mam-
malian Paleontology. He announced the third great
principle :

III —Law of Adaptation of Foot Structure in Ungulates by
Reduction, Accompanied by Shifting of the Metapodials.—
Kowalevsky’s ancestral type of ungulate or Protungulate,
like that of Huxley, was believed to possess five digits:

In the mean time the gifted John A. Ryder, of Philadel-
phia, was attacking the mechanical evolution of the feet
and teeth.

Cope, who had practically entered Mammalian Paleon-
tology in 1870, found a great field of facts lying fallow
before him, with the three above principles as a means of
interpretation. Keen to wed philosophy with anatomy, in
1873 he added to the generalizations of Huxley and Kow-
alevsky the additional principle :

IV.— The Ancestors of the Hoofed Animals possessed Buno-
dont or Hillock-like Teeth.—This prophecy was speedily veri-
fied by Wortman’s discovery of Phenacodus. This discovery
led Cope on to a re-classification of the entire group of Ungu-
lates by foot structure—the logical outcome of the move-
ment in which Owen, Kowalevsky, Huxley, Ryder, and
himself had participated. This centered about the following
principle:

V.—The Law of Taxeopody. That the primitive feet of
Hoofed Animals were Plantigrade, like those of the bear, with
serial unbroken joints—Thus he proposed in the early
eighties the four new Orders, two of which have been per-
manently adopted into Paleontology:

Cope. Marsh.
Taxeopoda. Protungulata.
Amblypoda. Amblydactyla.
Condylarthra. Holodactyla.

Diplarthra. Clinodactyla.

LIFE AND WORKS OF COPE. Xxxi

Kowalevsky, in 1873, had pointed out the significant
articulations of the metapodials; Cope here showed the
still greater importance of the mutual articulations of the
podials, firmly establishing thereupon the orders Condyl-
arthra and Amblypoda, uniting Owen’s Perissodactyla and
Artiodactyla into the Diplarthra, and by hypothetical
phyla connecting the Proboscidia and Hyracoidea with a
still-to-be-discovered plantigrade, bunodont stem, the “ pro-
tungulate” of Huxley and Kowalevsky. These generaliza-
tions, despite errors of excess and of detail which Riitimeyer
and Osborn have pointed out, constituted the first distinct
advance in mammalian classification since Owen demol-
ished Cuvier’s ‘“‘ pachydermata ;” they rank with Huxley’s
best work among similar problems, and afford a basis for
the phylogenetic arrangement of the hoofed orders which
has been adopted by all American and foreign paleontolo-
gists. Having thus raised the feet, a region of the body so
long neglected by the followers of Cuvier (with the excep-
tions noted), to a position of prime importance in classifica-
tion, it was his good fortune to discover in the collections
from the Puerco or Basal Eocene the following law:

VI.—Law of Trituberculy. That all Types of Molar Teeth
in Mammals originate in Modifications of the Tritubercular
Form.— It became apparent that the hoofed mammals had
sprung from clawed ancestors, but the Wasatch period was
too remote from the parting to furnish conclusive evidence.
This evidence came in a flood from the underlying Puerco
fauna, the systematic treatment of which constitutes the
most unique section of Cope’s work among the extinct
mammalia. From this material originated the above
great generalization—namely, that the primitive pattern
of the molar tooth consists of three tubercles. Around this
trituberculy centers the whole modern morphology of the
teeth of the mammalia and the establishment of a series of
homologies in the teeth of most diverse types, wholly un-

XxXxil LIFE AND WORKS OF COPE.

suspected in the ‘‘ Odontologies” of Cuvier and Owen, con-
necting the most ancient Mesozoic mammals with the most
modern and specialized types, and applying even in the
teeth of man. The forceand application of the tritubercular
law Cope clearly perceived, but left to others fully to work
outand demonstrate. It promises ultimately to give us the
key to the entire phylogeny of the mammalia, extending to
every division of the Marsupialia and Placentalia, and will
probably be found among the Monotremata.

Thus the final philosophical working basis for the evo-
lution of the hoofed, as well as the clawed, animals has been
gradually established, for, as Professor Marsh observes in
his monograph on the Dinocerata, ‘the characters of most
importance in the evolution of the Ungulates are the teeth,
the brain and the feet.’

It now only remained for Cope to take another step.
beyond Huxley and Kowalevsky, and, aided by fortunate
discoveries in the field, he demonstrated that the ancestors
of the hoofed animals were clawed animals, establishing
the Seventh Law:

VII.— The Hoofed Orders Converge towards the Clawed types
of Creodonta and Insectivora.

So much for the great generalizations which establish
Cope historical position in Mammalian Paleontology. These
are the mountain peaks, the points where exploration and
discovery were followed by happy inspiration in a chain of
contributions which includes his exposition of the faunal
succession of the Mammals from the base to the summit of
the Tertiary, as well as two or three discoveries of great
interest in Cretaceous. His most conspicuous work relates.
to the Puerco, with its extremely primitive hoofed and
clawed animals and primates. Here he established the
existence in this country of the Plagiaulacide and defined
the order Multituberculata. That from the Wasatch is.
perhaps next in value, and in succession rank his contribu-

LIFE AND WORKS OF COPE. XXXiil

tions from the John Day, Loup Fork, Blanco, Palo Duro and
Port Kennedy Bone Cave.

Asan explorer he had marked success, finding the unique
skeleton of Hyrachyus, of Loxolophodon, a name which was
telegraphed to the American Philosophical Society, and
converted by the operator into Lefalophodon. In the
Bridger, Cope himself found the lower jaw of Anaptomor-
phus, a little monkey with a dental formula like that of
man, which, owing to its extreme antiquity, occasioned
him a greater surprise than any discovery he ever made.
He also found the last of the great race of Uintatheres at
the top of Washakie Mountain of Central Wyoming. We
owe to him alone our knowledge of the scanty Wind River
fauna. From the White River Oligocene his materials were
poor and his work less satisfactory. From the rich Upper
Oligocene, with the assistance of Wortman, he secured fine
collections and has especially enriched our knowledge of
the Anchitheriide, Felide and Canide. From the Upper
Miocene Deep River and Loup Fork beds he has practically
contributed all that we know, especially of the Rhinoce-
roses, Horses, Mastodons, Camels, and other ruminants and
carnivora. Of the latter fauna his most complete papers
were upon the evolution of the Oreodontide. His latest
contributions to our knowledge of the fossil Mammalia
were upon the fauna of the Blanco and Palo Duro, or Good-
night beds of Texas, and the rich cave fauna from Port
Kennedy, Pa., brought together by his warm friend, Dr. H.
C. Mercer. It was his intention to cover the entire later Ter-
tiary in a second part of the “Tertiary Vertebrata ;’ many of
the plates and much of the MSS. of which volume are ready.

The “ Tertiary Vertebrata,’ Vol. III., of the Hayden
quartos, published in 1883, is his most imposing contribu-
tion to paleontology, including his studies of all the verte-
brate fauna of the Tertiary lakes west of the Rockies. This
work of over a thousand pages and eighty plates is said to

XXXIV LIFE AND WORKS OF COPE.

have been the despair of the public printer, owing to the
constant additions made while in press. It extends from
the Puerco to a portion of the Lower Miocene fauna. Be-
sides the full description and illustration of the great hoofed
orders above alluded to, it contains the full exposition of
the characteristic forms of Creodonta, an order of primitive
carnivora, which, as we have seen, he separated from the
Marsupialia in 1875, and in which he placed six families of
mammals from different parts of the world.

Before leaving the mammals it is fitting to speak of his
work upon “ kinetogenesis,” or the mechanical origin of the
hard parts of the body, especially of the teeth, vertebree, and
limbs. An invaluable paper by his friend and later col-
league, Ryder, put him upon this line of investigation, the
results of which he published in a long series of papers, cul-
minating in his memoir upon the “ Origin of the Hard Parts
of the Mammalia,” and in his collections of essays in the
“ Origin of the Fittest” and “ Primary Factors of Organic
Evolution.” One of his chief motives in these researches
was the demonstration, which he believed they afforded,
of the hereditary transmission of the effects of individual
efforts, use, and disuse; even if this motive is subsequently
shown to be an illusive one, by our future knowledge of the
real nature of evolution, these investigations lose little, if
any, of their intrinsic value. First, as in all his work, he
brings together an immense array of valuable facts and
observations ; second, he extends the principle of the inde-
pendent origin of similar structures ; third, he in most cases,
successfully establishes the actual mechanical adaptive or
teleological relations of the parts described; fourth, he
traces the course of phylogenic modification in a number of
important organs and thus establishes certain obscure
homologies, notably those in the teeth of Amblypoda,, Cory-
phodon and Uintathervwm.

As to his scientific character apart from his genius, which

LIFE AND WORKS OF COPE. xXxXxXV

is indefinable, we signalize his appreciation of the most
significant or diagnostic character in a group. Among his
fellow workers in the same field, whether upon the fishes,
amphibians, or mammals, he was quick to comprehend and
seize upon a strategic position. While others were plod-
ding on serenely in the description of facts, giving all an
equal value, Cope, with an eagle eye, would sweep down
upon some great distinctive fact and point out its supreme
importance. Thus he projected the Creodonta out of numer-
ous forms, such as Palxonictis, Hyxenodon, Arctocyon, which
had been discovered and studied for many years in France.
It is to be regretted that he did not more willingly surren-
der some of his own hypotheses. He clung to his er-
roneous mechanical explanation of the origin of Ungulate
foot structure long after it had been disproved by the present
writer. Like all of us, perhaps, he loved his own hypo-
theses, and once observed in jest in regard to a fossil which
opposed one of his theories, ‘I wish you would throw that
bone out of the window.”

He was no respecter of authority per se. Even if some-
times mistaken his fearless criticisms were chiefly animated
by high ideals and readiness to change the existing order of
things. He was full of cheer and determination when
things looked most unpromising, allowing nothing to dis-
turb the composure which is so essential to research. His
life, in fact, became a fine illustration of the happiness
attendant upon plain living and high investigation which
he foresaw at the early age of nineteen.

In this introduction the writer has drawn freely upon a biography
published in Science, also upon valuable materials which have been
furnished by Professor Theodore Gill, Cope’ lifelong friend, and
by Professors Baur and Dean. Professor Gill has revised the
proofs relating to Fishes ; those relating to Mammals have been
revised by the writer. General editorial revision has been made by

Dr. J. Perey Moore.

Syllabus of Lectures

ON THE

VERTEBRATA

CONTENTS.

INTRODUCTION

Supercntass I. HEMIcHoRDA
Class I. Helminthophya
Superctass II. Urocuorpa

Class I. Tunicata
Superciass III. CEpHALOCHORDA
Class I. Acrania.

Superciass IV CRANrIATA

Class I. Agnatha
Subclass I. Ostracophori
Subclass II. Cycliz
Subclass III. Marsipobranchii

Class II. Pisces
Subclass I. Elasmobranchii
Subclass II. Holocephali
Subclass III. Dipnoi
Subclass IV. Teleostomi

Class III. Batrachia
Subclass I. Stegocephali
Subclass II. Urodela
Subclass III. Salientia

Class IV. Monocondylia
Subclass I. Reptilia
Subclass II. Aves

Class V. Mammalia
Subelass I. Prototheria
Subclass II. Eutheria .

PAGE

21
28

52
D2

102
102
104

PREF ACE.

The present book is a corrected and extended edition of
The Syllabus of Lectures on Geology and Paleontology ;
Part III, Paleontology of the Vertebrata, published July
8, 1891. It was originally designed for use in the Exten-
sion Lectures of the University of Pennsylvania, and I
have used it as a text-book for the undergraduate classes in
the School of Biology of the University. The system
adopted is, with some minor exceptions, that which was
first presented by the author in the American Naturalist of
October, 1889. This was based upon original research on
the recent and extinct forms, with the exception of the
part relating to the Aves. This was taken, with little
modification, from Stejneger.

E. D. COPE.

University of Pennsylvania,
Philadelphia, 1897.

INTRODUCTION.

In the following pages the attempt is made to bring to-
gether the information which we possess as to the charac-
ters of the divisions of the Vertebrata above families which
are available for the determination of their relations by the
paleontologist. These characters, which are of necessity
those of the hard parts, must be of the first importance to
the discovery of the phylogenies, since the soft parts are
unavailable. It is, however, true that the relations of
these are close enough to render our inferences from the
former generally safe. Fortunately, also, the living rem-
nants of extinct groups are sufficiently numerous to enable
us to check our studies of the osteology. Thus we have
the Branchiostoma, the lampreys, the Ceratodus and Lepi-
dosiren, the Sphenodon, and the Monotremata, to which'‘to
refer when we desire to learn approximately the characters
of the soft anatomy of ancient forms.

All of the characters of the various divisions are not
given. In fact, when all extinct forms come to be known,
no divisionis likely to be defined by more than one character.
At present several characters may be often ascribed to
various divisions, but one of these will ultimately prove to
be the essential one. It is the object of the present synop-
sis to bring these definite characters into prominence ;
hence they are always stated first. The method of keys is
adopted as the most perspicuous method of exhibiting
them.

We are embarrassed in the endeavor to present the rela-
tions of the earliest and the lowest Vertebrata by a want
of knowledge of their structure, and by the absence from
our collection of numerous intermediate forms which must
have existed. Until our knowledge is more complete, the

8 : COPE

arrangement, especially of the contents of the class Agnatha,
must be regarded as largely provisional.

The ossification of the skeleton of the Vertebrata has
developed first on the exterior of the head and body and in
the sheath of the chorda dorsalis, and has then penetrated
inwards. The limbs have preceded in time the arches
(scapular and pelvic) to which they are, in the higher
forms, attached. Hence we find in such genera as Cepha-
laspis and Bothriolepis pectoral limbs without a scapular
arch, but with merely dermal ossifications to which they are
attached. This is parallel to the general absence of most of
the pelvic arch in fishes. The limbs themselves are sup-
posed to be radial ossifications in primitive longitudinal
folds of the body integument, some of which remain in
large part, as the dorsal fin of various fishes; while more
frequently but few of the radii remain, as in the limbs of
most Vertebrata. (Fig. 1.)

The use of this syllabus presupposes a knowledge of the
rudiments of vertebrate anatomy. The subject is presented
in zoological order, but tables are given, in which the tax-
onomic divisions are represented in their stratigraphic
position and succession.

BF An AF
Fia. 1.—Diagrammatic representation of primitive and derivative types of lateral
and median fins. .4, primitive condition, fins continuous; B. derivative condition,

fins distinct and specialized. D, dorsal fin; BrF, pectoral; BF, ventral; AF, anal;
SF, caudal; RF, dorsal; /F, second dorsal fin. From Wiedersheim.

VERTEBRATA 9

The branch Vertebrata* is divided into the following
superclasses :

No skull nor skeleton; notochord short,
anterior; nervous center a longitudi-
nal cord; Hemichorda.

No skull nor skeleton; notochord caudal
only; nervous center a ganglion; Urochorda.

No skull; notochord extending through-
out the body, included in a membran-
ous sheath, as is the cord-like nervous
axis above it; Cephalochorda.

A cartilaginous or bony skull and skeleton,
which extends throughout the body;
central nervous system a longitudinal
cord terminating in a brain within the
skull; Craniata.

Superclass I1.—HEMICHORDA.

There is but one class of Hemichorda:

Not metameric; no atrium; Helminthophya.

Class I—HELMINTHOPHYA.

Two orders of this class are known which differ as fol-
lows:

* The Vertebrata (or Chordata) as now extended, may be defined as celomate
Metazoa, possessing a longitudinal, axial, cellular supporting rod (the notochord),
derived from the hypoblast (except Hemicborda ?) and in the mesoblastic sheaths
of which the vertebral centra form in the higher types; with the walls of the
pharynx perforated by openings (gill slits, visceral clefts, stigmatz), placing its
lumen in communication with the exterior; with an epiblastic central nervous
system which is in whole or in part tubular, which lies dorsal to the notochord and
which is not perforated by the functional mouth ; and with the mesoblast segmented
(obscurely in Urochorda). In the different groups any of these characters may be
persistent, or temporary and embryonic, or adaptively modified and obscure.—ED.

10 COPE

Respiration pharyngeal; no external bran-
chie ; alimentary canal straight, travers-
ing the body lengthwise ; Enteropneusta.

Respiration wholly or in part performed by
external branching processes; alimen-
tary canal bent upon itself, anus at oral
extremity ; Lophopneusta.

The order of Enreropneusta includes but one family,
the Balanoglosside, which consists of animals of worm-
like form, which burrow in the soil of the seacoasts of
most continents between high and low tide. They have a
larva resembling that of the Echinoderma.

The order of LopHopneusta includes sessile, and so far
as yet known, compound forms, which inhabit the ocean
bed, attached to solid substances. There are two families,
the Cephalodiscide with gill-slits, and the Rhabdopleuride
in which gill-slits are wanting.

No extinct forms of Helminthophya are known.

UROCHORDA.

Superclass II.

There is but one class of Urochorda:

Not metameric; a mantle covering the body ;
respiration pharyngeal; heart distinct,
saccular ; Tunicata.

Class IL.—TUNICATA.

There are three orders of Tunicata as follows:

Cloaca posterior; caudal appendage per-
sistent. Individuals simple, free ; Perennichordata.

Cloaca posterior; caudal appendage ab-
sent or caducous; protostigmata un-

VERTEBRATA 11

divided ; mostly pelagic ; Thaliacea.

Cloaca dorsal; caudal appendage caduc-
ous; protostigmata divided into rows
of secondary stigmata ; mostly fixed; Ascidie.

The Tunicata have sac-like bodies, and are marine in
habitat. Some of the Tunicata are free-swimming, and
these may consist of single or conjoined individuals. Others
are attached, some by a peduncle and others sessile, separ-
ately or in colonies. No extinct species of Urochorda are
known. ‘The Thaliacea and Aseidie are degenerates from
the vertebrate line. The families are:

PERENNICHORDATA: Appendiculariide.
THaLiacea: Salpidee, Pyrosomide.

Ascip1a: Botryllide; Ascidiidee; Clavellinide.

Superclass If].—CEPHALOCHORDA.

The only class of the Cephalochorda is the following :

Walls of the body muscular myotomes; no
jaws nor extremities ; pharyngeal walls
fissured; heart a longitudinal vessel,
which gives off branchial vessels which
unite into an aorta; a liver and vena
cava present ; Acrania.

Class I—ACRANIA.

Of this class but one order is yet known:

Pharyngeal fissures enclosed externally by a
fold of the integument, which encloses a

12 COPE

chamber (atrium) which opens inferiorly ;
openings of alimentary canal at opposite
extremities; heart tubular ; Leptocardii.

The species of the only family, the Branchiostomide (the
lancelets), are of compressed, worm-like form, and burrow
in the soil of the shores of all oceans. No fossil remains
of them have been yet found.

Superclass IV.—CRANIATA.

The five classes of the Craniata are defined as follows:

I. No lower jaw nor pectoral arch.

Internal skeleton not ossified ; Agnatha.

II. Lower jaw and pectoral arch present.

A, Basicranial axis not ossified, supported by a
parasphenoid bone; vertebral column consist-
ing chiefly of intercentra; ribs myotomic ;
ANAMNIA.

Limbs represented by many-radiate fins,
which are also present on the median
lines of the body; a coracoid bone;
heart with two chambers; one or no
occipital condyle; no internal nares ; Pisces.

Limbs consisting of one basal element, two
propodials, and metapodials and digits;
no median fins; lower jaw attached to a
suspensorium, complex; no opercular
bones; a coracoid; heart with three
chambers; two occipital condyles; inter-
nal nares ; Batrachia.

AA, Basicranial axis ossified; no parasphenoid ;

VERTEBRATA 13

ribs intermyotomic; vertebral column consist-
ing chiefly of centra; an amnion and al-
lantois; AMNIOTA.

Limbs as in Batrachia; one occipital condyle;
a suspensorium of the lower jaw;
mandible segmented; a coracoid bone;
ankles between first and second rows of
carpal and tarsal bones; heart with three
or four chambers ; Monocondylia.

Limbs as in Batrachia; two occipital condyles;
no suspensorium of the lowerjaw ; man-
dible not segmented ; coracoid generally
not distinct; ankles between propodial
bones and carpus or tarsus; heart with
four chambers ; Mammalia.

The period of appearance and duration of each of these
classes is exhibited in the accompanying table. It will be
observed that the order of appearance corresponds with the
natural order of structural complexity, the simplest appear-
ing earliest and the most complex last. It must be
mentioned that it is probable that each of the classes ap-
peared a little earlier than the time assigned them in this
table. The discovery of the fossil remains is regarded as
the first positive indication of the presence of a species of a
given class. Spines uncertainly referred to sharks (Pisces)
have been found in the Middle and Lower Siluric in
Europe and North America respectively. Foot impressions,
probably of Batrachia, have been found in the Carbonic at
lower horizons than the bones. ‘Tracks, probably of birds,
have been found in the Trias. A fore limb, probably of a
mammal, has been found in the Permo-Triassic Karoo
formation of South Africa.

The geological range of these classes is as follows:

14 COPE

Agnatha | Pisces | Batrachia a oa Mammalia

Plistocene

Neocene

Tocexze .

Cretacic

Jurassic

Triassic

Carbonic

Devonic

Siluric

Ordovicic .

Cambric . .

Huronic

Class I—AGNATHA.

The known members of the class Agnatha are a very
small representation of those that once existed; and they
present a great variety of characters, having little affinity
with each other. Three subclasses are most distinctly in-
dicated :

An osseous dermal skeleton with lateral

limb-like appendages; Ostracophori.
An osseous axial skeleton; no dermal

skeleton; ?appendages; Cycliz.

VERTEBRATA 15

No osseous skeleton nor limb-like append-
ages; Marsipobranchit.

Of the Agnatha, extinct species of the subclasses Ostraco-
phori and Cyclie only are known.

Subclass I.—OSTRACOPHORI.

There are three orders of this subclass, as follows. In
none of them are nostrils present:

No pectoral appendages; exoskeleton in three
layers,—the internal lamellar, the middle
cancellous, the external vaso-dentine ;
no bone-corpuscles ; Heterostraca.

No pectoral appendages; exoskeleton in three
layers with bone-corpuscles, the middle
layer with vascular canals; Osteostraca.

Pectoral appendages; exoskeleton osseous;
with sensory grooves; Antiarcha.

The time of appearance and range in time of these or-
ders is displayed in the following table. They are all
Paleozoic.

Heterostraca. | Osteostraca. Anutiarcha.

Carbonic . ‘ Be ce |

Devonic. . .

Siluric

Ordovicic . . .. ame |

Cambric .

16 COPE

Fie. 2.—Bothriolepis canadensis Whiteaves; bead and carapace, 11um above, aud head from
below. From the Devonic of New Brunswick. From Whiteaves. (Antiarcha.)

The Ostracophori are the oldest known Agnatha, and
the oldest Ostracophori have been found in the Siluric, in
the Salina of Pennsylvania and the Ludlow rocks of Eng-
land, which have produced Heterostraca. The Osteostraca
appear first in the Ludlow beds in England, and extend to
the Lower Devonic (Old Red Sandstone). They appear
first in America at the latter horizon in New Brunswick.
The Antiarcha commence in the Old Red, and continue
into the Chemung of the Lower Carbonic system. The
principal genus of the Heterostraca is the Pteraspis; of
the Osteostraca is the Cephalaspis; and of the Antiarcha
is the Pterichthys. (Fig. 2.)

Subclass II.—CYCLIA.

Median diphycercal caudal fin; brain case
not segmented ; Saccocephala.

The order SaccocerHata is the only one known to per-

VERTEBRATA 17

tain to the Cyclie. It embraces the simple family of the
Paleospondylide based on the genus Paleospondylus,
which has been, so far, only found in the Carboniferous beds
of Scotland.

Subclass III -MARSIPOBRANCHII.

This subclass has two orders:
Branchial fissures communicating directly
with the pharynx; nasal sac perforating
the palate; Hyperotreti.
Branchial fissures communicating with a
common branchial passage which opens
into the pharynx; nasal sac not perforat-
ing palate; Hyperoarti.

To the Hyprrorreti belong two families, the Myxinidee
and the Bdellostomide; to the Hyprroarrt one, the Petro-
myzontide.

No extinct species of Marsipobranchii are known. They
include only the recent lampreys and hags.

Class [.—PISCES.

This class is divided into four subclasses :

I. No maxillary arch.
A. No dermal cranial ossifications nor opercular
bones; claspers present (in living forms).

A hyomandibular; palatopterygoid arch
distinct; Elasmobranchii.

No hyomandibular; palatopterygoid arch
fused with chondrocranium ; Holocephali.
AA. Dermal cranial ossifications and opercular

bones present; no claspers.

Suspensorium fused with chondrocranium ;
palatopterygoid arch free; Dipnoi.

18 COPE

II. A maxillary arch present.
Dermal cranial bones and opercula present;
no claspers; suspensorium distinct
from cranium; palatopterygoid arch
free ; Teleostomt.

With our present knowledge we find the first appearance
of these subclasses to be contemporary ; 7. ¢., in the earlier
part of the Devonic system, in the Corniferous Limestone
and its equivalents in other parts of the world. Spines,
possibly belonging to sharks (Onchus), have been found in
Lower Siluric (Clinton) in Pennsylvania, and the Upper
Siluric (Ludlow) in England, but the animals to which
they belong may not have been Pisces. The Devonic forms
referred to the Dipnoi (the Arthrodira), while probably be-
longing to that subclass, are not yet positively proven to be
such. Representatives of all four subclasses still exist.
The Elasmobranchii are seen in the sharks and rays; the
Holocephali in the Chimeras; the Dipnoi in the lung-
fishes; and the Teleostomi in the true fishes.

The primary divisions of the fishes, as above indicated,
are seen in the structure of the skull. After this the
structure of the fins demands attention. Some of the pe-
culiarities of fin structure are common to all the members
of a subclass, while others characterize subdivisions ‘of the
same. These members consist of a dermal portion and a
skeletal portion. ‘The former is external to the body-walls;
the latter is internal to those walls in the case of the median
fins, and external in the paired fins, when present. The
skeletal portion consists of two sets of segments. First
those to which the dermal rays or folds are attached are
called the basilars. These are articulated to a second set,
the axials. The segments are then termed the baseosts and
axonosts. In the paired fins the latter are articulated di-
rectly with the elements of the shoulder and pelvic girdles.
Of the latter only the pubic element is present in Teleostomi.
In the shoulder girdle we have elements termed scapula,

VERTEBRATA 19

coracoid, clavicle, and epiclavicle. There is sometimes an
interclavicle. The dermal portion of the fins may include
as supports certain “ fin-rays,” or it may include only hair-
like rods or “actinotrichia.” The latter characterize
primitive types. In the case of the median fins it is char-
acteristic of primitive fishes to have the baseosts and
axonosts articulated with the neural spines of the vertebrie.
In modern fishes the axonosts are not so articulated, and do
not correspond with them, while the baseosts are rudi-
mental or wanting. (Fig. 3.)

Ac ! H | .

Cee ye vx
ree Fe a
ar it Yea) el)

ial

Fiac.3.—Diagrams of actinophores; 4, entotetramerous; 5, ectetramerous; C, ),
ectrimerous; H#, ecdimerous; Ac, actinotrichia; Fr, fin-rays; ba, baseost; Ax. axv-
nost; Jc, intercentrum.

Remarkable modifications take place in the successive
evolution of the caudal fin. The primitive condition of the
vertebral column in the tail of a fish is straight, with spines
equally divergent above and below. This is the diphycercal
tail (Fig. 4, A). It persists in some modern fishes, e. g.,
the eels. In the next stage the axis is turned up at the
end, and the spines of the inferior side are spread out fan-
like, while those of the superior side are crowded together.
The dermal part of the fin may develop an angle on the
inferior spines (heemal spines), the result being a two-lobed
fin, in which the upper lobe is much larger than the lower.

au COPE

This is the heterocercal tail (Fig. 4, B, C,and D). In later
forms the hemal spines grow larger and the neural spines.
smaller, and the axis grows shorter by the abortion of the
terminal centra. As a result, the lobes of the caudal fin
become equal. In the most fully developed types the heemal
spines fuse into a fan-shaped bone which supports the fin..
This is the homocercal caudal fin (Fig. 3, E). In its growth
it generally passes through the diphycercal and heterocercal
stages before reaching maturity.

Ly
A UNREAL
MMM prs sq RUTTEN

ep mi gue py
<a
ge
we
hy

reranERTO

Ww.
SN

VERTEBRATA 21

Fig. 4.--Tails of fishes; A, diphycercal tail of Polypterus; B, dlphyheterocercal tail of Cocco-
steus; C, caudal fin of a shark (Centrina); D, end of vertebral column of trout (Salmo fario) ;
#, end of vertebral column of barbel (Barbus fluviatilis) ; ep, Epural; hy, hypural bones.

Subclass —ELASMOBRANCHII.

The sharks are divided into three orders, as follows:

Paired fins ptychopterygial; with axonosts
%

22 COPE

(metapterygium) enclosed in body-wall
and basilars only free; Acanthodit.

Paired fins archipterygial; unibasal; a basi-
occipital element ; Ichthyotoms.

Paired fins pluribasal (tribasal); no basioccip-
ital element; Selachwi.

The order AcanTHopiI presents the most primitive type
of paircd fins known in the Pisces. The vertebral axis
is notochordal, and the fins do not display any actino-
trichia or rays. The males have no claspers. Tail hetero-
cercal. In the known genera the fins are supported by a
large spine in the anterior border of each; and the integu-
ment is covered with small quadrate granules, which also
extend over the fins. The teeth are absent or minute, or
consist of a single compressed triangular cusp, with smaller
laterals.

Wn UN

OU

SOG NX \X AAANAN “i

Fie. 5.—Pleuracanthus lucius Ag. From the Coal Measures of Alsace. From Sauvage.
The Acanthodide have but one dorsal fin, while the Isch-
nacanthide and Diplacanthide have two. In the Dipla-
canthidz only have clavicles been observed; in the other

VERTEBRATA 23

two families they are wanting. Among Diplacanthide
the genus Climatius is remarkable in having a series of
spines on each side between the ventral and pectoral
fins, which indicate the position of the primitive lateral
fold, from which the paired fins are supposed to have been
derived.

Diplacanthide are Devonic, as are also Ischnacanthide.
Acanthodide belong to the Devonic and Carbonic.

Fig. 6.—Pectoral and ventral fins of Cladoselache x4 (from Dean). B, axonosts (mesv-
and meta-pterygium) ; R, baseosts; D, dermal portion of fin.

In the Icurnyoromi the notochord persists, and the cen-
tra are represented by segments; the neural spines, axo-
nosts, and baseosts are continuous, and the dermal rays are
actinotrichia (that is, hair-like and more numerous than
the baseosts). There are two families, the Pleuracanthide
and the Cladodontide, which differ in the form of the pec-
toral fin. In the former they are biserially pinnate (Fig. 5),
while in the latter they are uniserially pinnate (branches
—baseosts—on one side of the axis—axonosts—only) (Fig.
6.) The Pleuracanthidz have teeth with two principal equal
cusps, while the Cladodontide have a principal median cusp,

24 COPE

with or without smaller lateral cusps. The Cladodontide
embrace the largest and most formidable species, and be-
long to the Carbonic period. The Ichthyotomi are all con-
fined to the Carbonic system.

The Sexacuiz (sharks and rays) present two lines of re-
lation, or suborders, which differ as follows:

Vertebree, when developed, having the con-
centric laminee predominating over the
radiating lamine ; anal fin absent ; Tectospondylt.

Vertebree when developed, with the radiat-
ing lamine predominating over the con-
centric ; anal fin present ; Asterospondyli.

In the Tectospondyli the majority of living types have
the body depressed, so that the branchial fissures are on
the inferior surface. This type is seen in the skates, saw-
fishes, and rays. In the Asterospondyli, on the other hand,
the branchial fissures are lateral. There are several
families of Tectospondyli, which appeared at different
periods of geological time. They are as follows:

I. Snout unarmed.

A, The crowns of the teeth closely overlapping
each other like shingles.

Edges of crowns forming a grinding face; (1) Petalodontide.
AA, The crowns of the teeth not overlapping.
f&. Summits of crowns forming a grinding sur-
face.

(2) Psammodontide.
(8) Rajide.
(4) Rhinobatide.
(5) Trygonide.
(6) Myliobatide (eagle rays).

8. Summits of crowns elongate cusps.

VERTEBRATA 25
Body not flattened ; (7) Spinacide.

II. Snout produced into an ensiform weapon toothed
on both sides.

(8) Pristiophoride.
(9) Pristidex (saw-fishes).

These families have the following range in geological
time:

Plistocene |

Neocene |

Eocene .

Cretacic .

Jurassic.

Triassic .

Cithoule | |

Devonic

Siluric .

Ordovicie

Cambric |

Huronic |

26 COPE

Fig. 7.—Heptanchus griseus left pectoral fin, pluribasal type; SB, scapular arch;
NL, foramen; Pr, propterygium; Ms, mesopterygium; a, d, axis of metapterygium;
Ra, basilars; FS, actinotrichia. From Wiedersheim.

The families of the Asterospondyli differ as follows:
I. Teeth molariform.*

Teeth separate ; (1) Cestraciontide.
Teeth confluent ; (2) Cochliodontide.

II. Teeth with elevated cusps.
(3) Hexanchidx (Fig. 7).
(4) Scyllitdee.
(5) Lamnidee.
(6) Carchariide.

The distribution of these families in time is as follows:

* Except some Cestraciontide.

VERTEBRATA Q7

m
te
ih
cs

Plistocene. .

Neocene

Eocene. . .

Cretacic

Jurassic

Triassic

Carbonic é |

Devonic . be Fe tas

Silurie . 2...

Ordovicic

Cambric .....

Huronic .....

From these tables it appears that the Selachii with grind-
ing teeth are prior in geological age to those with teeth
especially appropriate to a carnivorous diet. The latter
reached their greatest perfection, as well as size and num-
bers, in the Neocene system, some of the Carcharodons hav-
ing been probably seventy feet in length.

All the families of Elasmobranchii are common to the
Old and New Worlds.

28 COPE

Subclass II.—-HOLOCEPHALI.

But one order of this class is known:

A single external branchial fissure; actino-
trichia present; basilars, axonosts, and
neural spines articulating with each
other; pectoral fin pluribasal, with
three axonosts and numerous basilars;
ventrals with elongate axonosts and
basilars ; Chimeroidei.

In all known Cutmrorper the teeth are large paired
bodies, one pair in the lower and two in the upper jaws,
composed of coarse vascular dentine. These contain
columns varying in number and shape, consisting of coarse
tubes with calcareous walls which terminate on the masti-
cating surfaces. Notochord persistent, the ossifications,
when present, consisting of delicate rings, more numerous
than the neural arches. In the existing forms the males
have claspers like those of sharks.

Chimeroidei appear in the Corniferous Limestone in Ohio,
Wisconsin and Jowa, and in the Eifel Limestone in Rhine
Prussia. They are found in Triassic and Jurassic beds in
Europe, and abound in the Cretacic of Europe and North
America, and New Zealand. They extend through the
Cenozoic beds of Europe and North America, and a few
species still exist. The known families are the Ptychodon-
tide (of doubtful reference), the Squalorajide, Myriacan-
thidee, and Chimeeride, the last named only still remaining
alive, in four genera.

VERTEBRATA 29
Subclass III.—DIPNOI.

Two orders of this subclass are known, as follows:

Paired fins rudimentary
or absent; pelvic ele-
ments (so far as
known) double, lat-
eral; body more or
less protected by
plates ; Arthrodira.

Paired finsarchipterygial
(7. é., unibasal); pel-
vic elements fused
on the middle line;
body without bony
plates (Fig. 8); Strenoidea.

The ARTHRODIRA are represent-
ed by three or four families, the
Coccosteide, the Asterosteide, and
the Mylostomide, all restricted to
Fig. 8.~Ceratodus forstert the periods of the Devonic sys-

Krefft; pectoral fin, unibasal : 7 i
type; 4, 6, axial elements; t+, tem, Their remains abound in

radials; FS, actinotrichia. From

ba samedi these horizons of Europe and North
America. They possess, behind the head, a segmented
dorsal plate and a compound ventral plate or plastron.
The anterior lateral superior plates articulate with the
posterior lateral cranial elements by a hinge-like joint.
Notochord persistent. These forms have some resemblance
to the Agnatha Antiarcha, but they have a well-developed
mandibular arch, and distinct pelvic elements. ‘Their teeth

are processes of the edges of the jaws.

The SIRENOIDEA commence in the Carbonic system and
continue to the present day, when representatives of two
genera, Ceratodus and Lepidosiren, exist in the fresh

30 COPE

waters of the Southern Hemisphere. The families are
four, the Dipteride, Phaneropleuride, and Ctenodontide,
where the skull is covered with small tessellated plates, and
the Lepidosirenide, where the bones of the skull are few
and large. The Dipteridee and Phaneropleuride are con-
fined to the Devonic, and the Ctenodontide to the Car-
bonic. The Lepidosirenide commence in the Permian
(Texas) and continue to the present time. The teeth of
Sirenoidea are plates with radiating ridges, or else processes
of the jaws.

Subclass IV.—TELEOSTOMI.

There are four superorders of the Teleostomi or true fishes,
which differ in-the structure of the fins:

I. Median fins each with a single bone representing
axonosts.
Paired fins unibasal ; Rhipidopterygia.
II. Median fins with numerous axonosts.

All paired fins with baseosts; pectorals only
with a distinct peduncle formed of

axonosts ; Crossopterygia.
All paired fins with baseosts; neither with
distinct peduncle of axonosts; Podopterygia.

Axonosts not forming a peduncle; pec-
torals only with developed baseosts ; Actinopterygia.

Superorder RHIPIDOPTERYGIA.
The orders of Rhipidopterygia are the following. They
all have actinotrichia in place of fin-rays:
I. Paired fins with the basilars arranged on each side
of the median axis, or archipterygial.
Median fins with basilars ; Taxistia.

II. Paired fins with the basilars arranged fan-shaped
at the end of the short axis.

VERTEBRATA 31

Median fins with basilars ; Rhipidistia.
Median fins without, caudal fin with, basilars; — Actinistia.

The Taxistia includes but one family, the Holoptychiide,
which is of Devonic age.

The Rhipidistia includes the Tristichopteride, from the
Devonic and Carbonic; the Osteolepide, from the same;
and possibly the Onychodontide, which are Devonic.

The Actinistia includes the single family of the Ceela-
canthide, which appears in the Lower Carbonic and ranges
to the Upper Cretacic, in both Europe and America. (Fig.
9).
In all of the Rhipidopterygia the tail is either heterocercal
or diphycercal, and the chorda dorsalis persists. The scales
have a layer of ganoine, which extends also on the head.
The latter has a well-defined, persistent transverse suture
separating the parietal from the frontal elements.

Fig. 9—Undina penicillata Miinst, (Ccelacanthidz); one-third natural alae showing rhipidopterygiar and

actinistious types of fins; j. jugular plates; b, scales of Unidina acutidens, From Zittel.
Superorder CROsSOPTERYGIA.
The superorder CRossopreryGIA includes two orders, as

follows:

32 COPE

Baseosts and axonosts well developed; actino-
trichia; no fin-rays; pectorals? unibasal ; Haplistia.

Baseosts rudimental; fin-rays; pectorals
tribasal ; Cladistia.

But one family is included in the Haplistia, the Tarasiidee,
from the Lower Carbonic of Scotland. The Cladistia are
represented by a family which is not known in the fossil
state, the Polypteride of the rivers of Africa. The vertebree
in this family are ossified and biconcave.

Superorder Poporreryeia.

The superorder Popopreryeta has also two orders. They
are thus defined :

Branchiostegal rays present ; _ Lysoptert.

Branchiostegal rays absent ; Chondrostei.

In these orders the notochord is persistent, and there are
either actinotrichia, or fin-rays, which are more numerous
than the baseosts. Tail heterocercal or diphycercal.

The Lysopteri includes four families, which differ as fol-
lows:

I. Tail heterocercal.

Teeth acute, external ; Palxoniscide.
Teeth obtuse, on palate and splenial ; - Platysomide.
No teeth ; ‘Chondrosteide.

II. Tail diphycercal.
Teeth present; scuta on body ; Belonorhynchide.

The Paleoniscide range from the Devonic to the Jurassic
inclusive; the Platysomide belong to the Carbonic ex-
clusively; the Belonorhynchide to the Trias; and the
Chondrosteide to the Jurassic.

The Chondrostei are degenerate representatives of the

VERTEBRATA 33.

Podopterygia. They are deficient in various normal ossifi-
cations, and have an additional series of membrane bones
in the middle line of the skull. The two families are the
Acipenseride, or sturgeons, and the Polyodontide, or
paddle-fishes. Both are represented at the present day in
the northern regions of both hemispheres, and both appear
first in geological time in the Eocene system.

Superorder ACTINOPTERYGIA.

In this superorder we have the finally specialized type
of the true fishes. This consists in the abbreviation of the
skeletal parts of the true fins,
so that the basilar elements
become sessile on thescapula.
(Fig. 10.) Coincidentally with
this result, the fin-rays of the
median fins become distinct-
ly developed and articulated
each with its corresponding
baseost or axonost. Fishes
of this superorder have, for
the most part, homocercal
tails, the superior lobe being
<5 contracted to the size of the
= inferior lobe, and the verte-

Coty Hea ‘ , :
ee at ye bree ossified; but in some of

Fre. 10,—Salmo farioL; leftshoulder- the lower and older types

girdle; Cm, posttemporal; D 1, epiclavi- ;

Ge; D, clavicle; D 2, postclavicle; S, both heterocercal tails and
scapula ; Co (Cl), coracoid; a, basilars ; :

L, scapular foramen ; HS, FS, fiin-rays. notochordal vertebree still re-
main. Diphycercal tails also continue well along the as-
cending scale.

The Actinopterygia fall into two tribes:
Ventral fins abdominal; a ductus pneumaticus;
no spinous dorsal fin; parietal bones not
usually separated by supraoccipital ; scales
usually cycloid ; Malacopterygia.

34 COPE

Ventral fins usually thoracic or jugular;
no- ductus pneumaticus; usually a
spinous dorsal fin; parietal bones us-
ually separated by the supraoccipital ;
scales usually ctenoid ; Acanthopterygia.

In the cartilaginous fin-rays and posterior ventral fins
the Malacopterygia more nearly approach the fishes of the
superorders considered in the preceding pages, than do the
Acanthopterygia. The persistence of the communication be-
tween the swim-bladder and the gullet (ductus pneumaticus)
is another indication of this affinity. Accordingly we find
representatives of the former in older periods than we do
the latter. They are abundant in the Jurassic and Cretacic
and later periods, while the Acanthopterygia are very rare
in the Cretacic, and are abundant first in the Eocene.

The orders of the Malacopterygia are the following:

‘I. Median fins with actinotrichia.

Basilars of median fins well developed;
notochord persistent ; (1) Docopteri.*

II. Median fin-rays equal to and articulating with
axonosts.

A. Metapterygium present ; anterior vertebree unmodi-
fied. (Holostei).

a, Vertebre complex, the pleurocentra
and intercentra distinct.

Anterior vertebree similar to others ; (2) Merospondyli.

aa, Vertebre with centra and intercentra complete
on part of the column at least ; amphiccelous;

Anterior vertebree similar ; (3) Halecomorphi.
aaa, Vertebree (intercentra) opisthoccelous.

* The position of this order is uncertain,

VERTEBRATA 85

Anterior vertebre similar; a precoracoid
arch and a coronoid bone ; (4) Ginglymodi.

AA. No metapterygium; anterior vertebre modified,
and with ossicula; vertebree amphiccelous. (Ostario-

phys).
3, A precoracoid arch.

7, No symplectic bone.

Anterior vertebre modified, and with ossi-
cula auditus; pterotic simple; parie-
tals not distinct ; (5) Nematognathi.

vr, A symplectic bone.

Anterior vertebree codssified, and with ossi-
cula auditus; pterotic simple; parie-
tals distinct ; (6) Plectospondyli.

88, No precoracoid arch.

Parietais present, distinct, not separated
by supraoccipital ; (7) Glanencheli.

AAA. No metapterygium; vertebre amphiccelous,
the anterior not modified and without accessory
ossicles. (Teleocephali).

3, Scapular arch suspended to cranium.
a, A symplectic.
y, A precoracoid arch.

Pterotic annular, including a cavity which
is closed by a distinct bone; parietals
distinct, not separated by supraoccip-

ital ; (8) Scyphophors.
Pterotic simple; parietals distinct, not sep-
arated by supraoccipital ; (9) Isospondyli.

Eight pectoral basilar bones, the external

36 COPE
pair opposite each other, all articulat-
ing with the scapula ; (10) Actinochiri.
vr, No precoracoid arch.

Pterotic simple; parietals separated by
supraoccipital ; baseosts distinct ; (11) Haplomi..

aa, No symplectic.
Baseosts fused into a single cartilage ; (12) Xenomi.

Pterotic simple ; parietals distinct, in con-
tact; a preoperculum and palatine
arch ; (13) Ichthyocephali.

88, Scapular arch free from cranium.
0, A symplectic bone.

Hyoid arches and pectoral baseosts de-
veloped ; (14) Holostomi.

06, No symplectic.

Opercular bones and five osseous branchial
arches, with ceratohyal ; (15) Enchelycephali..

Opercular bones and one osseous branchial
arch, and ceratohyal ; (16) Colocephali.

No opercular bones, nor ceratohyal, nor
osseous branchial arches ; (17) Lyomeri.

The families of the Malacopterygia are as follows:

Docopreri: Dorypteride.

MeErRosponDYLi: Sauropsidee; Pyenodontide; Stylodontide;
Spheerodontide ; Macrosemiide.

HatrecomorpHr: Amiidee (dog-fishes).
GineiyMopvI1: Lepidosteide (bony gars).

NemaToGNATHI: Siluride (cat-fishes); Hypophthalmide ;.
Aspredinide.

VERTEBRATA 37

PLecrosponpDyLI: Characinide; Sternopygide ; Cobitide ;
Cyprinide (carp); Catostomide (suckers).

GLANENCHELI: Gymnotide (electric eels).

ScyPHopHori: Mormyride; Gymnarchide.

IsosponDYLI: Dapediide ; Lepidotide ; Aspidorhynchide ;
Saurodontide (Fig. 11); Osteoglosside; Heterotide ;
Galaxiide; Clupeide (herring); Chirocentride ; Sal-
monide (salmon); Thymallide (grayling); Alepoceph-

alidee; Gonorhynchide; Sauride; Lutodiride; Aulo-
pide; Elopide; Albulide; Hyodontide; Notopteride.

AcTINOCHIRI: Erisichthide.

Hapromr: Esocide (pike); Stratodontide ; Umbride ; Cy-
prinodontide ; Amblyopside (blind fishes).

XeEnomi: Dalliide.
IcuTHyocePHALI: Monopteride.
Hotostom1: Amphipnoide.

ENCHELYCEPHALI: Nemichthyide; Anguillide (eels); Con-
grid (eels); Synaphobranchide ; Simenchelyide.

CoLocEPHALI: Murzenidee (eels).

Lyomeri: Saccopharyngide; Eurypharyngide.

22

can UME CLE
ARAN RRS ace CaN EKO =

Fic. 11.—Portheus molossus, Cope; isospondylous fish from the Upper Cretacic of Kan-
sas, one-twentieth natural size. Original.

The geological range of the orders of the Malacopterygia
is as follows:

COPE

38

o1uOIN Fy

oliqmes)

oLIn[IS

+ oTmoAay

oU0gIBy)

“+ OISSBITT,

* * OIssBIn er

D1IDBIJOIO

* + gua007y

* + 909008NT

LT

9T

ST

VL

tal

GL

mai

or

* 9u900}sI[ gq

VERTEBRATA 39

The orders of which no record is given are known only
in the living state. The Plectospondyli represent the
highest form of the Malacopterygia, while the Haplomi give
the connection with the Acanthopterygia through the Per-
cesoces of the latter. Tne Isospondyli offer the connection
with the Merospondyli. The Apodal line (the eels), in-
cluding orders Nos. 13 to 17, is a degenerate one, ending
in the greatly degenerate deep-sea forms of the Lyomeri.

The orders of the Acanthopterygia are the following :
A. Scapular arch suspended posterior to the cranium.

Maxillary bone distinct; no interclav-
icles; epibranchials and pharyngeals
present; inferior elements distinct; (1) Opisthomi.

AA. Scapular arch suspended to cranium by a post-
temporal bone.

a, Ventral fins abdominal.

Branchial arches developed, third su-
perior pharyngeal enlarged; gill-
fringes linear; no interclavicles; (2) Percesoces.

Epibranchials and superior pharyngeals
reduced in number; gill-fringes
linear; interclavicles present; (3) Hemibranchii.

Epibranchials and superior pharyngeals
wanting; gill-fringes in tufts ; (4) Lophobranchit.
ao, Ventral fins thoracic or jugular.
f, Anterior (spinous) dorsal fin expanded
into transverse lamin, sessile on cranium.
Cranium normal; (5) Discocephali.

88, Spinous dorsal fin not transversely ex-
panded.

40 COPE

7, Posttemporal projecting freely from
skull.

First vertebra united by suture to crani-
um; intercalaria united behind su-
praoccipital ; basilar pectoral bones
elongated ; (6) Pediculati.

Posterior cephalic region normal; the
anterior twisted so as to bring both
orbits on one side; inferior pharyn-
geals distinct; (7) Heterosomata.

Cranium normal, premaxillaries usually
codssified with maxillaries behind,
and the dentary with the articular;
pharyngeal bones distinct; (8) Plectognathi.

Cranium normal; bones of jaw dis- -
tinct; (9) Percomorphi.

77, Posttemporal an integral part of the skull.

Cranium normal; bones of jaws distinct;
pharyngeals separate; . (10) Craniomi.

The families of the preceding orders are as follows:

OpistHomi: Mastacembelide; Notacanthide.

Prrcesoces: Opheocephalide; Mugilidz (mullets); Ather-
inidee; Sphyreenidee (barracuda); Scombresocide (soft

gar).
HEMIBRANCHII: Pegasidee; Gasterosteide (stickle-back);
Fistulariide; Centriscidee; Amphisilide; Dercetide.

LorHosrancuir: Solenostomide; Syngnathide (pipe-
fishes); Hippocampidee (sea-horses).

DiscocePHALI: Echeneidide (remoras).

Pepicurati: Antennariide ; Lophiide (fishing frog).

VERTEBRATA 41

HeEtTeRosomMaTA: Pleuronectidee (flat-fishes).

PuecroGNaTHi: Triacanthide; Balistide (trigger-fishes) ;
Tetraodontide (bladder-fishes); Diodontide; Ostra-
clidee.

PEeRcoMoRPHI: (Anacanthini) Ophidiide; Gadide (cod);
Macruride; (Haplodoci) Batrachide (toad-fishes) ;
(Cyclopteroidea) Cyclopteride; (Scatophagoidea) Sca-
tophagide; (Epilasmia) Acronuride; Cheetodontide ;
(Rhegnopteri) Trichidiontide; (Distegi) Scorpeenide ;
Cottide (sculpins); Blenniide; Gobiide ; Platycepha-
lide; Rhamphocottide; Agonide; Heterognathide;
Gerreide ; Carangide’ (pompano); Sillaginide; Pristi-
pomatide; Scienide (maigres); Sparide; Percide
(perch); Berycide ; Scombridze (mackerel); Trichiuride ;
Xiphiadide ; (Labyrinthici) Osphromenide; Anaban-
tide; (Pharyngognathi) Embiotocide; Cichlide;
Labridee; Scaride.

Craniomi; Triglide (gurnards); Dactylopteride.

The table on the following page shows that no order of
the Acanthopterygia has become extinct. The Percomor-
phi display the greatest numbers and importance at the

present time. The Lophobranchii and Plectognathi are
degenerate types.

The geological distribution of these orders is as follows:

NotTEe.—The editor is indebted to Prof. Theodore Gill for several important
corrections of ambiguities in the definitions, and oversights in the arrangement of
the groups of fishes. None of these changes, however, affect the author’s views
on affinities, etc.

COPE

42

* ormomnyy

* o1lrquery

* OLMIS

dTUOAaCT

* o1uog.ABy)

OISSBLL,

orssean pe

o19v4AI,)

BUIV05F

9U9000 NT

or

9u900}sI[ J

VERTEBRATA 43
Class IIL—BATRACHIA.

There are three subclasses of Batrachia, as follows:

Basioccipital, supraoccipital, intercalare, and
supratemporal bones present; propodial
bones distinct ; Stegocephali.

Basioccipital, supraoccipital, and supratem-
poral bones wanting; propodial bones
distinct; no urostyle; Urodela.

Basioccipital, supraoccipital, intercalare, and
supratemporals wanting; frontals and
parietals connate; propodial bones con-
nate; lumbosacral vertebre united into
a urostyle; Salicntia.

Subclass I—STEGOCEPHALI.

There are four orders of Stegocephali.
a, One occipital cotyloid articulation.

Vertebral bodies represented by basal and
lateral elements (intercentra and centra); Ganocephali.

aa, Two occipital condyles;

Vertebree represented by distinct and incom-
plete intercentra and centra (pleurocen-
tra); atlas segmented ; Rhachitomi.

Centra and intercentra complete, making
two vertebral bodies to each neural arch; Embolomeri.

No centra; intercentra each supporting a
neural arch; Microsauri.

44 COPE

Fig. 12.—Trimerorhachis insignis Cope; parts of skeleton, natural size; a, occipital segment with
single cotylus ; 0, ec, posterior part of lower jaw; d, e, series of vertebra; d, side view, depressed ;
e, View obliquely from above, both showing the rhachitomous structure: 7, intercentra: p, pleuro-
centra. Original. From the Permian of Texas.

The Stegocephali are only known from the Carbonic and
Triassic systems. They are all more or less notochordal,
but a considerable range in this respect is found in the Mi-
crosauri. In some of them the notochord is cut off by
the vertebral ossification, while in others (Branchio-
saurus) the ossification is a mere sheath round the chorda.
In the Ganocephali and Rhachitomi the vertebral centra
are represented by a segment beneath each branch of the
neural arch, the pleurocentrum (Fig. 12, p); and these rest
below on a median inferior piece, the intercentrum (Fig.
12, 7). Inthe Embolomeri each of these segments is de-
veloped so as to form a disc, so that there are two vertebral
bodies to one arch. In the Microsauri the intercentra have
been fused with the pleurocentra, so that the body consists

VERTEBRATA 45

principally of the former, and the same structure is charac-
teristic of the remaining Batrachia. In the Reptilia, on
the other hand, the intercentra gradually disappear, being
represented in most of the types in the cervical and caudal
regions only. In the latter they support the chevron
bones.

Fig. 13—Trimerorhachis insignis Cope; skull. From the Permian bed of Texas.
Original,
Of the GANocEPHALI two families are known, the Tri-
merorhachide (Figs. 12-13), without, and the Archego-
sauride with neural spines of the vertebrae. They occur in

tho Coai Measures and the Permian of Europe and North
America.

46 COPE

The Racuitomi possesses but one family, the Eryopide,
which often reached a large size (Fig. 14). These are con-
fined to the Coal Measures. If the Labyrinthodontidee be-
long to this order they range also to the Trias inclusive, in
both continents.

The Empotomerr includes one family, the Cricotide,

NP
inp }
4 ‘ iy
he

ae “i

Fig. 14.—Eryops megacephalus Cope; skull, from above, one-fourth natural size.
From Permian bed of Texas. From Cope.

VERTEBRATA 47

which belongs to the Permian of Europe and North
America. (Fig. 15).

Fie. 15,—Cricotus heteroclilus Cope ; one-third natural size. From Permian of Texas,
a, head from aove; b, part of belly from below. From Cope.

The Microsauri embraces the following families: Bran-
chiosauridz ; Hylonomide ; Molgophide ; Phlegethontiide.
The smaller forms are mostly from the Coal Measures,
while some large ones occur in the Trias. Some of the
Hylonomide come from the Permian of North America and
Europe. They were mostly of small size, and their verte-
bre exhibit very various degrees of ossification. In some
the bodies are completely ossified, while in others the ossi-

48 COPE

fication forms a superficial layer. In the Labyrinthodon-
tide the ossification is more complete. The dentine may
be entire, or deeply inflected, so as to form straight or
labyrinthic folds.

Subclass II.—URODELA.

There are three orders of Urodela :
a, An os intercalare.
Palatine arch and vomer present; — Proteida.

aa, No os intercalare.

A maxillary arch and vomers; Pseudosauria.

No maxillary arch or vomers; Trachystomata.

Under the Prorerpa the only family known is the Pro-
teidee (Fig. 16.)

The Psrvposaurra embraces the following families:
Cryptobranchide; Amblystomide; Hynobiidee; Plethodon-
tide; Thoriidee; Desmognathide; Salamandride; Pleuro-
delide; Amphiumide; Ceeciliidie.

The TracHystomatra includes only the family of the Si-
renide.

A possible member of the Proteida occurs in the Coal
Measures, but certainly known members of the order are
not found in a fossil state. Of Pseudosauria fossil forms
are first found in the Laramie in America, and they are not
uncommon in the Neocene in Europe. Trachystomata are
not known fossil.

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50 COPE

Subclass III—SALIENTIA.

But one order of this subclass is recognized. It is thus
defined:

Vomers and palatopterygoid arch present; Anura.

The Anura has the families arranged under the follow-
ing suborders:
Eustachian foramina opening together on the
middle line; no tongue; coracoids con-
nected by a cartilage on each side ; Aglossa.

Eustachian foramina separate; a tongue; cor-
acoids connected by a separate cartilage
on each side, one overlapping the other ; Arcifera.

Eustachian foramina separate; a tongue; a
single median cartilage connecting all the
coracoids ; scapular arch free ; Firmisternia.

As in Firmisternia, but scapular arch artic-
ulated to skull ; Gastrechmia

The families included under these orders are as follows:

(Aglossa): Xenopide ; Pipide.

(Arcifera): Discoglosside ; Bufonide ; Dendrophryniscide;
Asterophrydide ; Pelodytide ; Seaphiopide ; Hylide ;
Cystignathide; Amphignathodontide; Hemiphractide.

(Firmisternia): Engystomide; Phryniscide; Dendroba-
tide ; Cophylide; Dyscophide; Colostethide; Ran-
ide; Ceratobatrachide.

(Gastrechmia): Hemiside.

Remains of Anura have been found in the Jurassic beds
of Colorado, but to which suborder they pertain is un-
known. They next appear in the Eocene of Wyoming.
Well defined forms are found in the phosphorites of France,
which include both Arcifera and Firmisternia. They con-
tinue to the present day.

The time relations of the orders of Batrachia are repre-
sented in the preceding table.

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52 COPE
Class IV.—MONOCONDYLIA.

There are two subclasses of Monocondylia:

Anterior limbs ambulatory, with numerous carpal
and metacarpal bones; two aorta roots; in-
tegument consisting partly of scales; Reptilia.

Anterior limbs volant, with the carpals and meta-
carpals more or less codssified and reduced in
numbers; integument consisting in part of
feathers; one aorta root; Aves.

Approximations between these subclasses exist at various.
points. Thus the Dinosaurian reptiles resemble birds in
the structure of the posterior limbs and pelvis; while
among birds the Saururee approach reptiles in the structure
of the manus and of the caudal vertebre. The definitions.
above given are, however, not violated.

Subclass I—REPTILIA.

The diversities in the orders of reptiles are seen chiefly
in the constitution of the posterior parts of the skull; the
shoulder-girdle presents a good many varieties, and the
limbs and vertebre exhibit others.

In order to understand the homologies of the elements
which make up the posterior region of the skull of reptiles.
generally, it is necessary to become acquainted with the skull
of the primitive order of the Cotylosauria, and the most nearly
allied subclass of the Batrachia, the Stegocephali. These
are represented in figures. In these types the temporal fis-
sure is crowned with an osseous roof which protects the
temporal muscles. This roof contains the elements of the
arches which extend from the orbit posteriorly to the sus-
pensorium of the quadrate bone. These elements are six
in number, and are named as follows, commencing with
those that lie next to the parietal and the frontal bones,

VERTEBRATA 53

the posterior of each pair being mentioned first: first
pair, supramastoid and postfrontal; second pair, squamosal
and postorbital; third pair, quadrato-jugal and jugal
(or malar). The appearance of two foramina in this
roof determines the presence of two arches, a superior or
postorbito-squamosal, and an inferior, or quadrato-jugal.
The presence of one foramen only, results in the develop-
ment of a single arch. Which of the two arches remains
will depend on the position of the foramen, and the de-
gree of excavation of the inferior edge of the temporal
roof. In the Plesiosauria this arch is the quadrato-jugal ;
in the Lacertilia it is the postorbito-squamosal. In
the Theromora it is postorbito-squamosal, but much of
the quadrato-jugal adheres to its lower border, as in the
Mammalia. In the Archosaurian series (with two post-
orbital bars) these elements are widely separated. In the
Testudinata parts of both bones may be combined into one,
or one only may be represented. The supramastoid bone
disappears as a distinct element early in the history of the
Reptilia. In the Theromora the quadrate bone grows
shorter as we approach the Mammalia, and coincidentally
the quadrato-jugal disappears, as for instance in the Cyno-

dontia (Seeley).
The quadrate bone is supported on a peduncle formed by

the transverse extension of the exoccipital, petrosal and
opisthothic bones. The opisthotic is generally early fused
with the exoccipital, but in the Ichthyopterygia and Testu-
dinata it is distinct, and takes the place of the petrosal as
a support of the quadrate in conjunction with the exoc-
cipital. In the Pythonomorpha a bone which occupies the
position of the terminal part of the opisthotic (or paroc-
cipital, which is the older name) issues from between the
exoccipital and petrosal, and supports the quadrate.
Whether this is homologous with part or all of the paroc-
cipital is an open question. For the present it is called, in

this book, the paroccipital, and it is probably a distinct

54 COPE

element from the opisthotic. In the Lacertilia it is ex-
cluded from the embrace of the petrosal and exoccipital,
and is carried on their extremity, and is in contact with
the quadrate. In the Serpentes, in consequence of the
loss of the arches, the quadrate is borne on the extremity
of this paroccipital, which generally projects freely from
the cranium.

The vertebre of Reptilia are either biconcave (amphi-
ccelous), or flat at both extremities (amphiplatyan), or ball
and socket. The latter is of two types: first, with the
socket in front and the ball behind or proccelous, which is
the most frequent; or second with ball in front and socket
behind, or opisthoccelous. Intervertebral articulations
other than those of the zygapophyses are found; as the
zygosphen and zygantrum, (Serpentes, Fig. 29, p. 81), and
the hyposphen and hypantrum (Dinosauri Saurischia, Fig.
22, p. 69), and some others.

In the scapular arch, a sternum, proscapula, presternum,
interclavicle and clavicle may be present or absent. Scap-
ula and coracoid are always present, and usually a pre-
coracoid.

Twelve orders of Reptilia are known:

I. The quadrate bone immovably fixed to the ad-
jacent elements by suture.

A. Scapular arch external to ribs; temporal region
with a complex bony roof; no longitudinal post-
orbital bars.

A tabular and supramastoid bones, anda
presternum; limbs ambulatory; ver-

tebree amphiccelous ; (1) Cotylosauria.

AA. Scapular arch internal to ribs; temporal region
with complex roof and no longitudinal bars.

A presternum ; limbs ambulatory ; (2) Chelydosauria.

VERTEBRATA 55

AAA. Scapular arch internal to ribs; sternum extend-
ing below coracoids and pelvis; one post-
orbital bar. :

No supramastoid ; a paroccipital; clav-
icle not articulating with scapula ; (8) Testudinata.

AAAA. Scapular arch external to ribs; one longitu-
dinal postorbital bar (Synaptosauria).

A supramastoid and paroccipital bones;
ribs two-headed on centrum ; car-
pals and tarsals not distinct in form
from metapodials; vertebree amphi-
ccelous ; (4) Icthyopterygia.

A supramastoid; paroccipital not dis-
tinct; a postorbito-squamosal arch ;
ribs two-headed; a clavicle ; obtu-
rator foramen small or none; ver-
tebree amphiccelous ; (5) Theromora.

*No supramastoid ; paroccipital not dis-
tinct ; a quadrato-jugal arch; scapu-
la triradiate ; no clavicle; ribs one-
headed ; (6) Plesiosauria.

AAAAA. Scapular arch external to ribs; two longitu-
dinal postorbital bars (paroccipital arch
distinct). (Archosauria).

a, A supramastoid bone.

Ribs two-headed; a clavicle and inter-
clavicle; acetabulum closed; no
obturator foramen; ambulatory ;
vertebree amphiccelous; (7) Pelycosauria.

aa, No supramastoid.

*Cope, Proc. Amer, Philos. Soc., 1894, identifies and describes a supra-mastoid in
the plesiosaurian skull.—ED.

56 COPE

Ribs two-headed ; interclavicle not dis-
tinct; external digits greatly elon-
gate to support a patagium for
flight ; (8) Ornithosauria.

Ribs two-headed ; no interclavicle ; ace-
tabulum open; ambulatory ; (9) Dinosauria.

Ribs two-headed ; an interclavicle; ace-
tabulum closed ; ambulatory ; (10) Loricata.

Ribs one-headed ; an interclavicle; ace-
tabulum closed, a large obturator
foramen ; ambulatory ; (11) Rhynchocephaha.

II. Quadrate bone loosely articulated to the cranium
and at the proximal end only. (Streptostylica).

No distinct supramastoid, nor opisthotic ;
one or no postorbital bar; scapular
arch external to ribs when present ;
ribs one-headed ; (12) Squamata.

It appears that four orders are Paleozoic in age, one of
which possibly continues into the Trias (Theromora).
Four are exclusively. Mesozoic, while four continue to
exist.

The Theromora present the greatest resemblances to the
Mammalia, while the Cotylosauria present resemblances to
the Stegocephalian Batrachia with which they agree essen-
tially in the structure of the temporal roof, in the presence
of the tabular and supramastoid bones.

The three plates which follow present diagrams of dorsal and lateral views of
all of the orders of Reptilia except the Chelydosauria. The bones are indicated by
a uniform system of abbreviations.—Ep.

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60 COPE

The time-histories of the orders of reptiles are represented
in the accompanying table :

Plistocene

Eocene .

Neocene |
Cretacic |
it

Jurassic |

Triassic . | | |
|

Carbonic | | |

|
|
|
|
|

|

|

Devonic |

Siluric

Ordovicic

Cambric

Huronic

The order CoryLosauria includes reptiles with amphiccel-
ous vertebre and of terrestrial habits. There are four families,
the Pariasauride, of robust form varying from the size of an
alligator to that of a cat, from the Permian beds of South
Africa and North America; the Diadectide (Fig. 17), with
hyposphen-hypantrum vertebral articulation, and teeth
with robust, molariform crowns, transverse to the jaws, from

VERTEBRATA 61

North America and Europe; the Pariotichidie, with several
‘Tows of teeth in the jaws, from North America; and the Elgi-
niide with very thin ossifications and light structure, from
the Trias of Scotland. This order is probably ancestral to all

F1G.17.—Empedias molaris Cope; inferior view of skull, one-third natural size.
A Cotylosaurian from the Permian of Texas.

other orders of Reptilia. The oldest reptile known is the
Isodectes punctulatus Cope from the Coal Measures of
Ohio. The South African forms Gorgonops and Proco-
lophum probably belong here, and are supposed to be types
of distinct families.

The CHELYDOSAURIA is an order of limited extent, with

62 COPE

present knowledge, as it includes but one family, the Oto-
celide, from the Permian formation of North America.
These reptiles possessed a carapace of transverse osseous
arches which extended across the back from side to side in
close contact. The anterior part of the scapular arch be-
low resembles the corresponding part of the plastron of a
tortoise. The temporal roof is excavated posteriorly for the
auricular meatus. The order is probably ancestral to the
Testudinata and the Pseudosuchia.

The order TesruprnaTa presents four subordinal modifi-
cations, as follows:

I. No descending processes of the parietal bones.
Vertebree and ribs free and separated from a

bony exoskeleton; no descending pro-
cesses of the parietals ; Athecee.

II. A carapace and plastron, and descending process of
parietals.

a, Sacral and caudal ribs articulating with neural
arches only.

Neck bending in vertical plane, last cervical
articulating with first dorsal by zygapo-
physes only; pelvis not anchylosed ;
marginal boues wanting or rudimental ; Trionychoidea.

aa, Sacral and caudal ribs articulating with body
of vertebree only.

As the last; but marginal bones present and
connected with ribs, and last cervical and
last dorsal vertebree articulating by
bodies; pelvis not anchylosed to plastron; — Cryptodira.

Neck bending in horizontal plane, the last
cervical and first dorsal vertebree articu-

VERTEBRATA 63

lating by bodies; pelvis anchylosed to
carapace and plastron ; marginal bones
present and connected with ribs; Plewrodiva.

The dAtheew includes the single family of the Dermo-
chelyde.

The Trionychoidea includes only the Trionychide.

The Cryptodira embraces the Cheloniide, Testudinidie
(Fig. 18), Cinosternide, Dermatemydide, Chelydride,
Baénide, and Adocide.

The Plewrodira includes the the Pleurosternide, Sterno-
theridee; Pelomedusidee, Plesiochelydide, Chelydidee, and
Carettochelydide.

The earlier tortoises are intermediate in character he-

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64 COPE

tween the Cryptodira and Pleurodira, and they have con-
tinued side by side to the present day. The Pleurodira
are, however, now confined to the Southern Hemisphere.
The Trionychoidea first appear in the Cretacic, and con-
tinue as the so-called soft-shell turtles of fresh water. The
Athecee first appear in the Eocene, and were then and still
are marine in their habits; but one species now exists, the
“leather-back turtle.”

Fra. 19.--Ichthyosaurus tenuirostis Conyb. ; one.sixth natural size. From the Lias
of Wiirtemburg. From Doederlein.

The order IcHrHyopreRyGIA embraces the families of
Ichthyosauride (Fig. 19) and Mixosauride. The Mixo-
sauridee have the bones of the forearm and leg distinguished
by their form from those of the carpus and tarsus, and they
thus approach more nearly other reptiles. They are the
oldest family, having inhabited European waters during
the period of the Trias. In the Ichthyosauride all the
bones of the limbs have the same form, except the humerus
and femur. They continued until the close of the Cretacic.
They were the especially marine Reptilia, having the
shape and habits of the Cetacea. They reached the length
of twenty-five feet during the Cretacic period. Their re-
mains have been found in all parts of the world except
Africa, but they were more numerous in the oceans now
covered by Europe than elsewhere. Some of the forms had
but few (Mureenosaurus) and others (Baptanodon) no teeth.
The typical genera had numerous teeth grooved at the
base. The vertebre are short and biconcave.

VERTEBRATA 65

The order THEeRoMoRA includes five suborders, and per-
haps others. They differ as follows :

I. Palate imperforate; interior nares posterior.
Teeth molariform ; one occipital condyle ; Placodonta.
II. Palate perforated anteriorly for internal nares.

a, Occipital condyle single.

Dentition complete, teeth compressed ; Theriodonta.

Teeth absent or reduced to one pair above; = lnomodonta.
aa, Occipital condyles two.

Teeth compressed ; Cynodonta.

Teeth molariform ; Gomphodonta.

Some of the suborders are imperfectly known. In all of
them it is probable, and in many of them certain, that the
supratemporal bone extends forward above the malar nearly
to the orbit, and that it sometimes includes, with the latter,
a foramen (Cynodonta), thus simulating two postorbital
bars. In the Gomphodonta and Cynodonta the two occipi-
tal condyles correspond with those of Mammals, and the
quadrate is so abbreviated as to render it highly probable
that these groups are the ancestors of the Mammalia. With
the extreme shortening of the quadrate the quadrato-jugal
bone disappears, and the supratemporal becomes the zygo-
matic part of the squamosal of the Mammal. In these
suborders and in many of the Theriodonta, the canine teeth
are greatly enlarged. The families are as follows:

Priacoponta : Placodontide, Trias of Europe.

THERIODONTA: ?Proterosauride, Permian of Europe; Rhyn-
chosauridee, Trias of Europe and India; Lycosauride,
Permo-Trias of South Africa and Europe.

Anomoponta; Dicynodontide (Fig. 20), Permo-Trias of
South Africa, Asia and Europe.

Cynoponta: Galesauride, Permo-Trias of South Africa.

GompHoponta: Tritylodontide, Permo-Trias of South
Africa.

66 COPE

In all of the suborders of the Permian of South Africa
where the parts are known, the obturator foramen and the
coracoid bone are small.

Fia. 20.—Lystrosaurus frontosus Cope; an Anomodont from South Africa, one-
third natural size.

The PrestosaurIA embraces the following families :
Plesiosauride, Nothosauride, and Lariosauride. The last
two families are only known from the Triassic of Europe.
The Plesiosauride range from the Jurassic to the Cretacic,

VERTEBRATA 67

inclusive, and include many species. ‘They are all adapted
for aquatic life, and were sometimes formed like the Cetacea ;
but others had very long necks, which could extend the
head to great depths or reach the surface when the body
was sunk. Some of the species reached a length of forty
or fifty feet. Their remains have been found in North and
South America, Europe, and New Zealand.

The Pelvcosauria includes the
families of the Bolosauridee and
Clepsydropide, which were carni-
vorous and abundantly provided
with teeth. They have been found
in the Permian of South Africa,
North America and Europe.

Two families enter the OrnI-
THOSAURIA, viz., the Pteranodon-
tide and the Pterodactylide. The
last-named family is furnished
with teeth, while the first-named
includes species which have a
toothless beak like that of a bird.
The Pterodactylide range from
the Trias to the Cretacic inclusive.
The earlier forms had a long tail,
and these continued through the »
Jurassic (Fig. 21). Some of the M
later forms had an edentulous
beak in front of the toothless por-
tion. The true Pterodactyles had
a much abbreviated tail and a !
long neck. H

The Ornithosauri were the fly- j/
ing order of reptiles, comparing y
with the other orders much as
bats compare with other orders en ee eee x

w. from the English Trias,
of Mammalia sternum. From Owen.

68 COPE

The Dinosauria embrace two suborders, as follows :
Pubic elements directed downwards ; Saurischia..
Pubic elements directed backwards; Orthopoda.

The Saurischia were mostly of carnivorous habits, while.
the Orthopoda were herbivorous. Both suborders com-.
mence in the Trias, and close with the end of the Postcre-
tacic. To the Saurischia belong two families, the
Cetiosauride and the Megalosauride, the former supposed
to be herbivorous, the latter carnivorous. The former have
robust inferior pelvic bones, and teeth with spoon-shaped
crowns. The latter have long, slender pubis and ischium, .
and sharp, knife-shaped teeth.

To the Cetiosauride belong the largest known Vertebrata
with ambulatory legs; the Camarasaurus supremus Cope
measuring seventy feet in length, and the Amphicelias
Jragilissimus Cope being considerably larger. These crea-.
tures had the dorsal vertebree hollow, and probably pene-
trated in life by branches from the lungs, while the tail
vertebrae and limb bones are solid. The former served as
floats and the latter as anchors as they walked on the sea
bottom. This family is only known from the Jurassic of —
Europe and North America (Fig. 22). The Megalosauridz
ranged throughout all Mesozoic time. Some of the species
were quite small, and others were gigantic, being the most
dangerous carnivorous animals that ever existed. Many of”
them were of kangaroo-like form, and in some of them the
vertebree were mere hollow shells. In most of them the.
limb bones were hollow.

FiG. 22.Catiosauride ; A, Camarasaurus supremus Cope ; cervical vertebra, from above and side,
one-tenth natural size; B&, do., dorsal vertebra, from behind, one-tenth natural size ; C, Amphicelias
altus Cope; femur, from behind, one-tenth natural size.

70 COPE

The Orthopoda includes the Scelidosaurids, Hypsirho-
phide, Agathaumide, Camptosauride, Iguanodontide and
Hadrosauride, all supposed to have been of herbivorous
habits. Representatives of these families have been found
in both Europe and North America, excepting the Scelido-
sauride, the members of which are so far only known
from Europe. They all include large forms, but the
most gigantic are members of the last four families.
These had an additional bone, the predental, which formed
a toothless extremity of the lower jaw, while the Aga-
thaumide had a corresponding toothless bone in front
formed of the premaxillary. In all the families ex-
cept the Agathaumidee the successional teeth appear on
the inner side of the base of the functional teeth, as in
lizards. In the Agathaumide they appeared under the
middle of the base, as in crocodiles. In all except Hadro-
saurides one row wasin functional use at a time; but in the
Hadrosauride two or three rows were used at once, Some
of the species of the latter had as many as 2,000 teeth ar-
ranged in four magazines, one in each jaw (Fig. 23). The
Agathaumide mostly had formidable horns on the head, at
the middle of the nose and over the eyes. Agathawmas
sylvestre Cope reached a length of forty feet, and had the
legs of sub-equal length. Many of the Hadrosauride were
of kangaroo-like proportions.

The Dinosauria include the great land reptiles of the
Mesozoic system. The order embraces no species adapted
for flight, and none adapted for a life in which movement
was made by paddles or limbs modified for swimming.
The order includes the largest land animals that have ever
lived, and their remains have been found in all continents
except the Australian.

VERTEBRATA

, from side ;

A

pe.

skull one-seventh natural size. From Co

beds of Dakota.

y:

clonius mirabilis Leid

B, from below. From the Laramie

—Di

F1a, 23

72 COPE

tf,
P”
iy |
i
Hy

|
iy

y':
H
pal I"

ee

Ae mM

3 Lee tar
Mauls ys fh Me
IN wel 7 4, ; AY Li : {

er
4 -

Fig. 24,—Belodon buceras Cope; skull, one-fourth natural’ size, From the Trias of the
Rocky Mountains. 4, the side; B, below; C, above. Original.

VERTEBRATA 73

The Lortcata includes three suborders, as follows:

Posterior nostrils opening in front of palate ;
nasals very short; premaxillary very
large ; external nostrils posterior, nasals
short ; Parasuchia.

Nasal bones very long, separating the small
premaxillaries; external nostrils an-
terior ; Pseudosuchia.

Nareal canal underroofed to behind larynx ;
no clavicle; pubis excluded from acetab-
ulum ; external nostrils anterior ; Eusuchia.

The Parasuchia and Pseudosuchia are restricted to the
Trias of both continents. The most important family of
the Parasuchia is the Belodontide (Fig. 24). The Pseudo-
suchia include the Aétosauride, the members of which
were completely incased in an armor of bony plates. To
the Eusuchia belong the true crocodiles. They commence
in the Jurassic with the Teleosauride and Goniopholidide,
which are followed in the Cretacic by the Crocodilide (Fig.
25) and Gavialide, which still exist.

The RHYNCHOCEPHALIA include two suborders:

Vertebree amphiccelous; axis undivided ; Sphenodontina.
Vertebre amphiplatyan ; dentatum of axis
separate ; Choristodera.

To the Sphenodontina belong the Protorosauride and
Paleohatteriide, from the European Trias, the Homeeosauri-
dee, from the European Jurassic, and the Sphenodontide,
which are represented by one species now living in New
Zealand. They are all of terrestrial habit, and of small or
medium size. The only family of the Choristodera is that
of the Champsosauride, which is found in the Postcretacic
of North America and the Eocene of Europe. The
species were aquatic, and reached the size of some of the
modern caymans. The dentine is inflected at the base of
the teeth, and the limbs were paddle-like. This order fur-
nishes the starting point for the great Archosaurian series.

74 COPE

Fig. 25.—Crocodilus acer Cope; skull, one-third natural size. From the Eocene
of Utah, Original.

The SquamatTa is an extended group which exists under
the following three subordinal forms:

Quadrate supported usually by exoccipital ;
paroccipital bone on end of exoccipi-
tal peduncle; frontal and parietal
little decurved; brain-case open in
front; roots of teeth dentinal ; Lacertilia.

VERTEBRATA 75

Quadrate supported by paroccipital, which
is deeply embraced by the exoccipital
and petrosal; frontal and parietal
little decurved, brain-case open in
front ; roots of teeth osseous ; Pythonomorpha.

Quadrate supported by paroccipital which
articulates with the brain-case direct-
ly; frontal and parietal decurved to
sphenoid, closing brain-case in front ;
teeth rootless ; Serpentes.

In general the Lacertilia have two pairs of limbs, and
the corresponding scapular and pelvic arches, but there are
many exceptions. The Pythonomorpha known have
two pairs of paddles in which all of the elements are dis-
tinguishable, and scapular and pelvic arches. In the Ser-
pentes limbs and the supporting arches are wanting, ex-
cepting rudiments of the posterior in a few forms.

The three suborders of Squamata first appear in the
Cretacic system. The Lacertilia and Ophidia still exist,
but the Pythonomorpha do not survive Mesozoic time.

The LacertTit1a embrace the following superfamilies:

I. Petrosal not produced anterior to semicircular canal,
and not articulating above with the parietal ; olfac-
tory lobes not under-arched; hemipenis mostly
calyculate.

Digits, including metapodials, in opposing

groups of two and three about a cen-

trale carpi and tarsi respectively ;

tongue papillose, extremity sheathed;

no clavicles ; 2 Rhiptoglossa.
Digits all directed forwards; clavicles
proximally simple; interclavicle

anchor-shaped ; tongue papillose, not
sheathed ; 3 Pachyglossa.

76 COPE

II. Petrosal produced anterior to semicircular canal,

not articulating above with the edge of the parietal.

a, Clavicle proximally expanded ; olfactory lobe
underarched by frontal.

Tongue papillose or smooth; hemipenis
calyculate ; 4 Nyctisaura.

ao, Clavicle proximally simple; olfactory lobes
underarched by frontal.

Vertebree amphicelous; no supratem-
poral arch; tongue papillose ; 5 Urolatoidea.

Vertebre procceelous; a supratemporal arch;
interclavicle anchor-shaped; tongue
smooth ; hemipenis flounced ; 6 Thecaglossa.

Vertebre procclous, no supratemporal
arch; interclavicle simple; tongue
papillose; hemipenis flounced ; 7 Helodermatoidea.

aaa, Clavicle simple proximally, olfactory lobes
not underarched by frontal.

Interclavicle cruciform ; tongue papillose;
hemipenis flounced; 8 Diploglossa.

aaaa, Clavicle proximally expanded; olfactory
lobes not underarched.

Vertebre proccelous ; tongue scaly ; hemi-
penis flounced or plicate ; 9 Leptoglossa.

III. Petrosal produced anterior to the anterior semi-
circular canal, articulating above with the border
of the parietal.

a, Olfactory lobes underarched by frontals, no
supratemporals, nor ceratohyals; cervical and
caudal intercentra codssified with the middle
of the centra.

VERTEBRATA 77

Palatine and pterygoid foramina; an epi-
pterygoid ; tongue papillose ; 10 Annielloidea.

No palative nor pterygoid formina; no
epipterygoid ; 11 Annulati.
A group which does not enter any of the above super-
families is the Dolichosauria (1.) On account of ignorance
of its essential characters, its position cannot be definitely
fixed, but it differs from other Lacertilia in the large num-
-ber of cervical vertebree, which is more than nine, accord-
‘ing to Owen. Teeth pleurodont.

The families of these superfamilies are the following :

Dolichosauria ; Dolichosauriide.

Rhiptoglossa ; Chameleonide.

Pachyglossa ; Agamide, Iguanide (Fig. 26.)

Nycisauria ; Kublepharide, Geckonide.

Uroplatoidea ; Uroplatide.

Thecaglossa ; Varanide.

Helodermatoidea ; Helodermide.

Diploglossa ; Zonuride, Pygopodide, Anguide, Xenosauri-
dee.

Leptoglossa ; Xantusiide, Teiidee, Lacertide, Gerrhosauride,
Scincide, Acontiide, Anelytropide.

Annielloidea ; Aniellide.

Annulati ; Chirotide, Amphisbeenide, Trogonophide.

Fig, 26.—Skull of Iguana tuberculata Linn ; external side view, natural size.

The recent forms of the Lacertilia are much more numer
ous than the known fossil forms.

COPE

78

The geologic history of the Lacertilia is as follows:

Column No. 1 represents the Dolichosauria.

OMMOIN|T

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OTMOAR,

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* oIsseldy,

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9119009 NT

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Or

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9u90048I| q

VERTEBRATA 79

The Dolichosauria are only known
from European formations. Some
of the species are snake-like in form.
Acrodonta are best known from the
same region, but one species is known
from the American Eocene. It may
belong to the Rhiptoglossa. The
extinct Iguania and Leptoglossa are
all European as far as known, while
Diploglossa are known from both
continents.

The PytHonomorpna (Fig. 27) in-
clude two families, the Mosasauridee
and the Plioplatecarpide. The
Mosasauride were the predominant
type of sea-saurians during the Cre-
tacie period in North America,
and they were common in Europe
and in New Zealand, and some
species have been found in Brazil.
The Plioplatecarpide are only known
from the Upper Cretacic in Eu-
rope. Some of the species of Mosa-
saurus and Liodon reached a length
> of fifty feet. Their limbs were short,
inflexible paddles, and the pelvic
bones were very slender and feeble.

The Opuipia include the follow-
ing superfamilies :

A. Supratemporal intercalated
in the cranial walls. (Angio-
stomata.)

a, No ectopterygoid; pala-
tines bounding choanz

posteriorly ; ethmotur-
binal forming part of

Fie, 27.—Clidastes _propy- roof of mouth; rudi-
thon Cope; skull one-half natur- :
alsize. From the Upper Creta- ments of a pelvis. (Sco-

ic of Alabama. dt
cic of Alabama lecophidia.)

80 COPE

Maxillary bone fixed to prefrontal and pre-

maxillary ; a pelvis; (1) Catodonta (blind snakes).
Maxillary bone vertical and free from all
others; no pelvis; (2) Epanodonta (blind snakes).

aa, An ectopterygoid; palatines not bounding
choane posteriorly.

Maxillary bone free, horizontal ; (8) Tortricina.
AA. Supratemporal attached scale-like to cranial walls,

produced freely posteriorly ; ectopterygoid present.
(Eurystomata.)

Maxillary bone horizontal, not articulated

with the prefrontal by a ginglymus,

hence not erectile ; (4) Colubroidea.
Maxillary bone vertical, not reaching pre-

maxillary, articulating with the prefron-

tal by a ginglymus, and to the ectoptery-

goid without imbrication, and bearing a

perforated tooth; (5) Solenoglypha (vipers and pitvipers).

By far the greater number of snakes belong to the su-
perfamily Colubroidea. These fall into four primary
tribes which differ as follows:

I. No grooved or channeled teeth.
Rudimental posterior extremities ; Peropoda.
No rudiments of extremities; Aglyphodonta.
II. Some teeth grooved or channeled.
One or more grooved teeth on the posterior

part of the maxillary bone ; Opisthoglypha.
A channeled or perforate tooth at the an-
terior end of the maxillary bone; Proteroglypha.

Fig. 28.—Xenopeltis unicolor Reinwt, from Siam; skull, natural size. Original.
(Asinea. )

VERTEBRATA 81

The Peropoda are believed to be the most primitive
snakes, from which the other divisions have been derived.
The line to the Proteroglypha and Solenoglypha is an as-
cending one, while that to the Tortricina and Catodonta is
a descending one.

The families embraced by these superfamilies are as fol-
lows:

Catodonta ; Stenostomide.

Epanodonta ; Typhlopide.

Tortricina ; Tortricide, Uropeltidee.

Colubroidea ; (Peropoda) Pythonide, Boide (Fiz. 29), Char-
inide, Ungualiide ; (Aglyphodonta), Xenopeltide (Fig.
28), Acrochordide, Nothopide, Colubride ; (Opistho-
glypha) Dipsadide ; (Proteroglypha), Elopide, Naji-
dee, Dendraspidide, Hydrophide.

Solenoglypha ; Causidee, Atractaspididee, Viperidee, Crotali-
dee.

The paleontology of the snakes is very imperfectly known.
The oldest known genus, Symoliopsis Sauv., from the Neo-
comian of France, may be either one of the Catodonta or
Epanodonta. The oldest forms of Asinea (harmless snakes)
are Pythonide and Boide; they occur in both continents.

Fic. 29—A, Paleophis littoralis Cope: B, P. halidanus Cope; vertebr Both
from the Eocene of New Jersey. Cope.

82 COPE

The largest are the species of Paleophis Owen, which occur
in the Eocene of New Jersey and England; they equaled
the largest existing pythons in dimensions (Fig. 29). The
venomous snakes appear in the Neocene in both Europe
and North America.

The geological distribution of the superfamilies of snakes
is as follows:

Plistocene

Neocene

Eocene

Cretacic

Jurassic

Triassic

Carbonic

Devonic

Siluric

Ordovicic

Cambric

Huronic

VERTEBRATA 83

Subclass IJ.—AVES.

There are two superorders of the birds as follows :

Metacarpal and carpal bones all distinct, the
digits with ungues; caudal vertebre
numerous, unmodified ; clavicles united ;
pelvic elements distinct ; teeth present ; Saurure.

K.Sch.gez,

Ps)

Fic. 30.—Archeopteryx lithographica Wagn. From Middle Odélite of Bavaria
much reduced. From Dames.

84 COPE

Metacarpal and carpal bones reduced in num-
ber, codssified; ungues wanting or
single; caudal vertebree reduced in num-
ber, the terminal areas usually codssified; Eurhipidure-

The Saurur# includes but one order, which is defined’
as follows:
Vertebre biconcave; feathers arranged in
one series on each side of the caudal ver-
tebree ; teeth present ; Ornithopappt..

To this order but one family belongs, viz., the Arche-
opterygide.

This family is represented by one genus, Archeopteryx
from the Odlite (Middle Jurassic) of Solenhofen, Bavaria.
lt was furnished with long feathers on the wings, adapting
it for flight, and the long lizard-like tail had a row of long
feathers on each side of it. It is the oldest known type of
birds, and is an important one as showing the near connec-
tion of birds with reptiles. (Fig. 30.)

The Evuruiripur# include four or five tribes which dif-
fer as follows:
Ischium free from ilium posteriorly ;*
palate dromeognathous (7. ¢., maxillo-
palatines articulating with vomer
which is between them, palatines not
articulating directly with sphenoid
rostrum); no teeth ; (1) Ratite.

Ischium free from ilium posteriorly; pal-
ate? dromegnathous ; teeth; (2) Odontolee.

Ischium free from ilium posteriorly ; pal-
ate not dromeognathous(?); teeth; (3) Odontotorme.

Ischium codssified with ilium posteriorly ;
palate not dromeognathous; feathers

*With some apparent exceptions.

VERTEBRATA

distributed in areas, those of the wings
much differentiated ;

Ischium codssified with ilium posteriorly ;
no teeth; feathers universally distrib-
uted and not differentiated on wings;

The Ratitz are the ostriches and their alli

85

(4) Huornithes.

(5) Impennes.
es; the Odon-

tole and Odontotorme are toothed birds; the Impennes

are the penguins, and the Euornithes are the

remaining, or

typical birds. The geological distribution of these tribes

is as follows:

Plistocene

Neocene ... |

Eocene

Cretacic . ... | | |
|
{
tT

Jurassic

Triassic

Paleozoic . .

The Ratit# includes the following orders :

Sternum without keel; clavicles; wings rudi-
mental ;

Sternum without keel; no clavicles; wings
rudimental ;

Sternum with keel; clavicles; wings rudi-
mental ; =

Sternum with keel; clavicles; wings func-
tional ;

Struthiones..
Apteryges..
Gastornithes..

Crypturt..

86 COPE

The families belonging to these orders are the following :
Struthiones ; Struthionide (ostriches), Rheide, Casuariide,
Dromeeidee, Dinornithide (Fig. 31), Aipiornithide.

Apteryges ; Apterygide (kiwis).
Gastornithes ; Gastornithide.
Crypturi ; Crypturidee (tinamus).

Of the above families the Struthionidiv are remarkable
in having the pubes forming a ventral symphysis. The
Rheide are equally remarkable in having a ventral sym-
physis of the ischia. The geological range of these orders
is as follows:

|
Struthiones _ Apteryges | Gastornithes Crypturi

Neocene . ...

Plistocene

Eocene

Cretacic . .. .

Jurassic .

Triassic

Carbonic .

Devonic

Siluric .

Ordovicic

Cambric. ..

Huronic. ...

VERTEBRATA 87

Extinct Struthiones are known from all the great regions
of the Old World, but none from the New. The extinct
Apteryges come from the Australian realm only. The
Gastornithes are known from the Eocenes of both Europe
and North America. Gastornis has persistent sutures of
the skull, and a tooth-like process on the upper jaw. G.
edwardsii was nine feet high. The species of Dinornis (Fig.
31) from the Plistocene of New Zealand were all of large
size, some of them reaching twelve feet in height. One
species is known from Australia.

Fic. 31.—Dinornis parvus Owen. From Plistocene of New Zealand. Irom
Owen. JI, ilium; is, ischium; pp, pubis; s/, sternum; B, tarsometatarsus.,

The tribe OpoxroLtc includes but one order:

Teeth in a groove; sternum without keel ;
wings rudimental; pelvic bones free
posteriorly ; vertebre saddle-shaped; = Dromeopapp?.

The Drom«opraprr has but one family, the Hesperorni-
thidee, which have been only found so far in the Upper

88 COPE

Cretacic of Middle North America. They have many of
the characters of the divers (order Cecomorphe).

To the OponrororM#& one order only is referred. It is
thus characterized :

Teeth in sockets; sternum keeled; wings
well developed; ischium and pubis free
posteriorly ; vertebree biconcave ; Pteropappi.

The family of the Ichthyornithide is the only one
known to belong to the Prrroparri. Its members are
known from the Upper Cretacic of North America and the
Lower Cretacic of England.

The EvornirHEs include numerous suborders, which are
defined as follows:

1. Maxillopalatines united across the middle of the
palate. (Desmognathe).

A. Four toes directed forwards (pamprodactyl-
ous).

Toes webbed ; .no basipterygoids ; (1) Steganopodes.

Toes free; vomer unossified; no_basi-
pterygoid processes ; (2) Colioidei.
AA. Three toes directed forwards.*
Short basipterygoid processes; toes gen-
erally webbed ; preecocial ; (8) Chenomorphe.

No basipterygoid processes; bill and legs
slender; toes generally free; altri-

cial ; (4) Herodii.
Bill and claws hooked ; toes free; ver-
tebree saddle-shaped ; altricial ; (5) Accipitres.

Bill hooked; toes free; vertebree opis-
thocelous; rostrum movably ar-

*Except Cuculide and Rhamphastide, which are zygodactylous.

VERTEBRATA 89

ticulated with skull; basiptery-
goids ; (6) Heterospondyli.

Toes free; vertebree saddle-shaped ; ros-
trum fixed ; (7) Coceygomorphe.

AAA. First and fourth toes directed backwards
(zy godacty lous).

Rostrum freely articulated with the
skull; vertebree opisthoccelous ; (8) Psittact.

AAAA. First and second toes directed backwards
(heterodactylous).

Basipterygoids present; heteropelmous; (9) Trogonoidei.

II. Maxillopalatines not united across the palate ;
vomer narrowed and acute in front. (Schizo-
guathe.)

A. Toes three forwards (anisodactylous).

Schizorhinal ; toes webbed ; (10) Ceconorphe.
Toes free; legs long; feathers with after-

shaft ; preecoces ; (11) Gralle.
No basipterygoids; lachrymal bones co-

ossified with rostrum ; toes free ; (12) Opisthocome.
Toes free; hallux rudimental ; (13) Galline.

Toes free; hallux well developed; two
carotid arteries ; (14) Pullastre.

Toes free; hallux well developed; one
carotid artery; basipterygoids; (15) Micropodinidci.*

AA. First and fourth toes directed backward
(zy godacty lous).

* Family Trochilidiv.

90 COPE

No ceca coli; no interclavicle; one
carotid artery ; (16) Picoidet.

III. Maxillopalatines not united on median line;
vomer single, truncate, and excavated in front.
(Aigithognathe.)

A. Toes three in front (anisodactylous).

Toes free; hallux well developed; tar-
sometatarsus with five tendinous
canals; basipterygoids wanting or
rudimental; sternum with two
notches; no ceeca coli; one carotid
artery ; (17) Passeres.

AA. Four toes directed forwards (pampro-
dacty lous).
Toes free; no basipterygoids; sternum
entire posteriorly; tensor patigii
brevis muscle attached to a tendon
which extends to the manus; no
ceecacoli ; (15) Micropodioidet.*

The cormorants, pelicans, and boobies of the Stegano-
podes appear in the Eocene. The Eocene representatives
of the Chenomorphe are primitive flamingoes, the true
ducks and geese not appearing before the Neocene. The
Eocene Accipitres are Falconide, the owls not appearing
before the Neocene. The kingfishers are the earliest repre-
sentatives of the Coccygomorphe, appearing in the Eocene;
the remaining families are not known prior to the Neocene.
The Phasianide of the Galline appear in the Middle Eo-
cene. It is thus evident that the majority of the families
of the Euornithes are not known prior to the beginning
of the Neocene system.

The time distribution of the Euornithes is as follows, as
far as known:

* Family Cypselidex.

91

VERTEBRATA

| 1
ae

Neocene

Eocene . |

10

11

12

13

=
|
|

Cretacic ©

Jurassic

Triassic

Carbonic ;

Devonic

Siluric .

|
|
|
|
|
|
|

Ordovicic ,

|

Cambric |

Huronic

He COPE

us
B

Fig. 32.—Diagrams exhibiting the more important modes of distribution of the deep plantar
tendons in Euornithes. A, nomopelmous (schizopelmous); B, synpelmous (desmopelmous) ;
C, antipelmous; D, heteropelmous; jth, flexor longus hallucis; /pd, flexor perforans digitorum
v, vinculum, Jto IV, lst to 4th toes. (After Stejneger).

D

The digits of birds are arranged in several different orders.
The usual type with three toes forwards and one backwards
is termed anisodactylous; that with the first and fourth
backwards and second and third forwards, is syndactylous-
When the first and second are directed backwards and the
third and fourth forwards the arrangement is termed he-
terodactylous; and when all four are directed forwards the
foot is said to be pamprodactylous. The flexor tendons of
the toes are arranged differently in these different digital
arrangements, although not always identically in the same.
The arrangement usual in the anisodactylous foot is the
schizopelmous system, where the flexors of the digits 2-3-4
have a common stem, while that of digit 1 is distinct.
In some anisodactylous feet, however, all the- flexors or ten-
dons are more or less fused ; this is the desmopelmous sys-
tem; it occurs also in many zygodactylous feet. In an-
other type of zygodactylous foot (that of the Picoidei) we
have the antipelmous arrangement. Here all the tendons
are fused excepting that of third, which is distinct. Finally
in the heterodactylous foot we have the heteropelmous

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RTEBRATA

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94 COPE

tendons, where the first and second form a common stem,
and the third and fourth a common stem, following the ar-
rangement of the digits. (Fig. 32).

The palatal structure must be consulted in attempting
the discrimination of the orders of birds. There are four
types of structure, as follows: First, the dromeognathous
palate (Fig. 33 D) has the anterior roof continuous and
formed of the fused maxillopalatines and vomer. In this
case, also, the vomer and pterygoid bones exclude the
palatine from contact with the presphenoid, by intervening
between the two. In the desmognathous palate (Fig. 33 C)
the maxillopalatines join each other on the middle line,
excluding the vomer, which is free and above them or
aborted. Both palatines and pterygoids reach the pre-
sphenoid bone at its basipterygoid processes. In the schiz-
ognathous palate (Fig. 33 B) the maxillopalatines do not
meet on the middle line, and the vomer terminates in a free
acute apex above, and more or less posterior to them. The
fourth type (the egithognathous, Fig. 33 A) is a modifica-
tion of the schizognathous, differing only in the form of the
vomer. The anterior extremity of the latter is flattened,
often expanding and notched or excavated on the anterior
margin.

The families of the EvorniruHes are as follows:

Steganopodes ; Pheetonide, Fregatide, Pelecanide, Sulide,
Phalacrocoracide, Plotide.

Colioidei ; Coliide.

Chenomorphe ; Palamedeide, Anatide, Phcenicopteride.

Herodii ; Ibidide, Ciconiide, Baleenicipitidee, Ardeide.

Accipitres ; Cathartide, Falconide, Pandionide, Strigide.

Heterospondyli ; Steatornithide.

Coccygomorphe ; Cuculide, Coraciidee, Alcedinide, Upupi-
dee, Musophagide, Todide, Momotide, Bucerotide,

VERTEBRATA 95

Rhamphastide, Caprimulgide, Bucconidie, Indicatori-
dee.

Psittaci ; Psittacidee.

Trogonoidei ; Trogonidie.

Cecomorphe ; Colymbidee, Heliornithide, Alcidee, Laride,
Procellariidee.

Gralle ; Chionide, Thinocoride, Glareolidee, Dromadide,
Charadriide, Otidide, Eurypygiide, Rhinochetide,
Cariamidee, Psophiide, Gruide, Rallide.

Opisthocomi , Opisthocomide.
Galline , Tetraonide, Phasianide.
Pullastre ; Cracidee, Megapodiide, Pteroclidie, Didide,
Columbidee.
Micropodioidei, Cypselidee, Trochilide.
Picoidee ; Picide, Yngide.
Passeres. This order is divided into five superfamilies, as
follows:
I. Tensor patagii brevis picarian ; Menwrorder.
II. Tensor patagii brevis passerine.
A. Syrinx mesomyodian,

Tendons of flexor muscles of foot desmo-

pelmous ; ELuryleemoidet.
Tendons of foot schizopelmous; syrinx bron-

chiotrachial ; Tyrannoidet.
Tendons of foot schizopelmous; syrinz tra-

chial ; Formicaroidei.

AA. Syrinx acromyoctian.

Tendons of foot schizopelmous ; Passerovdet.

The families of these superfamilies are the following :

Menuroidei ; Menuride, Atrichornithide.

Eurylemoidei ; Eurylemide.

96 COPE

Tyrannoidei ; Xenicide, Philepittide, Pittide, Tyrannide,
Cotingide, Phytotomide.

Formicaroidei ; Conopophagide, Pteroptochide, Formi-
cariidee.

Passeroidei ; Alaudide, Motacillide, Timaliide, Liotrichide,
Muscicapide, Turdide, Cinclide, Troglodytide,
Chameide, Hirundinide, Artamide, Laniide, Paride,
Paradisiidee, Corvide, Sturnide, Meliphagide, Nec-
tariniidee, Certhiide, Ploceide, Tanagride, Icteride,
Fringillide.

Fig. 34.—Diagrams of a tracheal and a bronchial syrinx. t.c, trachelo-clavicular
muscle. (After Newton.)

Fie. 35—Lateral and dorsal views of the acromyodian syrinx of the raven (Corvus
corax), B, I, II, IV, second, third and fourth bronchial rings. 1, m. tracheo-
bronchalis ventralis ; 2, m. tracheo-bronchialis obliques; 3 and 4, m. tracheo-bron-
chialis dorsalis longus et brevis ; 5, m, syringeus ventralis ; 6, m. syringeus ventralis-
lateralis ; 7, m. syringeus dorsalis. (After Newton.)

VERTEBRATA 97

Fic. 36.—Diagrams of the elbow muscles of Icterus (A), showing the passerine
type and Nenina( B)., showing the picarian type. b, biceps; emr/, extensor meticarpi
radialis longus; A, humerus; s, shoulder; s7, secondarv remiges: ¢, triceps; tpd,
tensor patagii brevis; ¢pl, tensor patagii longus. (After Newton.)

To the Impenyes but one order belongs. This is the

Ptilopteri :

[lium not anchylosed with sacrum; bones of
wing not foldable on each other; meta-
carpals not separated; hallux directed for-
wards ; feathers scale-like ; vertebree opis-
thoccelous ; Ptilopteri.

The Ptilopteri includes the recent family of Aptenody-
tide or penguins. They first appear in the Upper Eocene
of New Zealand. No species, living or extinct, have been
found in the Northern Hemisphere.

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-ue ‘9 Suury seuvolye seydatq so uvrprosoqoid ‘g ‘edog snawmrid snpoosuayg Jo snopodvaxry ‘vy ‘equ[nsuy jo eanjonzys pedivo Jo sodéT—9g “Ol

v

2 ‘ainSy oy Jo w Si ayy Moray | pur Gg 47 UL S4zoy ay. Worf Taqunu sySip oq} g pure KP uy ‘ado wnpn719v)
wn7tay}0.1ga0,7 JO {Azoepouay) suorptrpdip ‘qb adog warjpoorp WHIAIYPIAINTT JO (spoupossiiad; snosmgpwepdrp "Gg sadeg sndojunydaja wvopoydnsog
nine snpoopwyg Ju suopodoaxry ‘fr—Isivy equpnusun jo sods E— oe ‘OT

jo snopadsjqure $9 Staury suunop spydaig Jo Wernasoqoad "gy todo, s

"eYSVIQON Jo oUV000N (1eddqQ) Y1Oq dnoy oy. Wor
‘potddns suoniod pepeysun ey, “adop ‘qd “W Jo UOTZO9T[00 UT NOJOTeYs Woy poioysoyY “poonper yon ‘ Apray snaws uopounjy—1E “OLA

ma un

VERTEBRATA 101

Class V—MAMMALIA.

The primary characters of the Mammalia are seen in
their limbs and in their teeth. The general characters of
the skeleton may be learned from Fig. 37 (Zlurodon sxvus
Leidy). The mutilate type of limbs is seen in C‘ctotheriwm
cephalus Cope (Fig. 45, facing page 108). The phalanges are
connected by integument and form an inflexible paddle, and
in the typical forms the elbow is also inflexible. The dif-
ference between unguiculate and ungulate phalanges may
be seen by comparing Fig. 37 and Fig. 55 (page 119) with
Figs. 38 and 39. The compressed curved form of the former,
adapted for prehension, is easily distinguished from the flat
generally wide type seen in the latter, which is adapted for
support. The different carpal and tarsal types of the Ungu-
lata are seen in Figs. 38 and 39. A, is the taxeopodous; B,
the proboscidian; C, the amblypodous; and D and FE, the
diplarthrous, represented by the Perissodactyla (D) and Ar-
tiodactyla (E) respectively.

The molar types of dentition in historical and develop-
mental order to the quadritubercular are represented in Fig.
40. Forms up to No. 5, inclusive, predominate in the Un-
guiculata ; and 6 and 7 with their derivatives predominate
in the Ungulata. The derivatives of Nos. 6 and 7 are
formed by the development of ridges or crests which con-

()

t

B
3B is g
3 BE

Fie, 40.—Diagrams of types of mammalian dentition, from Osborn; 1, haplodont; 2, pro-
todont; 8, triconodont; 4, tritubercular superior and inferior; 5, tritubercular superior,
tuberculo sectorial lower; 6, quadritubercular superior, quinquetuberculur inferior; 7, quad-
ritubercular, both jaws. pr, protocone ; pa, paracone ; me, metacone.

102 COPE

nect the cusps longitudinally or transversely, forming vari-
ous patterns, two of which are represented in Figs. 59
(page 124) and 63 (page 128). Such types are termed
lophodont. Those in which the crests and the grooves be-
tween them become respectively elevated and profound are
termed ptychodont, Figs. 51 (page 114) and 60 (page 125).

Two subclasses are known to belong to the Mammalia:

An interclavicle ; a large coracoid articulating
with the sternum ; Prototheria.

No interclavicle; coracoid very small, codssified
with scapula; not reaching sternum ; Eutheria.

The Prototheria have one existing order, the Monotre-
mata, and it is supposed, with much probability, that two
orders which appear in the Trias and continue until the
Eocene, inclusive, belong to it. If this classification is
admitted, the Eutheria have their first representatives in
the Postcretacic, and their latest are now numerous.

Of the Prororneria, there are probably three orders of
which species are known, but the location of the first two
enumerated below is not certain.

Incisors reduced; molars with compressed
cutting crowns, and undivided roots ; Protodonta.

Incisors much enlarged; molars with tuber-
cular grinding surfaces, and distinct
roots ; Multituberculata.

No teeth at maturity ; Monotremata.

The families are the following :
Protodonta ; Dromatheriide.

Multituberculata; Plagiaulacide, Chirogide, Polymastodon-
tide. Monotremata ; Ornithorhynchide.

VERTEBRATA 103

The geological range of these orders is shown by the
table below :

Protodonta Multituberculata Monotremata

Plistocene .

Neocene. .

Eocene . . ; |

Cretacic. .

Jurassic. .

Triassic . .

Carbonic

Devonic. .

Siluric

Ordovicic .

Cambric .

Huronic

Species of Proroponra have been found thus far only in
the Triassic beds of North Carolina. Their molar teeth
bear some resemblance to those of some of the correspond-
ing teeth of the Theromorous reptilian genus Dimetrodon.
The oldest MuLTITUBERCULATA appear In the Trias of South
Africa in the Karoo beds, and in the Upper Trias (Keuper)
of Wiirtemburg (Europe). They are more abundant in the
Jurassic of England and of the Rocky Mountain region of

104 COPE

Fig. 41.—Ptilodus medievus

Cope; mandibular ramus; a, b, Fia. 42.—Chirox plicatus Cope; palate and molar
4, c, $. Original. From Puerco teeth, from below, 3-2 natural size. From Puerco bed
bed of New Mexico. of New Mexico. Original.

America, and in the later Cretacic of North America
and of Patagonia. In the latest Cretacic (Puerco) of New
Mexico the largest forms occur, which equal an adult kan-
garoo (Figs. 41 and 42). The last of them are found in
the lowest Eocene in the north of France. Extinct and re-
cent MonoTtREMATA are restricted to the Australian realm.

The Eurueria are represented by the following numer-
ous orders:

I. Marsupial pelvic bones (generally); palate per-
forated ; (vagina double; placenta wanting; corpus
callosum rudimental; cerebral hemispheres small).
(Didelphia.)

One deciduous molar tooth shed; the
rest permanent ; (1) Marsupialia.
II. No marsupial bones; palate generally entire;
(one vagina; placenta and corpus callosum* well
developed). (Monodelphia.)
A. Posterior limbs wanting, or represented by

* More recent researches than those of Osborn in 1886, confirm the view of Owen
that this element is the hippocampal commissure. —H. F. O.

VERTEBRATA 105

minute rudiments; anterior limbs oar-like.
(Mutilata.)

Elbow joint inflexible; carpals discoid,
and, with the phalanges, separated
by cartilage; lower jaw without as-
cending ramus; (2) Cetacea.

Elbow joint flexible; carpals and pha-
langes with close articulations;
mandible with ascending ramus; (8) Strenia.

AA. Posterior limbs present; ungual phalanges
compressed and curved on one or all the
feet.* (Unguiculata.)

3, Carpal and tarsal bones generally in linear
series.

y, Teeth without enamel; no incisors.

Limbs ambulatory ; hemispheres small ; (4) Edentata.

77, Teeth with enamel; incisors present.

No postglenoid process; mandibular con-
dyle not transverse; mastication
proal; limbs not volant; hemis-
pheres small ; (5) Glires.

Anterior limbs volant; hemispheressmall; (6) Chiroptera.

A postglenoid process; mandibular con-
dyle transverse ; mastication orthal ;
scaphoid and lunar bones not coal-
lesced;+ hemispheressmall, smooth; (7) Bunotheria.

A postglenoid process; limbs not volant,
with a scapholunar bone; mastica-
tion orthal; hemispheres larger,
convoluted ; (8) Carnivora.

106 COPE

ff, Carpal and tarsal bones alternating ;
faceted.

Anterior limbs prehensile; mandibular
condyle and mastication transverse; (9) Chalicotheria.

AAA. Posterior limbs present ; ungual phalanges
not compressed and hooked.{ (Ungulata.)

8, Carpal, and usually tarsal bones in linear
series.§

Limbs ambulatory; teeth with enamel; (10) Taxeopoda.

BB, Carpal bones alternating externally ;
tarsals in linear series.

Limbs ambulatory, median digits
longest; teeth with enamel ; (11) Toxodontia.

88, Tarsal bones alternating ; carpals linear
or reversed diplarthrous.

Cuboid bone partly supporting navicu-
lar, not in contact with astragalus ;
no canine teeth ; (12) Proboscidea.

£288, Both tarsal and carpal series more or
less alternating ; the distal row inwards.

Os magnum not supporting scaphoides;
cuboid supporting astragalus; su-
perior molars tritubercular ; (13) Amblypoda.

Os magnum supporting scaphoides; su-
perior molars quadritubercular ; || (14) Diplarthra.

The geological range of these orders is as follows:

* Except Mesonychide, some Glires, and posterior feet of some Edentata.
t+Except Talpa and Erinaceus. {Except in the Hapalide. §Except in Dendrohyrax.
||Except Pantolestide.

107

VERTEBRATA

otmOIn yy

o1qurey

a * OIOIAOPIQ)

* ounig

o1m0Aagy

* o1moqie)

=e | OISSBIL,

-_ Hh a he nm oIssBin¢?

. _ 6 d1IORIOIO

wie iG is a ia 4 a08 * sua00ny

ree Sia me oe ; or hei 9119909 NT

Te aera | | i j ; oh ousd0}8I[ J
i ur a | nO | «| | «|e twits |e | *« 7 # le

108 COPE

The Marsupialia fall into two suborders, which differ as
follows:

: 4or5 °
Incisors 3554 Polyprotodontia.

i. 1to3 - .
Incisors 755 Diprotodontia.

The Polyprotodontia are carnivorous and insectivorous
in their habits, and their present range is Australasia and
the two Americas. They are the older of the suborders,
being represented in the Eocenes of Europe and North
America by spe-
cies of opossums
(Fig. 43). Genera
probably allied oc-
cur in the Post-
cretacic of North
America. Nolarge

forms are known.

: : Fia. 43.—Peratherium fugax Cope; anterior part of
The Diprotodontia skull. From Lower Neocene (White River bed) of

are now restrict- Colorado. X 2. Original.

ed to Australasia, and the extinct forms belong to the Plis-
tocene of the same region. These include some very large
kangaroos, with still larger animals of the genera Noto-
therium and Diprotodon. The Diprotodon australis Owen
was as large as a rhinoceros, but though allied to the kan-
garoo, was typical of a different family. The feet were
plantigrade, and the fore limbs were larger than the hind
limbs.

The Creracna are represented by three suborders, which
differ as follows:

External nostrils at middle of muzzle;
temporal fosse elongate, approxi-
mated; teeth ; Archeoceti.

External nostrils at base of muzzle; tem-
poral fossee short, lateral; teeth ; Odontoceti.

VERTEBRATA 109

External nostrils at base of muzzle; tem-
poral fosse short, lateral; no teeth,
but transverse horny laminz on the
upper jaw; Mysticett.

Fie. 44.—Zeuglodon cetoides Owen; skull, much reduced. From Eocene of
Alabama, From Gaudry.

The Archeeoceti are few in number, and are chiefly repre-
sented by the gigantic Zeuglodon (Fig. 44.) They are not
known before or after the Eocene system, nor out of the
geographical area of the Northern Hemisphere. The
Odontoceti first appear in the older Neocene, and the oldest
family, the Squalodontide, have the posterior teeth two-
rooted, as in the Zeuglodons. The other families with one-
rooted teeth were contemporary with them, and are repre-
sented in the present seas by numerous types, as the dol-
phins, sperm whales, etc. The Mysticeti, or whalebone
whales, appear in the Lower Neocene, and are still abun-
dant. They include the largest vertebrates. (Fig. 45.)

The Odontoceti and Mysticeti are probably independent
descendents of the Archeoceti. The former retain their
dental and rib,characters, but the nasal bones are more ab-
breviated than in the Mysticeti. In certain Mysticeti of the
Neocene the frontals as well as the nasals are somewhat
elongate, and some (e. g., Cephalotropis) have temporal
ridges.

Srrenta first appear in a very generalized family in the
Eocene beds of some West Indian Islands. In the Neo-
cene of Europe and North America forms approaching
more nearly to the existing types are not rare, especially

110 COPE

in the south of Europe. At present, species exist on the
shores of the continents in the warmer latitudes. (Fig. 46.)

The succession of types in this order is measured by
the modifications in the dentition. In the Eocene Pro-
rastomide we have a nearly normal dentition with dis-
tinct canines. In the succeeding forms canines are want-
ing and the incisors are either enlarged or disappear.
Thus, in the Neocene Halitheriide we have nearly nor-
mal molar dentition, with incisors reduced in number and
functioning as-digging tusks. The existing dugongs con-
tinue the enlarged incisors, but the molars are reduced in
number, and have become simple prisms. The also
existing Manatees have lost the incisors, retaining nor-
mal molars which have abnormally increased in number.
In the lately extinct Rhytinide all dentition has disap-
peared..

The Eprenrata have been generally restricted to the
Southern Hemisphere, although during the Neocene they
ranged as far north as the Augean Sea in Europe, and to
latitude 46° in North America during the Plistocene. They
first appear. in the Eocene of Patagonia, and were ex-
tremely common in the Neocene throughout tropical
America, where they are still abundantly represented.
A few species still remain in the Ethiopean and Paleo-
tropical geographical realms. The Megatheriide of the
South American Neocene were of large and gigantic size,
the largest species pertaining to the genus Megatherium,
which ranged in North America to South Carolina. The
Glyptodontidee were covered with an. immovable carapace
consisting of bony tesseree, somewhat like that of arma-
dillos. They varied in size from that of a sheep to that
of a rhinoceros. Species were abundant in the Neocene of
South America, when they also ranged north to Texas,
Florida, and Kansas. The most ancient (Eocene) Edentata
display traces of enamel on the teeth.

VERTEBRATA 111

The Grires are found in the strata of all regions, but in
reduced numbers in the Eocene in all countries except
temperate South America, where they were abundant at
that age. There are four suborders, which differ as fol-
lows:

I. Fibula not articulating with caleaneum; ankle and
elbow not tongued and grooved; one pair of incisors
in upper jaw.

Incisive alveolus not passing]into centre
of ramus of lower jaw; fibula dis-
tinct ; Hystricomorpha.

Incisive alveolus penetrating ramus; fibula

distinct ; Seiuromorpha.

Incisive alveolus penetrating ramus; fibula
coossified with tibia ; Myomorpha.

II. Fibula articulating with calcaneum ; ankle and el-
bow tongued and grooved; two incisors in upper
jaw.

Inferior incisor penetrating ramus; fibula
and tibia codéssified ; Lagomorpha.

The Hystricomorpha (porcupines, cavies, etc.) are
abundant in the Eocene of South America, and are pres-
ent in the Upper Eocene of France. They are more
abundant in the Neocene of South America and France,
and are present in the Upper Neocene of North America. At
present they are cosmopolitans, excepting Australia, but
they chiefly abound in South America. (Fig. 47.) The

COPE

112

/

fos

~

-y

a

a

-
mene ae"

Fie. 47.—Castoroides ohioensis F

tocene of Ohio.

uch reduced. From Plis-

m

; skull,

oster ;
From Hall and Wyman.

ys delicatissimus Leidy; Skull. From Eocene of

Fic. 48.—Plesiarctom

Wyoming. Original.

VERTEBRATA 1138

Sciuromorpha only occur fossil in the Northern Hemi-
sphere, and are the prin-
cipal Glires of the Eo-
cene. From that peri-
od they became more
abundant in both North
America and Europe,
reaching their highest
expression in the beaver,
and spread into South
America. (Figs. 48-49.)

The Myomorpha
(mice, rats, etc.) have a
few supposed represen-
tatives in the Eocene of
Patagonia, but only cer-
tainly begin in the Neo-
cene in the Northern
Hemisphere. The dor-
mice are peculiar to Eu-
ropean beds, and the

Fic. 49.—Plesiartomys delicatissimus Leidy ; pocket-gophers to North

a, humerus; b, proximal ends of ulna and radius; America. They cover
2, d, tibia, distal end; e, /, astragalus ; all natural : :
: : ae the earth, including a

size. From Eocene of Wyoming.
few forms in Australia,

2

at the present period. (Fig. 50.)

The Lagomorpha (rabbits) appear first in the lowest Neo-
cene of North America (White River), and continue to the
present day. They appear in Europe at about the same
time, but not in South America until the Plistocene. (Fig.
51.)

The Curroprera (bats) includes two primary divisions at
the present time, the insectivorous (Animalivora) and the
frugivorous (Frugivora), which differ as follows:

Crowns of molar teeth Vs; those of opposite

114 COPE

jaws interlocking in mastication ; Animalivora.
Crowns of teeth obtuse, not interlocking in

mastication ; Frugivora.

Fig. 51.—Paleolagus haydenii Leidy;

Fia. 50.—Entoptychus crassiramis a rabbit from the lower Neocene of Col-

Cope; a pocket-gopher from the Middle orado. Original; natural size; a, ante-

Neocene of Oregon, naturalsize. Origi- rior half of skull, from below ; b, ¢, lower

nal. c, lower jaw from above. jaw; d,tibia and fibula, from front; e,
tibia and fibula, from below.

Animalivora appear in typical forms in the Lower
Eocene (Wasatch) in North America, and the Upper Eocene
(phosphorites) in France.

The Frugivora (flying foxes) present a degenerate denti-
tion; they are not known in the fossil state.

The BunorHERrtIA present four suborders, which are char-
acterized as follows:

Molars 8 or more; incisors several, with closed

roots ; Pantotheria.
Molars 7 or less; incisors with closed roots,

not enlarged ; fibula distinct; otic bulla

developed ; Creodonta.

VERTEBRATA 115

Molars 7 or less; canine generally small, in-
cisors generally enlarged; fibula generally
united with tibia ; Insectivora.

Molars 7 or less; incisors few, growing con-
tinually from open roots ; Tillodonta.

Incisors much enlarged, growing from persis-
tent pulps, the superior with enamel in
anterior and posterior bands, and hence
truncate; * Taeniodonta.

The Pantotheria are the most ancient of the Eutherian
Mammalia, appearing rather abundantly in the Jurassic of
Europe and North America. They are all of small size.
The Creodonta represent the Carnivora in the Postcretacic
and Eocene systems. They have (except in one family)
more than one sectorial molar, and these are true molars,
and not premolars. They have (except in one family)
strong canines. They range in size from that of an opos-
sum (Stypolophus sp.) to that of a grizzly bear (Hemipsalo-
don sp.). They are found in the horizons mentioned in
North America, Europe, and South America. A few species
(Hyzenodon, etc.) remain over in the Lower Neocene
(White River). (Fig. 52.)

Undoubted Insectivora appear in the Upper Eocene of
France, but supposed members of the suborder occur in the
Lower Eocene of the same. In America none are certainly
known prior to the Lower Neocene (White River). They
are rare in all formations in America, but they are abund-
aut in the Middle Neocene of Europe, and in later beds,
where forms of moles, shrews, and hedgehogs are abundant.
The enlargement of the incisors seen in the Insectivora
reaches its extreme in the Tillodonta, where they grow
from persistent pulps, asin the Glires. These animals first
appear in the Postcretacic of North America, and are not

*Tertiary vertebrata, 1883, p. 186. This order is equal to the Calamodonta of
Cope and the Ganodonta of Wortman.—ED.

116 COPE

FIG. 52.—Deltatherium fundaminis Cope; portions of skull, two-thirds -
natural size. From Puerco beds of New Mexico. a, skull profile; b, from
below: c, fragment of lower jaw, from above; d, do., from outer side.

Fie. 53.—Calumodon simplex Cope; lower jaw, one-third natural size.
From Wasatch Eocene of Wyoming. 0, from above; cd, an isolated molar..

VERTEBRATA 117

uncommon in the Eocene of America, with one form in the
Eocene of Switzerland. Later than that age they are un-
known. (Fig. 53). The Creodonta are probably the an-
cestors of the Carnivora and Insectivora, and the Tillodonta
of the Glires. The Pantotheria are ancestors of the Creo-
donta. The time relations of the Bunotherian orders are
expressed in the following table:

Pantotheria Tnsectivora Tillodonte

Plistocene

Neocene

Eocene

|

|
Cretacic . . . . | ? | 9
F |
Jurassic |

Triassic

Carbonic . is os |

Devonic . .

Silurie. ....

Ordovicic

Cambric. .. .

Huronic . .. . |

True Carnivora are definitely known from the beginning
of the Neocene, and it is probable that they already existed

118 COPE

in the Upper Eocene in Europe. There are two suborders,
which differ as follows:

Digits distinct ; posterior limbs free ; Fissipedia.

Digits united into paddles by integument;
hind limbs partly enclosed in general in-
tegument ; - Pinnipedia.

The Fissipedia have their closest connection with the
Creodonta, where the latter exhibit exceptionally a denti-
tion like the dogs, in the family Miacide. From the dogs
development has pursued two principal lines. The one has
tended to an omnivorous diet, and has terminated in the
bears. The other has tended to an exclusively carnivorous
diet, and has ended in the cats, which display the charac-
ters of the order in the greatest perfection. True bears
appear in the Plistocene, and cats (Fig. 54) and hyenas in
the Upper Miocene, in the Northern Hemisphere. In the
Southern Hemisphere cats do not appear until the Plisto-
cene (Fig. 55), and hyzenas do not occur in the Western

Fie, 54.—Nimravus gomphodus Cope, two-fifths natural size; left side of
skull. From Miocene of Oregon. 3-4, premolars; 1-2, true molars.

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120 COPE

Hemisphere. Dogs are very abundant throughout the
Neocene, except in the southern continents, where they do
not appear till the Plistocene (Fig. 37).

The Pinnipedia appear later in geologic time than the
Fissipedia, no forms being known of an age prior to the
Middle Neocene. The earliest forms are of two types re-
lated to the true (earless) seals and walruses respectively.
Of the primitive affinities of the Pinnipedia nothing is
known, but they are suspected to have had connection with
the Creodonta.

The CHALICOTHERIA is a group which includes but few
species, which have been found in India, Europe, and
North America. They possibly appear in Europe in the
Upper Eocene, and occur in the successive stages of the
Neocene, beyond which they did not continue. Their
characters are very peculiar, including long fore legs and
short hind ones, with claws and digits like sloths, but den-
tition like perissodactyle ungulates. The largest species
equaled a grizzly bear.

The order Taxroropa includes the following suborders :

I. ? No clavicle.

Astragalus not interlocked laterally with the
‘tibia; fibula not articulating with cal-
caneum; head of astragalus rounded ;
canines ; no anapophyses; Condylarthra.

Astragalus not interlocked with tibia; fibula
articulating with caleaneum ; astragalus
head flat; canines ; Litopterna.

Astragalus interlocking at side with tibia ;
fibula not articulating with calcaneum ;
head of astragalus flat; no canines; Hyracoidea.

II. Clavicles present.

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VERTEBRATA 121

Incisors growing from persistent pulps;

anapophyses ; Daubentonioidea.*
Incisors with closed roots ; anapophyses ; Quadrumana.
Incisors with closed roots; no anapo-

physes ; Anthropomorpha.

The Condylarthra are characteristic of the Postere-
tacic in North America, and the Lower Eocene of both
continents. They have not yet been detected in the
Southern Hemisphere. Their characters connect insepar-
ably the Ungulata and Unguiculata divisions, since the
carpus and tarsus are like those of the latter, while the
ungues are hoofs or semihoofs. The molars are trituber-
cular, quadritubercular or lophodont. The best known
genus is Phenacodus (Fig. 56), which is the synthetic type
of all ungulates. In the known Condylarthra there are
five digits on all the feet.

The Litopterna are only known, so far, from the Cenozoic
of South America, ranging throughout the entire series.
The teeth present the variations seen in the Perissodactyla,
some being bunodont and some lophodont, but the buno-
dont forms are tritubercular, like the lowest Condylarthra.
This group displays a remarkable reduction in the digits
as in the Perissodactyla, passing to three and one digit, as

in the horse line.
The Hyracoidea are only known in the recent state in

Africa and Western Asia. Their molar dentition is lopho-
dout, while the incisors approximate somewhat the rodent
pattern. In like manner Daubentonioidea are only known
as recent in Madagascar. They are allied to the lemurs.
Quadrumana appear in the Lower Eocene in North

America and Europe as lemurs (Fig. 57). True monkeys
do not appear until the Middle Neocene in the Old World,

while they are absent from North America, and occur in
the Plistocene of South America. The Anthropomorpha
(Fig. 58) appear first in the Middle Neocene of France and

* The horny coverings of the terminal phalanges of all but the first digit in the
only known genus, are claw-like.

122

COPE

Fia. 57.—Necrolemur antiquus Filh; alemur from the Upper Eocene of Fraiee,

natural size.

From Filhol,

the Upper Neocene of India, but man does not appear

until the Plistocene in both America and Europe.

The

exact stage of the Plistocene at which man’s remains have
been found is not clearly ascertained, but he was contem-
porary with various extinct species and genera of Mam-

malia in both Europe and North and South America.

The

time history of the Taxeopoda may be expressed as follows :

Plistocene

1

2

3

4

5

6

Neocene-

Eocene .

Cretacic

Jurassic

Triassic .

Carbonic

Devonic

Siluric .

Ordovicic

Cambric

Huronic

VERTEBRATA 128

The Toxoponrra are confined to the South and Central
American continents. They appear first in the Eocene of
Patagonia and continue through the Plistocene system.
These later representatives of the genus Toxodon reached
the size of the largest species of rhinoceros. The teeth are
lophodont, and early began to be prismatic, and to grow

Fie. 58—Dryopithecus fontani Lart.; anthropoid ape from Middle Neo-
cene of France; ramus of lower jaw, natural size. From Gaudry.

from persistent pulps. This condition appeared first in the
incisors, some of which resemble somewhat those of ro-
dents, and later in the molars. There are two suborders of
Toxodontia which differ as follows :

Femur with third trochanter ; a clavicle ; Typotheria.

Femur without third trochanter; no clavicle; Barytheria.

The known species of Typotheria have five anterior, and
four or five posterior digits, and they are of medium and

124 COPE

small size. The known Barytheria have three digits on
each foot, and the later forms reached large proportions,
with heavy body and short legs.

Fre. 59.—Coryphodon elephantopus Cope ; Skull; from below, 2-9 natural size.
Original. From the Wasatch Eocene of New Mexico,

The AMBLYPODA are, restricted to the Northern Hemis-
phere, and to the Postcretacic and Eocene systems, where
their remains abound both in Europe and North America.
They are the only order of Ungulata with superior molars
constructed on a tritubercular basis, and the inferior molars
on the tuberculo-sectorial pattern. The earlier forms from
the Puerco are the smallest; those of the Lower Eocene
(Suessonian, Wasatch) are next in size (Fig. 59); while
those of the Middle Eocene (Bridger), which have been
found in North America only, are often.of gigantic size
(Dinocerata) (Fig. 60), and have the cranium adorned with
horny processes.

There are three well-marked suborders of this order,
with the following characters :

Astragalus with a head ; femur with third
trochanter ; superior incisors present; Taligrada.

Astragalus without head; a third trochanter
and superior incisors ; Pantodonta.

VERTEBRATA 125

Astragalus without head ; no third trochanter
nor superior incisors ; Dinocerata.

The Taligrada are confined to the Postcretacie epoch ;
the Pantodonda to the Lower, and the Dinocerata to the
Upper Eocene.

The order Prososcrpr is unknown prior to Middle Neo-
cene time in the northern part of the Eastern and Western
Hemispheres, and late Neocene or Plistocene time in South
America. In other parts of the Southern Hemisphere they
are unknown, although one of the two living species (Lle-
phas africanus) is restricted to Africa (Fig. 60). The earliest
forms of Proboscidia are Dinotheria and Mastodons (Fig 61),
which reached huge dimensions. Elephants appear in late
Neocene times in India, and spread in later epchs over
Noath America and Europe. In America their remains are

Fic. 60 —Elephas indicus L; superior molar tooth, reduced. From Tomes,
after Owen.

not known from south of the valley of Mexico. The hairy
mammoth (Elephas primigenus Blum.) was a contemporary
of prehistoric man in the Old World, and probably in the
New.

The DiptartHra include the most specialized types’ of
Mammalia as regards the structure of the skeleton, denti-
tion, and digestive system, but they are inferior to the an-
thropomorphous suborder of the Taxeopoda in the structure
of the brain.

There are two suborders, as follows:
Feet with the third digit of predominating

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Fig, 62.—Lozolophodon cornutus Cope ; skeleton, restored, 1-20 natural size. Original, From the Bridger Eocene of Wyoming.

128 COPE

dimensions; distal end of astragalus not
forming a ginglymus; Perissodactyla.

Feet with the digits 3 and 4 predominating
and subequal; astragalus with a distal
ginglymus ; Artiodactyla.

The PrerissopacryLa exhibit various series of forms in
which the toes diminish in number, from four in front and

Fie. 63.—Aphelops meyalodus Cope, one-sixth natural size; B, inferior
view of cranium, Original. Hornless rhinoceros from the Loup Fork Neo-
cene, Colorado.

: N. A. Original,
. D. Cope.

130 COPE

three behind, to one on all the feet (the horse). They may
be considered under two heads with respect to the structure
of their superior molars, viz.: first those in which the exter-
nal wall of the crown does not form two Vs (Rhinocero-
toidea), and those in which such Vs are present, with the
angles directed inwards (Equoidea). Each series includes
several families, mostly extinct; in the former the rhinoceros
(Fig. 63) and tapir, and in the latter the horse, are still
living. The earliest forms belong to the former division,
and some of them (Hyracotherium, Fig. 64) have the super-
ior molars almost quadritubercular. They appear first in
the lowest Eocene. Three-toed horses first appear in the
Lower Neocene, and one-toed horses in the Upper Neocene.

The Artiodactyla are represented by a great variety of
forms, which differ primarily in their dental characters.
Thus the oldest type (Trigonolestoidea) has tritubercular
superior molars. Of the remainder, one series (Sudidea)
have the molars quadritubercular, or more highly tubercu-
lar. The remaining types have the tubercles of the molars
more or less flattened on one side, so as to give crescentic
figures on section (Fig. 65), and are hence called seleno-
dont. In some of these there are five crescents (Anthraco-
theroidea) ; in the other there are only four. The latter may
have most of the premolars simple (Cameloidea, Fig. 65) or
complex (Bodidea). Some types of the former and all of the
latter lack superior incisor teeth, and have the cuboid and
navicular bones codssified. The Bodidea culminate in forms
with horns or bony processes of. the skull, which may be
permanent (Bovide, or annually shed (Cervide). These
divisions appear in geological time in the order of structu-
ral modification as here mentioned. This appearance is
represented, together with the divisions of the Perissodac-
tyla, in the following table:

131

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VERTEBRATA

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132 COPE

Fig. 65.—Protolabis transmontanus Cope ; skull, one-third natural size, A
Camelvid from the Ticholeptus bed of Oregon. a, from left side; b, from below.
Original.

Of the superfamilies which continue to the present day
the existing species are few in number, with the exceptiom
of the Bodidea. All except the Bodidea are now restricted
to tropical and subtropical regions, and while the majority
of Bodidea have the same distribution, there are many
species in temperate regions, and a few dwell within the-
Arctic Circle.

The above table includes only the most important of the-
superfamilies of the Diplarthra. Those of the Perissodac-
tyla are equally distributed on both sides of the Northern
Hemisphere, but only modern forms of the Equoidea appear:
in South America and in late Neocene time. Pantolestoi-
dea have been so far found in North America only, and
the other superfamilies are relatively rare in that continent,
except the Cameloidea, which are more abundant than im
the Old World. This and the Bodidea only appear in South.
America in the late Neocene,

As in the Perissodactyla there is a reduction of the digits.

VERTEBRATA 133

in most of the lines of the Artiodactyla, which reaches its
extreme in the Cameloidea, and in the Bodidea. In the
most specialized types of these superfamilies but two digits
remain, and the metacarpals of these are fused into a single
bone, the “cannon bone.” This structure first appears in
time in the latest Neocene. (Fig. 66.)

Fic, 6s.—Anterior feet of Artiodactyla, with both series of carpals, except in
No. 4. From Kowalevsky. No.1, Hippopotamus; 2, Hyopotamus; 3, Dorcathe-
rium: 4, Gelocus ; 5, Cervus.

The families embraced in the orders of Eutherian Mam-
inalia are the following :

Marsuprarra ; (Polyprotodontia) ; Triconodontide, Amphi-
theriidee, Myrmecobiide, Dasyuridee, Didelphide, Per-
amelidee ; (Diprotodontia) ; Phascolomyide, Phalangist-
ide, Tarsipedide, Diprotodontidee, Macropide, Thyla-
coleonide. :

Ceracea; (Archeocet); Zeuglodontide ; (Odontoceti) ; Squal-
odontide, Platanistide, Physeteride, Del phinidee, (Mys-
tacocett) ; Baleenide.

StreNIA; Prorastomide, Halitheriide, Manatidee, Halicori-
de, Rhytinide.

1384 COPE

Epenrata ; Orycteropodide, Manide, Bradypodide, Mega-
theriide, Myrmecophagide, Dasypodide, Glyptodon-
tide.

Guires; (Hystricomorphx) ; Paradoxomyide, Hystricide,
Echinomyide, Octodontide, Capromyide, Caviide,
Chinchillidee ; (Sciuromorpha) ; Sciuridee ; (Myomorpha) ;
Dipodide, Muride, Myoxide, Saccomyide, Microtide,
Lophiomyide, Bathyergidee; (Lagomorpha) ; Leporidee.

CHIROPTERA ; (Animalivora) ; Phyllostomide, Desmodontide,
Rhinolophidee, Noctilionide, Vespertilionide, Embal-
lonuride; (Frugivora) ; Pteropide.

BunoTHERIA ; (Pantotheria); Amblytheriide ; (Creodonta) ;
Mesonychide, Esthonychide, Arctocyonide, Miacide,
Hyeenodontide ; (Insectivora) ; Leptictide, Centetide ;
Galeopithecide, Tupzide, Solenodontide, Macrosceli-
didee, Talpidee, Adapisoricide, Mythomyide, Scalopide,
Chrysochloride, Erinaceide, Myogalide, Soricide ;
(Tillodonta) ; Tillotheriide ; (Teniodonta); Ectoganide,
Stylodontide.

Carnivora; (Fissipedia) ; Cercoleptidee, Procyonide, /Elu-
ride, Canide, Bassaridide, Mustelide, Protelide, Arc-
ticidee, Viverride, Cynictide, Suricatide, Cryptoproc-
tide, Nimravide, Felide, Hyenide; (Pinnipedia) ;
Phocide, Otariide, Odobeenide.

CHALICOTHERIA ; Chalicotheriide.

Taxeopopa ; (Condylarthra) ; Phenacodontide, Pleuraspido-
theriidee, Meniscotheriide ; (Litopterna) ; Proterotherii-
de, Macraucheniide, Astrapotheriide ; (Hyracoidea) ;
Hyracide ; (Daubentonioidea), Chiromyide, Mixodec-
tidee ; (Quadrumana) ; Adapide, Anaptomorphide, Tar-
siide, Lemuridee, Hapalide, Cebide, Cercopithecide;
(Anthropomorpha) ; Simiidee, Hominide.

Toxoponira ; (Typotheria), Atryptheriide, Interatheriide,

VERTEBRATA 135

Protoxodontide, Mesotheriidee ; (Barytheria) ; Xotodon-
tide, Toxodontide.

Prososcrpra ; Dinotheriide, Elephantide.

AmBLypopa ; (Taligrada) ; Periptychide, Pantolambdide ;
(Pantodonta) ; Coryphodontide ; (Dinocerata) ; Uinta-
theriide.

DipLartHRa; (Perissodactyla) ; Lophiodontide, Triplopodi-
dee, Ceenopide, Hyracodontide, Rhinoceride, Tapiride,
Lambdotheriide, Menodontide, Paleotheriide, Equi-
de ; (Artiodactyla,) ; Trigonolestide, Eurytheriide, An-
oplotheriide, Dichobunide, Czenotheriide, Anthraco-
theriidee, Xiphodontidee, Suidee, Hippopotamide, Mery-
copotamidee, Dichodontide, Oreodontide, Poebrotherii-
dee, Protolabidide, Camelide, Eschatiide, Tragulide,
Moschide, Bovide, Cervide.

ERRATA.

Page 95, 5th line from bottom, for acromyoctian read
acromyodian.

Page 124, 8th line from bottom, for Fig. 60 read Fig. 62.

Page 134, under BunorHerta, transfer Leptictide and
. Centetidee from Insectivora to Creodonta.