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 Syllabus of lectures on the vertebrate / 
 
 3 1924 024 781 704 
 
Cornell University 
 Library 
 
 The original of tiiis book is in 
 tine Cornell University Library. 
 
 There are no known copyright restrictions in 
 the United States on the use of the text. 
 
 http://www.archive.org/details/cu31924024781704 
 
publications 
 
 OF THE 
 
 iriniversit^ of pennsi^lvania. 
 
 Syllabus of Lectures 
 
 ON THE 
 
 VERTEBRATA 
 
 BY 
 
 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. 
 
Press of Levy type Co., Engravers and Printers, Philadelphia. 
 
THE LIFE AND WORKS OF COPE 
 
 ILLUSTKATING THE TRAINING OF A NATURALIST AND THE 
 
 ESSENTIAL CHARACTERISTICS OF A GpEAT 
 
 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. As a 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 Pirn 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. V 
 
 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 scle»" 
 otic plates look at the eye — thee will see these in it." Av 
 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 : 
 
 " I 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. I am 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 Spelerpinse, 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 ours) 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 
 gliUinosum — 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 tliat lie was sensitive to being laughed at for 
 his interest in these animals. 
 
LIFE AND WORKS OF COPE. Vll 
 
 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,f 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 Salamandridse, 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 
 
 fA word used by French writers of the time to express lines of evolutionary 
 descent. 
 
via 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 Lselaps. 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, Pmiheus, Platecarpus, Clidastes, Compsemys 
 (type of the Amphichelydia, Lydekker), Protostega, and 
 Agaihaumus. 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 zoologists. 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 COrE. XI 
 
 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, studj'ing 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- 
 oiDBS had been originally introduced into zoology by Agas- 
 
Xll LIFE AND WOEKS 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 Miiller 
 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- 
 ble 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 pentadactyl 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. 
 11- P- ^■) 
 
 * Fishes having the mouth surrounded hy specialized membrane bones as in the 
 Teleosts. 
 
LIFE AND WORKS OF COPE. XUI 
 
 "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- 
 lachii (Cuvier, 1817), the Dipnoi (Miiller), the Crossopterygii 
 (Huxley) with paired fins arranged so as to form a fringe 
 around a central lobe, and the Adinopteryi. 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 vertebrae for the 
 combinations of Physostomi* 
 
 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 
 and 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 WOKKS OF COPE. 
 
 of living types he broke up into a number of groups by an 
 originality of analysis as marked as in 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- 
 achostomi, Scyphophori, Pledospondyli, Haplomi, Enchelyceph- 
 ali, Colocephali, Percesoces and Hemibranchii. 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 Pledospondyli and the separation of the 
 eel-like fishes into different groups. 
 
 The order Pledospondyli (fishes with the anterior verte- 
 brse coalesced, as distinguished from Isospondyli (Cope) in 
 which all the vertebrae are separated) was framed for the 
 numerous fresh-water fishes comprised in the families Ga- 
 tostomidse (Suckers), Cyprinidse, (Carps, etc.,), Cobitidse 
 (Loaches), Sternopygidx (Carapos, etc), and the numerous 
 South American and African fishes representing the Cyprin- 
 oideans, constituting the families Characinidas and Ery- 
 thrinidse. 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 
 
 * This section upon the classification of the Teleostei is directly quoted from Pro- 
 fessor Gill's Memorial Address before the American Philosophical Society as far as 
 the middle of page xyi, A number 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 Pledospondyli 
 were the " anterior four vertebrae much modified, and with 
 ossicula auditus," also the presence of a prgecoracoid arch. 
 But from this group he differentiated two large orders,* 
 distinguished by simple anterior vertebrae, including the 
 form (Gymnotus, the electric eel) which had been pre- 
 viously placed in the same family with the Sternopygidse, 
 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 "vertebrse unmodi- 
 fied " was the extreme reverse of the truth ; they were found, 
 in fact, to have the first four vertebrse 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 vertebrse " 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 prsecoracoid." Again, the com- 
 bination of " four vertebras co-ossifled, and with ossicula auditus " was attributed 
 to another order {Nematognathi), but within the same order a group was admitted 
 with the attribute of " vertebrse unmodified," although it was remarked that the 
 fishes in question, "the Hypophthalmidw, are indeed scarcely to be referred to this 
 order." 
 
LIFE AND WORKS OF COPE. 
 
 successively dissipated, and the order Fledospondyli, modi- 
 fied by the addition to it of his Qlanencheli, 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 Linnsean order 
 Apodes — an order that is Linnsean, however, only in name 
 and in type. 
 
 Professor Gill closes with the following critique : 
 
 " I cannot consider his removal of the ' subclass Dipnoi ' to a posi- 
 tion between the Holocephali and his Elasmohranchii {Plagiostomi,) 
 or his combination of his old subclass Orossopterygia with his sub- 
 class Adinopteri as improvements. But the subdivision of the 
 Crossoptergyia into Rhipidopterygia and Orossopterygia (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. XVll 
 
 Fossil 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 tlie labors of Owen, Huxley, and 
 Cope. Cope's interest in the ancestry of the fishes was 
 naturallj' intensified as he became a more convinced evolu- 
 tionist, and b}' 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 vertebi'ate 
 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 Palaeozoic 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- 
 
XVIU LIFE AND WOEKS OF COPE. 
 
 ing the bearing of the foregoing facts upon published 
 schemes of classification of the Elasmobranchii, 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 Palseozoic 
 Pleuracanths with the skulls of the so-called Didymodus, 
 Cope, from the Permian of Texas. (Catalogue, Pt. I., p. 
 xxiii.) 
 
 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 Asterolepidss." (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 Tarrasiidse. 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 Adinistia 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 Extinct 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- 
 Tfiary Divisions of the Salmdndridas, with description's 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 ; II. Bufoni- 
 formia ; III. Arcifera ; TV. Raniformia. 
 
 At the age of 25 he described his first extinct Amphibiar 
 
XX LIFE AND WOBKS 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 {Cceci- 
 lia), Proteida {Nedurus- 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 Air-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 tlie typical coal measures and Permian ex- 
 tended to Iowa and Illinois, leading to the determination 
 of Cricotus, which in 1880 f 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 rnade 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 vertebrae, but 
 also the great primitive representatives of the Reptilia. 
 The suborders Rachitomi and Emboloineri have been 
 grouped as Temnospondyli in contrast with the specialized 
 Lahyrinthodontia 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 pres^ious contributions, but he anticipated that 
 
 » Trans. Amer. Phil. Soc, read 1868, pub. 1869. See also Proc. Phila. Aead. Nat. 
 Sci., 1868, p. 211. 
 
 t 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. As a 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 vertebrae of living amphibia are composed of inter- 
 centra and are, therefore, not homologous with the true 
 centra (pleurocentra) of reptiles, birds, and mammals. 
 
 * " The Batrachia of North America." Bull. No. 34, U. S. Nat. Museum. 
 
XXll LIFE AND WORKS OF COPE. 
 
 Classification of the Reptilia by Characters op 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 SqvM- 
 mata* a group proposed by Oppel to include the lizards 
 and snakes. Two years later he made his first remarks 
 about the Dinosaur Lselaps,f and in 1867 he compared 
 the carnivorous Dinosaurs with the birds. J 
 
 " 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 Archceopteryx presented points of affinity. The second, 
 and one not less striking, is by the Dinosauria of the orders Oonio- 
 
 » Proo. Acad. Phila., 1864, p. 224. 
 
 t IjCelaps aquilunguis. Cope. Proc. Acad. Nat. Sci., Phila., July, 1866, pp. 275- 
 279. 
 
 t Ibid., 1867, pp. 234-235. 
 
LIFE AND WORKS OF COPE. XXUl 
 
 poda and Symphypoda. He showed the essential differences between 
 the ordinary Dinosauria 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 Lcelaps, 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 Batkygnathus, 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 Dinosauria were pubes, having a position similar to those 
 of the Orocodilia. 
 
 Also, that a species of Lcelaps had been observed in France, by 
 Cuvier, which was different from the Lcdaps aquilunguis, and which 
 he proposed should be called Lcelaps gallicus. 
 
 Gompsognathus, 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 vertebrae 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 WOEKS OF COPE. 
 
 the long-tailed Ratitce (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 
 ctos."t 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 Ehynchocephalia 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. Ichthyopttrygia : Order Ichthyopterygia. 
 
 B. Extremities differentiated. 
 
 II. Streptostylica : Orders Lacertilia, Pythonomorpha, 
 Ophidia. 
 
 III. Synaptosauria : Orders Rhynchocephalia, Testudi- 
 nata, Sauropterygia. 
 
 IV. Archosauria : Orders Anomodontia, Dinosauria, 
 Grocodilia, Ornithosauria. 
 
 In 1875 the large volume " 21ie Vertebrataof the Cretace- 
 ous Formatio7is of the West " appeared, as Vol. II of the 
 Rep. U. S. Geol. Surv. Territ. (302 pp., PI. LVII.). This 
 work contains extensive descriptions, especially of the 
 
 ♦Popular Science Review, 1868, pp. 237-247. 
 tProe. Assoc. Adv. Sci., XIX, pp. 191-247. 
 
LIFE AND WORKS OF COPE. XXV 
 
 Mosasaurs, also of Testudines, Crocodilia, Plesiosaurs and 
 Dinosaurs (Iguanodontia) : Agathaiimas, Cope {Triceratops, 
 Marsh); Hadrosaurus, Leidy; (Trachodon, Leidy; Didonius, 
 Cope, Laosaurus, 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 Pelycosaurla 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 Pelycosaurla, 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 Diadectidas. In 
 a skull of Empedias he described two occipital condyles, 
 being misled by the missing basioccipital. In 1883t he 
 placed his genera Pariotichus and Pantylus in a new family, 
 Pariotichidse, characterized by the over-roofing of the 
 temporal fossae and the presence of the supra-occipital and 
 par-occipital plates (intercalare. Cope). He now found the 
 basioccipital in position and the order Cotylosauria was 
 given up. In 1890 (March 12th), Cope, however, again em- 
 ployed the Cotylosauria as a suborder under the Theromora 
 distinguishing three families: Careiasauridse, Pariotichidse, 
 Diadectidse. In 1882J 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. 
 tProc. Amer. Philos. Soc, p. 631. 
 tProc. Boy. Sou., Vol. 44, p. 383. 
 §Trans. Am. Phil. Soc, Vol. XVII. 
 
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, Otocoelidse, 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. J The MSS. for this 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. Philoc Soc, Vol. XXXIV, 1896 (Feb. 2d), pp. 436-457, Pis. VII-X. 
 
 tibid., Vol. XXXV, pp. 112-139, Pis. VII-X, Aug. 15, 1896. 
 
 {Many preliminary papers have appeared for this publication, of which the 
 following are named : 
 
 " An analytical Table of the Genera of Snakes." Proc. Am. Philos. Soc, 1886 pp 
 479-499 
 
 " The Osteology of the Lacertilia." Proc. Am. Philos. Soc, Vol. XXX, 1892, pp. 
 185-221. Pis. II- VI. < 'ff 
 
 "On Degenerate Types of Scapular and Pelvic Arches in the Lacertilia." Journ. 
 Morphol., Vol. VII, 1892, pp. 233-2-14, PI. XIII. 
 
 "The Classification of the Ophidia." Trans. Amer. Philos. Soc, Vol. XVIII. 
 April 15, 1895, pp. 186-219, Pis. XIV-XXXIII. 
 
 " On the Hemipenes of the Sauria." Proc. Acad. Nat. Sci., Phil., August. 1896, 
 pp. 461-467. . . B . 
 
LIFE AND WORKS OF COPE. XXVll 
 
 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 ior 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 as a 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 
 
XXVlll LIFE AND WORKS OF COPE. 
 
 will be fouad to correspoad 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.) 
 
 Evolution of the Mammals. 
 
 Cope's most numerous and voluminous vs^ritings 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 Solipedes, 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 Perissodadyla 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 Palaeontology, 
 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 bj^ Lartet, and has been subsequently elaborated 
 and demonstrated by Marsh. 
 
 II. — 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 
 Faloplotherium 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 Palseontology. 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. Clinodadyla. 
 
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, firmlj' 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 Trituhercular 
 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 — namel}^ 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- 
 
XXXII 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 force and application of the tritubercular 
 law Cope clearly perceived, but left to others fully to work 
 out and 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 cla>ved, 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 Plagiaulacidse and defined 
 the order MuUituberculata. That from the Wasatch is- 
 perhaps next in value, and in succession rank his contribu- 
 
LIFE AND WORKS OF COPE. XXxiii 
 
 tions from the John Day, Loup Fork, Blanco, Palo Duro and 
 Port Kennedy Bone Cave. 
 
 As an explorer he had marked success, finding the unique 
 skeleton of Hyrachym, of Loxolophodon, a name which was 
 telegraphed to the American Philosophical Society, and 
 converted by the operator into Lcfalophodon. 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 Aiichitheriidse, Felidse and Canidse. 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 Oreodontidse. 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 (Jreodonta, 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, vertebrae, 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- 
 2)hodon and Uintatheriiim. 
 
 As to his scientific character apart from his genius, which 
 
LIFE AND WORKS OF COPE. XXXV 
 
 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 Palseonictis, Hyasnodon, Ardocyon, 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 Soienoe, also upon valuable materials which have been 
 furnished by Professor Theodore Gill, Cope'S^-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. Percy Moore. 
 
Syllabus of Lectures 
 
 ON THE 
 
 VERTEBRATA 
 
CONTENTS. 
 
 NTRODUCTION 
 
 Superclass I. 
 
 Hemichorda 
 
 Class I. 
 
 Helminthophya 
 
 Superclass II. 
 
 Urochorda 
 
 Class I. 
 
 Tunicata 
 
 Superclass III. 
 
 Cephaloghorda 
 
 Class I. 
 
 Acrania . 
 
 Superclass IV 
 
 Craniata 
 
 Class I. 
 
 Agnatha 
 
 Subclass I. Ostracophori 
 
 PAGE 
 
 7 
 
 9 
 9 
 
 10 
 . 10 
 . 11 
 11 
 12 
 14 
 15 
 
 Subclass II. Cyclise . 16 
 
 Subclass III. Marsipobranchii 17 
 
 Class II. Pisces . .17 
 
 Subclass I. Elasmobranchii .... 21 
 
 Subclass II. Holocephali . . 28 
 
 Subclass III. Dipnoi . . 29 
 
 Subclass IV. Teleostomi . 30 
 
 Class III. Batrachia 43 
 
 Subclass I. Stegocephali . .43 
 
 Subclass II. Urodela . 4S 
 
 Subclass III. Salientia . . 50 
 
 Class IV. Monocondylia .52 
 
 Subclass I. Reptilia . . . . 52 
 
 Subclass II. Aves . . . 83 
 
 Class V. Mammalia . 102 
 
 Subclass I. Prototheria 102 
 
 Subclass II. Eutheria . .... 104 
 
PREFACE. 
 
 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 M'as 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 division is 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 
 
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. 
 
 Fig. ]. — Diagrammatic representation of primitive and derivative types of lateral 
 and median fins. A^ primitive condition, fins continuous; B, derivative condition, 
 flns distinct and specialized. D, dorsal fin; BrF, pectoral; BF, ventral; AF, anal; 
 8F, caudal; RF, dorsal; FF, second dorsal fin. From Wiederslieim. 
 
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 I.— HEMICHORDA. 
 
 Helminthophya. 
 
 There is but one class of Hemichorda: 
 Not metameric ; no atrium ; 
 
 Class I.— HELMINTHOPHYA. 
 
 Two orders of this class are known which differ as fol- 
 lows: 
 
 *Tlie Vertebrata (or Chordata) as now extenfled, may be defined as coelomate 
 Metazoa, possessing a longitudinal, axial, cellular supporting rod (tlie notochordj. 
 derived from the hypoblast (except Hemichorda ?) 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, stigmatse), 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- 
 chiae ; 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 Enteropneusta includes but one family, 
 the Balanoglossidse, 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 Cephalodiscidee with gill-slits, and the Rhabdopleuridse 
 in which gill-slits are wanting. 
 
 No extinct forms of Helminthophya are known. 
 
 Superclass II.— UROCHORDA. 
 
 There is but one class of Urochorda : 
 
 Not metameric ; a mantle covering the body ; 
 respiration pharyngeal ; heart distinct, 
 saccular ; Tunicata. 
 
 Class I.— 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 ; Ascidise. 
 
 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 Aseidiae are degenerates from 
 the vertebrate line. The families are : 
 
 Perbnnichoedata : Appendiculariida'. 
 
 Thaliacea : Salpidfe, Pyrosomidse. 
 
 AsciDi/E : Botryllidfe ; Ascidiidas ; Clavelliuidae. 
 
 Superclass HI.— 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 Branchiostomidse (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 lower jaw ; man- 
 dible not segmented ; coracoid generally 
 not distinct ; ankles between propodial 
 bones and carpus or tarsus ; heait with 
 four chambers ; Mammalia. 
 
 The period of appearance and duration of each of these 
 classes is exhibited in the accompanjdng 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 
 
 COPK 
 
 
 Agn 
 
 atba 
 
 Pisces 
 
 Batrachia 
 
 Monocon- 
 dylia 
 
 Mammalia 
 
 Plistocene . . . 
 
 
 
 — 
 
 
 
 
 
 
 
 
 
 Neocene 
 
 
 
 
 Eocene . 
 
 
 
 
 
 
 
 
 
 
 
 
 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; Cydias. 
 
VERTEBEATA 
 
 No osseous skeleton nor limb-like append- 
 ages; 
 
 15 
 
 Marsipobranch ii. 
 
 Of the Agnatha, extinct species of the subclasses Ostraco- 
 phori and C3'clia3 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. 
 
 j\ntiarcha. 
 
 Carbonic. . . . . 
 
 
 1 
 
 1 
 
 Devonic ... 
 
 
 
 
 Siluric . . . 
 
 
 
 
 
 Ordovicic 
 
 
 
 
 
 Cambric 
 
 
 
 
16 
 
 COPE 
 
 Fig. 2. — Bothriolepis canadensis Whiteaves; bead and carapace, Iroin above, and bead from 
 below. From the De-vonie of New Brunswick. From Whlteaves. (Antiareha.) 
 
 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.— CYCLIiE. 
 
 Median diphycercal caudal fin ; brain case 
 
 not segmented ; Saccocephala. 
 
 The order Saccocephala is the only one known to per- 
 
VERTEBEATA 17 
 
 tain to the Cyclise. It embraces the simple family of the 
 Paleospondylidse 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 Hypeeotreti belong two families, the Myxinidse 
 and the Bdellostomidse; to the Hyperoarti one, the Petro- 
 myzontidse. 
 
 No extinct species of Marsipobranchii are known. They 
 include only the recent lampreys and hags. 
 
 Class II.— 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 ; Teleostomi. 
 
 With our present knowledge we find the first appearance 
 of these subclasses to be contemporary ; i. e., 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 Chimseras ; 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. kSome 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 vertebia'. 
 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 
 
 / 
 
 / 
 / 
 
 D 
 
 / / 
 
 B 
 
 D 
 
 D 
 
 rr 
 
 Ba 
 Ak 
 /Vs. 
 
 FxG.3. — Diagrams ot actinopliores; A, eutotetramerous; B^ ectetramerous; C, />. 
 ectrimerous; E^ ecdimeroas; Ac, actinotrichia; i^r, fin-rays; Ba, baseost; Ax. axu- 
 nost; Ic, 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 (haemal spines), the result being a two-lobed 
 fin, in which the upper lobe is much larger than the lower. 
 
20 
 
 COPE 
 
 This is the heterocercal tail (Fig. 4, B, G, and D). In later 
 forms the hsemal spines grow larger and the neural spine's 
 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 haemal 
 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 heterocercali 
 stages before reaching maturity. 
 
VEETEBEATA 
 
 21 
 
 Fig. 4, — Tails of fishes; A^ diphycercal tail of Polypterus; B, dlphyheterocercal tail of Cocco- 
 steus; C, caudal fin of a shark (Centrina); Z), end of vertebral column of trout {Salmo fario); 
 E, end of vertebral column of barbel (Barbus fluviatilis) ; ep, Epural; hy, hypural bones. 
 
 Subclass I.— ELASMOBRANCHII. 
 
 The sharks are divided into three orders, as follows: 
 Paired fins ptychopterygial ; with axonosts 
 
22 
 
 COPE 
 
 (metapterygiura) enclosed in body-wall 
 
 and basilars only free; Acanthodii. 
 
 Paired fins archipterygial ; unibasal ; a basi- 
 
 occipital element; Ichihyotomi. 
 
 Paired fins pluribasal (tribasal) ; no basioccip- 
 
 ital element ; Selachii. 
 
 The order Acanthodii presents the most primitive type 
 of paired 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 clas'pers. 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. 
 
 Fig. S.—Pleuracanthus lucius Ag. From the Coal Measures of Alsace. From 8auvage. 
 
 The Acanthodidse have but one dorsal fin, while the Isch- 
 nacanthidse and Diplacanthidse have two. In the Dipla- 
 canthidse only have clavicles been observed; in the other 
 
VEETEBEATA 
 
 23 
 
 two families they are wanting. Among Diplacanthidse 
 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. 
 
 Diplacanthidse are Devonic, as are also Ischnacanthidse. 
 Acanthodidge belong to the Devonic and Carbonic. 
 
 Fig. 6.— Pectoral and ventral fins of Cladoselache xi/g (from Dean). B, axonoats (meso- 
 and meta-pterygium) ; R, baseosts ; D, dermal portion of fin. 
 
 In the IcHTHYOTOMi 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 Pleuracanthidse 
 and the Cladodontidse, 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 Pleuracanthidse have teeth with two principal equal 
 cusps, while the Cladodontidse have a principal median cusp, 
 
24 COPE 
 
 with or without smaller lateral cusps. The Cladodontidse 
 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 Selachii (sharks and rays) present two lines of re- 
 lation, or suborders, which dififer as follows : 
 
 Vertebrae, when developed, having the con- 
 centric laminae predominating over the 
 radiating laminae ; anal fin absent ; Tedospondyli. 
 
 Vertebrae when developed, with the radiat- 
 ing laminae predominating over the con- 
 centric ; anal fin present ; Aster ospondyli. 
 
 In the Tedospondyli 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 Aslerospondyli, 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) Petalodontidas. 
 
 AA, The crowns of the teeth not overlapping. 
 
 /?. Summits of crowns forming a grinding sur- 
 face. 
 
 (2) Psammodontidse. 
 
 (3) Rajidse. 
 
 (4) Rhinobatidse. 
 
 (5) Trygonidx. 
 
 (6) Myliobatidse (eagle rays). 
 
 j9/3. Summits of crowns elongate cusps. 
 
VEETEBEATA 
 
 Body not flattened ; 
 
 25 
 
 (7) Spinacidse. 
 
 II. Snout produced into an ensiform weapon toothed 
 on both sides. 
 
 (8) Pristiophmidss. 
 (9) Pristidse (saw-fishes). 
 
 These families have the following range in geological 
 time : 
 
 
 1 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
 Plistocene 
 
 
 
 
 
 
 
 
 — 
 
 — 
 
 
 — 
 
 
 
 
 
 
 Neocene 
 
 
 
 
 
 
 
 
 
 
 
 
 
 Eocene . 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 Cretacic . 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 Jurassic . 
 
 
 
 
 
 
 
 
 
 
 Triassic , 
 
 
 
 
 
 
 
 
 
 
 C&,rbonic 
 
 
 
 
 
 
 
 
 
 
 
 
 Devonie 
 
 
 
 
 
 
 
 
 
 
 Siluric . 
 
 
 
 
 
 
 
 
 
 
 Ordovicic 
 
 
 
 
 
 
 
 
 
 
 Cambric 
 
 
 
 
 
 
 
 
 
 
 Huronio j 
 
 
 
 
 
 
 
 
 
26 
 
 (»PE 
 
 Fig. I.—Heptanchus griseus left pectoral fin, pluribasal type; SB, scapular arch; 
 JVL, foramen; Fr, propterygium; JKs, mesopterygium; a, 6, axis of metapterygium; 
 Sa, basilars; FS, aotinotricliia. From Wiedersheim. 
 
 The families of the Asterospondyli differ as follows : 
 
 I. Teeth molariform.* 
 
 Teeth separate ; (1) Cestraciontidae. 
 
 Teeth confluent ; (2) Cochliodontidse. 
 
 II. Teeth with elevated cusps. 
 
 (3) Hexanchidse (Fig. 7). 
 
 (4) Sq/lliidse. 
 
 (5) Lamnidse. 
 
 (6) Carchariidse. 
 
 The distribution of these families in time is as follows : 
 
 * Except ^ome Cestraciontidae. 
 
VEJRIEBKATA 
 
 27 
 
 ;!-4 
 
 Plistocene . . 
 
 
 1 
 
 
 f 
 1 
 
 
 
 Neocene . 
 
 ■ i 
 
 1 
 
 
 Eocene ... . 
 
 
 
 Cretacic . . 
 
 
 ' 
 
 
 " 
 
 Jurassic 
 
 
 
 i 
 
 ! 
 
 
 
 Triassic . . . . 
 
 
 =- — 
 
 
 
 
 
 Carbonic 
 
 
 
 
 ! 
 
 i 
 
 Devonic . . . . 
 
 
 
 
 
 Siluric 
 
 
 
 
 
 
 Ordovicic . . . 
 
 
 
 
 
 
 Cambric 
 
 
 
 
 
 
 Huronic 
 
 
 
 
 
 
 From these tables it appears that the Seiachii 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; Chimseroidei. 
 
 In all known CniMiEROiDEi 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. 
 
 Chimseroidei appear in the Corniferous Limestone in Ohio, 
 Wisconsin and Iowa, 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- 
 tidse (of doubtful reference), the Squalorajidse, Myriacan- 
 thidse, and Chimseridse, 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 fins archipterygial 
 (;'. e., unibasal); pel- 
 vic elements fu$ed 
 -?j- on the middle line; 
 body without bony 
 plates (Fig. 8); Sirenoidea. 
 
 The Arthrodira are represent- 
 ed by three or four families, the 
 Coccosteidse, the Asterosteidse, and 
 the Mylostomidse, all restricted to 
 Kr^ft; %7SZti'"iX u^Tbtai ^^^ P^riods of the Dcvouic sys- 
 llS!As'^>%i^tnotrt^iTkok tem. Their remains abound in 
 wiedersheim. ^j^^gg horizous of Europc 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 Dipteridse, Phaneropleuridse, and Ctenodontidge, 
 where the skull is covered with small tessellated plates, and 
 the Lepidosirenidse, where the bones of the skull are few 
 and large. The Dipteridse and Phaneropleuridse are con- 
 fined to the Devonic, and the Ctenodontidse to the Car- 
 bonic. The Lepidosirenidse 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 ; Bhipidopterygia. 
 
 II. Median fins with numerous axonosts. 
 All paired fins with baseosts; pectorals only 
 
 with a distinct peduncle formed of 
 
 axonosts ; Crossopierygia. 
 
 All paired fins with baseosts ; neither with 
 
 distinct peduncle of axonosts ; Podopterygia. 
 
 Axonosts not forming a peduncle ; pec- 
 torals only with developed baseosts ; Adinopterygia. 
 
 Superorder Rhipidoptekygia. 
 The orders of Ehipidopterygia 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; Adinistia. 
 
 The Taxistia includes but one family, the Holoptychiidse, 
 which is of Devonic age. 
 
 The Rhipidistia includes the Tristichopteridse, from the 
 Devonic and Carbonic ; the Osteolepidse, from the same ; 
 and possibly the Onychodontidse, which are Devonic. 
 
 The Actinistia includes the single family of the Coela- 
 canthidse, 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. 
 
 Ssfc/.^ 
 
 Fig 9—ZMdina penicillata Miinst, (Coelacanthidse) ; one-third natural size, showing rhipidopterygiar and 
 actinistious types of flns ; ^^ jugular plates ; 6, scales of Unidina acutidens. From Zittel. 
 
 Superorder Crossopterygia. 
 The superorder Crossopterygia 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 Tarasiidse, 
 from the Lower Carbonic of Scotland. The Cladistia are 
 represented by a family which is not known in the fossil 
 state, the Polypteridae of the rivers of Africa. The vertebrae 
 in this family are ossified and biconcave. 
 
 Superorder Podopteeygia. 
 The superorder Podopterygia has also two orders. They 
 are thus defined : 
 
 Branchiostegal rays present ; Lysopteri. 
 
 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 ; Palseoniscidse. 
 
 Teeth obtuse, on palate and splenial ; Platysomidse. 
 
 No teeth ; Chondrosteidse. 
 
 II. Tail diphycercal. 
 
 Teeth present ; scuta on body ; Belonorhynchidae. 
 
 The Palseoniscidse range from the Devonic to the Jurassic 
 inclusive ; the Platysomidse belong to the Carbonic ex- 
 clusively ; the Belonorhynchidae to the Trias ; and the 
 Chondrosteidse to the Jurassic. 
 
 The Chondrostei are degenerate representatives of the 
 
VERTEBRATA 
 
 3S 
 
 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 
 Acipenseridse, or sturgeons, and the Polyodontidse, 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 the scapula. 
 (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 
 contracted to the size of the 
 inferior lobe, and the verte- 
 brae ossified; but in some of 
 the lower and older types 
 both heterocercal tails and 
 notochordal vertebi'te 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 ; Malacoptcrygia. 
 
 Jlaf 
 
 Fig. va.—Salmo fario L ; left shoulder- 
 girdle; Cm, posttemporal ; D 1, epiclavi- 
 cle ; D, clavicle ; D 2, postclaricle ; S, 
 scapula : Co (CI), coracoid; Ba, basilars; 
 L, scapular foramen ; US, FS, flin-rays. 
 
34 COPE 
 
 Ventral fins usually thoracic or jugular ; 
 nO' ductus pneumaticus; usually a 
 spinous dorsal fin; parietal bones us- 
 ually separated by the supraoceipital ; 
 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 vertebrse unmodi- 
 fied. (Holostei). 
 
 a, Vertebrse complex, the pleurocentra 
 and intercentra distinct. 
 
 Anterior vertebrse similar to others ; (2) Merospondyli. 
 
 aa, Vertebrse with centra and intercentra complete 
 on part of the column at least ; amphicoelous ; 
 
 Anterior vertebrse similar ; (3) HalecomorpM. 
 
 aaa, Vertebrse (intercentra) opisthocoelous. 
 
 * The position of this order is uncertain, 
 
VERTEBEATA 35 
 
 Anterior vertebrae similar; a precoracoid 
 
 arch and a coronoid bone ; (4) Ginglymodi. 
 
 AA. No metapterygium ; anterior vertebrae modified, 
 and with ossicula ; vertebras amphicoelous. (Ostario- 
 physse). 
 
 ;9, A precoracoid arch. 
 
 Y, No symplectic bone. 
 
 Anterior vertebrte modified, and with ossi- 
 cula auditus ; pterotic simple ; parie- 
 tals not distinct ; (5) Nematognathi. 
 
 JX, A symplectic bone. 
 
 Anterior vertebrae coossified, and with ossi- 
 cula auditus ; pterotic simple ; parie- 
 tals distinct ; (6) Pledospondyli. 
 
 j3j^, No precoracoid arch. 
 
 Parietals present, distinct, not separated 
 
 by supraoccipital ; (7) Glanencheli. 
 
 AAA. No metapterygium ; vertebrae amphicoelous, 
 the anterior not modified and without accessory 
 ossicles. (Teleocephali). 
 
 /5, Scapular arch suspended to cranium. 
 
 «, A symplectic. 
 
 ;', A precoracoid arch. 
 
 Pterotic annular, including a cavity which 
 is closed by a distinct bone ; parietals 
 distinct, not separated by supraoccip- 
 ital ; (8) Scyphophori. 
 
 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) Adinochiri^ 
 
 YY, 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. 
 
 /?/3, Scapular arch free from cranium. 
 
 d, A symplectic bone. 
 
 Hyoid arches and pectoral baseosts de- 
 veloped ; (14) Holostomi. 
 
 00, 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 : 
 
 DocoPTEEi : Dorypteridse. 
 
 Mbeospondyli : Sauropsidse; Pycnodontidse ; Stylodontidaj ; 
 Sphserodontidse ; Macrosemiidse. 
 
 Halecomorphi : Amiidse (dog-fishes). 
 
 GiNGLYMODi : Lepidosteidse (bony gars). 
 
 Nematognathi : Siluridee (cat-fishes); Hypophthalmidte :, 
 Aspredinidse. 
 
VEETEBRATA 
 
 87 
 
 Plectospondyli : Characinidse ; Sternopygidaj ; Cobitidse ; 
 CyprinidEe (carp) ; Catostomidse (suckers). 
 
 Glanencheli: Gymnotidae (electric eels). 
 
 ScYPHOPHORi : Mormyridse ; Gymnarchidse. 
 
 IsospoNDYLi : Dapediidse ; Lepidotidse ; Aspidorhynchidse ; 
 Saurodontidse (Fig. 11) ; Osteoglossidse ; Heterotidse ; 
 Galaxiidse ; Clupeidte (herring) ; Chirocentridse ; Sal- 
 monidfe (salmon) ; Thymallidse (grayling) ; Alepoceph- 
 alidse; Gonorhyuchidfe ; Sauridse; Lutodiridse; Aulo- 
 pidse; Elopidte; Albulidee; Hyodontidse; Notopteridse. 
 
 AcTiNOCHiRi : Erisichthidse. 
 
 Haplomi : Esocidse (pike) ; Stratodontidse ; Umbridse ; Cy- 
 prinodontidae ; Amblyopsidse (blind fishes). 
 
 Xenomi : Dalliidse. 
 
 IcHTHYOCEPHALi : Monopteridse. 
 
 HoLOSTOMi : Amphipnoidse. 
 
 Enchelycephali : Nemichthyidse; Anguillidse (eels) ; Con- 
 gridse (eels) ; Synaphobranchidse ; Simenchelyidse. 
 
 CoLocEPHALi : Mursenidse (eels). 
 
 Lyomeri: Saccopharyngidse; Eurypharyngidse. 
 
 Fig. 11.— FortTi^us moiossits, 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: 
 
38 
 
 COPE 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CD 
 
 
 
 
 
 
 
 
 
 
 
 
 i-H 
 
 [ 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 1— 1 
 
 
 
 
 
 
 
 
 
 
 
 
 (M 
 
 
 
 
 
 
 
 
 
 
 
 
 i~t 
 
 [ 1 
 
 
 
 
 
 
 
 
 ! 1 
 
 O 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CO 
 
 1 1 
 
 
 
 
 
 
 
 
 ) i 
 
 
 00 
 
 
 
 1 
 
 
 
 
 
 
 
 
 t- 
 
 
 
 
 
 
 
 
 
 
 
 
 CD 
 
 1 
 
 
 
 
 
 
 
 
 
 lO 
 
 
 
 
 
 
 
 
 
 
 
 Tl< 
 
 
 
 
 
 
 
 
 
 
 
 
 
 M 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 <M 
 
 
 
 
 1 
 
 
 
 
 
 i 
 
 
 
 i-H 
 
 
 
 
 
 
 
 — 
 
 
 
 
 
 
 s 
 .s 
 
 a 
 
 <u 
 
 a 
 
 ■s 
 
 .2 
 
 1-5 
 
 .S 
 
 1 
 
 3 
 
 ■§ 
 
 > 
 Q 
 
 .2 
 
 ■ 
 
 i 
 
 o 
 
 'a 
 o 
 
 u 
 3 
 
 w 
 
VEETEBEATA 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) Lophobranchii. 
 
 aa, Ventral fins thoracic or jugular. 
 
 j3, Anterior (spinous) dorsal fin expanded 
 into transverse laminse, sessile on cranium. 
 
 Cranium normal; (5) Discocephali. 
 
 ^/3, Spinous dorsal fin not transversely ex- 
 panded. 
 
40 COPE 
 
 Y, Posttemporal projecting freely from 
 skull. 
 
 First vertebra united by suture to crani- 
 um ; intercalaria united behind su- 
 praoccipital ; basilar pectoral bones 
 elongated; (6) Pedieulati. 
 
 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 
 coossified with maxillaries behind, 
 and the dentary with the articular; 
 pharyngeal bones distinct ; (8) Plectognathi. 
 
 Cranium normal; bones of jaw dis- 
 tinct; (9) Percomorphi. 
 
 YT, 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: Mastacembelidse ; Notacanthidee. 
 
 Pbecesoces: Opheocephalidse; Mugilidse (mullets); Ather- 
 inidse; SphyrEenidse (barracuda); Scombresocidse (soft 
 gar). 
 
 Hemibranchii : Pegasidse; Gasterosteidse (stickle-back); 
 Fistulariidse ; Centriscidse ; Amphisilidse ; Dercetidse. 
 
 LoPHOBRANCHii : Solenostomidse; Syngnathidse (pipe- 
 fishes); Hippocampidee (sea-horses). 
 
 DiscocEPHALi : Echeneididse (remoras). 
 
 Pediculati : Antennariidge ; Lophiidse (fishing frog). 
 
VERTEBEATA 41 
 
 Heterosomata : Pleuroiiectidse (flat-fishes). 
 
 Plectognathi : TriacanthidEe ; Balistidfe (trigger- fishes) ; 
 Tetraodontidse (bladder-fishes) ; Diodontidge ; Ostra- 
 ciidfe. 
 
 Pbrcomorphi: (Anacanthini) Ophidiidse ; Gad idee (cod); 
 Macruridse ; (Haplodoci) Batrachidse (toad-fishes) ; 
 (Cyclopteroidea) Cyclopteridse ; (Scatophagoidea) Sca- 
 tophagidse ; (Epilasmia) Acronuridse ; Chsetodontidse ; 
 (Rhegnopteri) Trichidiontidse ; (Distegi) Scorpsenidge ; 
 Cottidfe (sculpins) ; Blenniidse ; Gobiidse ; Platycepha- 
 lidse; Rhamphocottidse ; Agonidse; Heterognathidse ; 
 Gerreidse ; Carangidee' (pompano); Sillaginidse ; Pristi- 
 pomatidse ; Sciaenidse (maigres) ; Sparidse ; Percidse 
 (perch) ; Berycidse ; ScombridEe (mackerel); Trichiuridse ; 
 Xiphiadidse; (Labyrinthici) Osphromenidse ; Anaban- 
 tidse; (Pharyngognathi) EmbiotocidEe ; Cichlidse; 
 Labridae; Scaridse. 
 
 Craniomi; Triglidse (gurnards); Dactylopteridse. 
 
 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 : 
 
 Note.— The editor is indebted to Prof. Tlieodore 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. 
 
42 
 
 COPE 
 
 o 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CT> 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 00 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 r- 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 so 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 U3 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 ^ 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CO 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 IM 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 r-t 
 
 
 
 
 
 
 
 
 
 
 
 
 
 a) 
 
 s 
 
 1 
 
 g 
 
 O 
 
 'S 
 
 OS 
 
 1 
 
 1-5 
 
 
 'n 
 o 
 
 o 
 
 .2 
 
 'n 
 o 
 
 > 
 
 ft 
 
 
 6 
 
 1 
 
VERTEBEATA 43 
 
 Class III.— 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 vertebrfe united into 
 a urostyle; Saliodia. 
 
 Subclass I.— STEGOCEPHALI. 
 
 There are four orders of Stegocephali. 
 
 «, One occipital cotyloid articulation. 
 
 Vertebral bodies represented by basal and 
 
 lateral elements (intercentra and centra) ; Ganocephali. 
 
 aa, Two occipital condyles; 
 
 Vertebras 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 
 
 t;oPK 
 
 Fig. 12.— Ti-i7nerorhachis insignis Cope ; parts of skeleton, natural size ; a, occipital segment with 
 single cotylus ; 6, Cy posterior part of lower jaw ; d, e, series of vertebrae ; d, side view, depressed ; 
 e, view obliquely from above, both showing the rhachitomous structure : i, 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, i). In the 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. 
 
 
 I 
 
 Fig. 13 — Trimerorhachis insignia Cope; skull. From the Permian bed of Texas. 
 Original. 
 
 Of the Ganocephali two families are known, the Tri- 
 merorhachidse (Figs. 12-13), without, and the Archego- 
 sauridse with neural spines of the vertebrae. They occur in 
 the Coal Measures and the Permian of Europe and North 
 America. 
 
46 
 
 COPE 
 
 The Rachitomi possesses but one family, the Eryopidse, 
 which often reached a large size (Fig. 14). These are con- 
 fined to the Coal Measures. If the Labyrinthodontidse be- 
 long to this order they range also to the Trias inclusive, in 
 both continents. 
 
 The Embolomeri includes one family, the Cricotidse, 
 
 N 
 
 \ 
 
 Fig. U.— 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). 
 
 
 Fig. 15, — Cricotus heteroclitus Cope ; one-third natural size. From Permian of Texas, 
 a, head from aove ; &, part of belly f rfim below. From Cope. 
 
 The MiCROSAURi embraces the following families : Bran- 
 chiosauridse ; Hylonomidae ; MolgophidfB ; Phlegethontiidae. 
 The smaller forms are mostly from the Coal Measures, 
 while some large ones occur in the Trias. Some of the 
 Hylonomidae come from the Permian of North America and 
 Europe. They were mostly of small size, and their verte- 
 brae exhibit very various degrees of ossification. In some 
 the bodies are completely ossified, while in others the ossi- 
 
48 COFE 
 
 fication forms a superficial layer. In the Labyrintliodon- 
 tidse tiae ossification is more complete. The dentine may 
 be entire, or deeply inflected, so as to form straight or 
 labyrinthic folds. 
 
 Subclass II.— UEODELA. 
 
 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 tlie Proteida the only family known is the Pro- 
 teida (Fig. 16.) 
 
 The Pseudosauria embraces the following families : 
 Cryptobranchidge; Amblystomidse ; Hynobiidse; Plethodon- 
 tidse; Thoriidse; Desmognathidse; Salamandridse ; Pleuro- 
 delidse; Amphiumidse; Cseciliidise. 
 
 The Trachystomata includes only the family of the Si- 
 renidse. 
 
 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. 
 
M 
 
 m c^ "" 
 2 ®"^ 
 
 O JS 3 
 
 a 
 
 3N 
 
 O O d 
 
 S S. 3 
 S i3 g 
 
 «•■■'". 
 a-" 
 
 ■a ►< M 
 
 S<0 ,- 
 
 ■a« 
 
 
 
 ^Sb- 
 
 ag.; 
 
 (O O 0^ 
 
 go? CO 
 
 M S~i CO 
 
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 ordgrs are as follows : 
 
 (Aglossa) : Xenopidse ; Pipidse. 
 
 (Arcifera) : Discoglossidse ; Bufonidse ; Dendrophryniscidse ; 
 Asterophrydidse ; Pelodytidse ; Scaphiopidse ; Hylidse ; 
 Cystignathidse; Amphignathodontidse; Hemiphractidse. 
 
 (Firmisternia) : Engystomidse ; Phryniscidse ; Dendroba- 
 tidse ; Cophylidse; Dyscophidse; Colostethidse; Ran- 
 idse ; Ceratobatrachidse. 
 
 (Gastrechmia) : Hemisidse. 
 
 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. 
 
VERTEBRATA 
 
 51 
 
 a 
 
 
 
 
 
 
 
 i 
 
 
 
 
 
 
 
 
 
 
 
 s 
 
 t 
 
 p 
 
 s 
 
 1 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 'S 
 o 
 
 
 
 
 1 
 
 
 
 ] 
 
 
 
 
 
 
 3 
 
 I 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 p. 
 
 s 
 
 c 
 
 g 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CD 
 
 a 
 8 
 S 
 
 
 □ 
 o 
 
 'a 
 
 3 
 i-5 
 
 •fl 
 
 'S 
 
 'a 
 o 
 > 
 
 ft 
 
 _3 
 
 o 
 '3 
 
 1 
 
 o 
 
 i 
 
 "3 
 
 o 
 s 
 
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 coossified 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 Saururse approach reptiles in the structure 
 of the manus and of the caudal vertebrae. 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 vertebrae 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 tiie 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,. 
 
VEETEBRATA 63 
 
 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 corK 
 
 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 vertebrae of Reptilia are either biconcave (amphi- 
 coelous), 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 procoelous, which is 
 the most frequent ; or second with ball in front and socket 
 behind, or opisthocoelous. 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 Eeptilia 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, and a 
 presternum; limbs ambulatory; ver- 
 tebrse amphicoelous ; (1) Cotylosauria. 
 
 AA. Scapular arch internal to ribs ; temporal region 
 with complex roof and no longitudinal bars. 
 
 A presternum ; limbs ambulatory ; (2) Ghelydosauria. 
 
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 ; (3) 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 ; vertebrse 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- 
 tebrae amphicoelous ; (5) Theromora. 
 
 * No supramastoid ; paroccipital not dis- 
 tinct; a quadra to-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 ; 
 vertebrse amphicoelous ; (7) Pelycosauria. 
 
 aa, No supramastoid. 
 
 *Cope, Proc. Amer. Philos. Soc, 189J, identifies and describes a supramastoid in 
 tlie plesiosaurian skull. — Ed. 
 
56 COPE 
 
 Eibs 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) Rhynchocephalia. 
 
 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 Reptllia except the Chelydosauria. The bones are indicated by 
 a uniform system of abbreviations. — Ed. 
 
VERTEBEATA 
 
 57 
 
 s 
 
 e 
 
 ■Vj 
 
 8 
 
 
58 
 
 COPE 
 
VERTEBKATA 
 
 59 
 
60 
 
 COPE 
 
 The time-histories of the orders of reptiles are represented 
 in the accompanying table : 
 
 
 1 
 
 2 
 
 3 
 
 4 
 
 
 5 
 
 6 
 
 7 
 
 8 
 
 9 
 
 10 
 
 11 
 
 12 
 
 Plistoeene 
 
 
 
 
 
 --= 
 
 
 
 
 
 
 
 
 — 
 
 = 
 
 — 
 
 
 
 
 Neocene 
 
 
 
 
 
 
 
 
 
 
 
 
 Eocene . 
 
 
 
 
 
 
 
 
 Cretacic 
 
 
 — 
 
 — 
 
 
 ^- 
 
 
 
 — 
 
 — 
 
 — 
 
 — 
 
 
 Jurassic 
 
 
 
 
 
 ? 
 
 Triassic . 
 
 
 ! 
 
 
 
 = 
 
 
 
 
 
 
 
 Carbonic 
 
 
 1 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 Devonic 
 
 
 
 
 
 
 
 
 
 
 
 Siluric 
 
 
 — 
 
 
 
 
 
 
 
 
 
 
 
 Ordovicic 
 
 
 
 
 
 
 
 
 
 
 
 Cambric 
 
 
 
 
 
 
 
 
 
 
 
 
 Huronic 
 
 ' i 
 
 
 
 
 
 
 
 
 
 
 
 The order Ootylosauria includes reptiles with amphicoel- 
 ous vertebrse and of terrestrial habits. There are four families, 
 the Pariasauridee, 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 Diadectidse (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 PariotichidtB, with several 
 rows of, teeth in the jaws, from North America; and the Elgi- 
 niidse with very thin ossifications and light structure, from 
 the Trias of Scotland. This order is prohably ancestral to all 
 
 4^ ^tt< - -^ ' 
 
 V»..J ' 
 
 Fig. 17. — Empedlas motaris Cope ; inferior view of sliull, one-third natural size. 
 A Cotylosaurian from the I^ermian 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- 
 ilophum 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- 
 coelidge, 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 Testudinata presents four subordinal modifi- 
 cations, as follows : 
 
 I. No descending processes of the parietal bones. 
 Vertebrse and ribs free and separated from a 
 
 bony exoskeleton ; no descending pro- 
 cesses of the parietals ; Athecse. 
 
 II. A carapace and plastron, and descending process of 
 parietals. 
 
 «, 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 bones wanting or rudimental ; Trionychoidea. 
 
 aa, Sacral and caudal ribs articulating with body 
 of vertebrse only. 
 
 As the last ; but marginal bones present and 
 connected with ribs, and last cervical and 
 last dorsal vertebraj articulating by 
 bodies; pelvisnotanchylosed to plastron; Cryptodira. 
 
 Neck bending in horizontal plane, the last 
 cervical and first dorsal vertebrse articu- 
 
VEKTEBRATA 
 
 63 
 
 lilting by bodies; pelvis anchylosed to 
 carapace and plastron ; marginal bones 
 present and connected with ribs; Pleui-odira. 
 
 The Athccpe includes the single family of the Dermo- 
 chelyda\ 
 
 The Trionychoidea includes only the Trionychidfc. 
 
 The Cryptodira embraces the Cheloniidte, Testudinidai 
 (Fig. 18), Cinosternidse, Dermatemydidas, Clielydridfe, 
 Baenidfe, and Adocidce. 
 
 The Plcurodira includes the the Pleurosternidse, Steruo- 
 theridfe; Pelomedusidee, Plesiochelydidee, Chelydidfe, and 
 Carettochelydidse. 
 
 The earlier tortoises are intermediate in character be- 
 
 Fig. IS.—Testudo laUcuneus Cope ; carapace, from above, one-fourth natural size. 
 From tlie White River Neocene bed of Nebraska. 
 
64 COPE 
 
 tweeii 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 
 Athecse first appear in the Eocene, and were then and still 
 are marine in their habits ; but one species now exists, the 
 " leather-back turtle." 
 
 Fiti. 19. — lehUiyosavrus tenuirostis Conyb. ; one. sixth natural size. From the Lias 
 of Wiirtembiirg. From Doederlein. 
 
 The order Ichthyopterygia embraces the families of 
 Ichthyosauridse (Fig. 19) and Mixosauridse. 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 Ichthyosauridse 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 (Mursenosaurus) and others (Baptanodon) no teeth. 
 The typical genera had numerous teeth grooved at the 
 base. The vertebrte are short and biconcave. 
 
VERTEBEATA 65 
 
 The order Theromora includes five saborders, 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 ; Thcriodonta. 
 
 Teeth absent or reduced to one pair above ; Anoinodonfa. 
 
 aa, Occipital condyles two. 
 Teeth compressed ; Ci/nodonta. 
 
 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 theTheriodonta, the canine teeth 
 are greatly enlarged. The families are as follows : 
 Placodonta : Placodontidse, Trias of Europe. 
 Theriodonta: PProterosauridee, Permian of Europe; Rhyn- 
 
 chosauridse, Trias of Europe and India ; Lycosauridas, 
 
 Permo-Trias of South Africa and Europe. 
 Axomodonta; Dicynodontidse (Fig. 20), Permo-Trias of 
 
 South Africa, Asia and Europe. 
 Cynodonta : Galesauridse, Permo-Trias of South Africa. 
 Gomphodonta : Tritylodontidse, 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. 
 
 Fig. 20. — Lystrosaurus frontosus Cope ; an Anomodont from South Africa, one- 
 third natural size. 
 
 The Plesiosaueia embraces the following families : 
 Plesiosauridse, Nothosauridse, and Lariosauridse. The last 
 two families are only known from the Triassic of Europe. 
 The Plesiosauridse range from the Jurassic to the Cretacic, 
 
VEETEBRATA 
 
 67 
 
 inclusive, and include man}' 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 Pelycosauria includes the 
 families of the Bolosauridse and 
 Clepsydropidse, which were carni- 
 vorous and abundantly provided 
 with teetJi. They have been found 
 in the Permian of South Africa, 
 North America and Europe. 
 
 Two families enter the Orni- 
 THOSAUKiA, viz., the Pteranodon- 
 tidse and the Pterodactylidfe. 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 Pterodactylidse 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 
 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. 
 
 The Ornithosauri were the fly- 
 ing order of reptiles, comparing 
 with the other orders much as 
 bats compare with other orders 
 of Mammalia. 
 
 Fig. iX.—Dimorphodamacronyx 
 Ow. from the English Trias. B, 
 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 
 Cetiosauridse and the Megalosauridse, the former supposed 
 to be herbivorous, the latter carnivorous. The former have 
 robust inferior pelvic bones, and teeth vi^ith spoon-shaped 
 crowns. The latter have long, slender pubis and ischium, 
 and sharp, knife-shaped teeth. 
 
 To the Cetiosauridse belong the largest known Vertebrata 
 with ambulatory legs ; the Camarasaurus supremus Cope 
 measuring seventy feet in length, and the Amphicoelias 
 fragilissimus Cope being considerably larger. These crea- 
 tures had the dorsal vertebrse hollow, and probably pene- 
 trated in life by branches from the lungs, while the tail 
 vertebrse 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 Megalosauridse 
 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 
 vertebrse were mere hollow shells. In most of them the 
 limb bones were hollow. 
 
Fig. 22.^Cetiosauridae ; 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, Amphiceelias, 
 allies Cope ; femur, from behind, one-tenth natural size. 
 
70 COPE 
 
 The Orthopoda includes the Scelidosauridse, Hypsirho- 
 phidse, Agathaumidse, Camptosauridse, Iguanodontidse and 
 Hadrosauridse, all supposed to have been of herbivorous 
 habits. Representatives of these families have been found 
 in both Europe and North America, excepting the Scelido- 
 sauridse, 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- 
 thaumidee had a corresponding toothless bone in front 
 formed of the premaxillary. In all the families ex- 
 cept the Agathaumidse the successional teeth appear on 
 the inner side of the base of the functional teeth, as in 
 lizards. In the Agathaumidse they appeared under the 
 middle of the base, as in crocodiles. In all except Hadro- 
 sauridse one row was in functional use at a time ; but in the 
 Hadrosauridse 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 
 Agathaumidse mostly had formidable horns on the head, at 
 the middle of the nose and over the eyes. Agaihaumas 
 sylvesfre Cope reached a length of forty feet, and had the 
 legs of sub-equal length. Many of the Hadrosauridse 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. 
 
VEETEBEATA 
 
 
 f5 
 
 3 f>' 
 
 ? ^i 
 
 -<i^.^ 
 
 Fig. 23. — Diclon ius mirabilis heidy -, skull one-seventh natural size. F'rom Cope. A, from side ■ 
 J5, from below. From the Laramie beds of Dakota. ' ' 
 
72 
 
 COPE 
 
 I I 
 
 11 
 
 m 
 
 -«. 
 
 Fig. 24,— Belodon buceras Cope ; skull, one-fourtli natural' size. From the Trias o£ the 
 Kooky Mountains. ^, the side ; -B, helow; C, above. Original. 
 
VERTEBEATA 73 
 
 The LoEiCATA 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 Belodontidfe (Fig. 24). The Pseudo- 
 suchia include the Aetosauridae, 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 Teleosauridse and Goniopholididse, 
 which are followed in the Cretacic by the Crocodilidse (Fig. 
 25) and Gavialidse, which still exist. 
 
 The Rhynchocephalia include two suborders : 
 Vertebrae amphicoelous ; axis undivided ; Sphenodontina. 
 Vertebrae amphiplatyan ; dentatum of axis 
 
 separate ; Choristodera. 
 
 To the Sphenodontina belong the Protorosauridae and 
 Paleohatteriidae, from the European Trias, the Homoeosauri- 
 dae, from the European Jurassic, and the Sphenodontidae, 
 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 Champsosauridae, 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.—CrocodUiis acer Cope ; skull, one-third natural size. From the Eocene 
 of Utah. Original. 
 
 The Squamata 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. 
 
VERTEBEATA 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 Lacertilia 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 Nydisaura. 
 
 aa, Clavicle proximally simple ; olfactory lobes 
 underarched by frontal. 
 
 Vertebrae amphicoelous ; no supratem- 
 
 poral arch ; tongue papillose ; 5 Urolatoidea. 
 
 Vertebrae procoelous; a supratemporal arch; 
 interclavicle anchor-shaped ; tongue 
 smooth ; hemipenis flounced ; 6 Thecaglossa. 
 
 Vertebrae procoelous, 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. 
 
 Vertebrae procoelous ; 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 coossified with the middle 
 of the centra. 
 
VEETEBRATA 
 
 77 
 
 Palatine and pterygoid foramina ; an epi- 
 
 pterygoid ; tongue papillose ; 10 Amiielloidea. 
 
 No palative nor pterygoid formina; no 
 
 epipterygoid ; 11 Annulati. 
 
 A group whichi 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 vertebrae, which is more than nine, accord- 
 ing to Owen. Teeth pleurodont. 
 
 The families of these superfamilies are the following : 
 
 Dolichosauria ; Dolichosauriidee. 
 
 Rhiptoglossa ; Chamseleonidse. 
 
 Pachyglossa ; Agamidse, Iguanidae (Fig. 26.) 
 
 Nydsauria ; Eublepharidse, Geckonidse. 
 
 Uroplatoidea ; Uroplatidse. 
 
 Thecaglossa ; Varan idse. 
 
 Helodei^matoidea ; Helodermidse. 
 
 Diploglossa ; Zonuridse, Pygopodidse, Anguidse, Xenosauri- 
 
 dse. 
 Leptoglossa ; Xantusiidse, Teiidse, Lacertidse, Gerrhosauridae, 
 
 Scincidae, Acontiidae, Anelytropidae. 
 Annielloidea ; Aniellidse. 
 Annulati ; Chirotidse, Amphisbaenidse, Trogonophidae. 
 
 I'r 
 
 Tinx,fi 
 
 Fig. 26.— Skull of Iguana tubereulata Linn ; external side view, natural size. 
 
 The recent forms of the Lacertilia are much more numer 
 ous than the known fossil forms. 
 
78 
 
 COPE 
 
 The geologic history of the Lacertilia is as follows : 
 Column No. 1 represents the Dolichosauria. 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 o 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 o 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CO 
 
 
 
 
 
 
 
 
 
 
 
 
 
 b- 
 
 
 
 
 
 
 
 
 
 
 
 
 
 CD 
 
 
 
 
 
 
 
 
 
 
 
 
 
 lO 
 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 Tf" 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 W 
 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 i 
 
 o 
 
 § 
 
 o 
 
 i 
 
 1f1 
 
 .2 
 
 _o 
 
 'S 
 
 o 
 
 o 
 
 '3 
 
 o 
 > 
 
 
 o 
 n3 
 
 
 
 
 S 
 
 ^ 
 
 w 
 
 
 ^ 
 
 ^ 
 
 
 ft 
 
 JE 
 
 O 
 
 6 
 
VEETEBEATA 
 
 79 
 
 t^ 
 
 h 
 
 < 
 
 4 
 
 >. 
 
 P 
 
 
 ? I 
 
 ^ 
 
 Fig. 27. — Olidastes vropy- 
 ttoTiCope; skull one-half natur- 
 al size. From the Upper Greta 
 cic of Alabama. 
 
 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 Pythonomoepha (Fig. 27) in- 
 clude two families, the Mosasauridse 
 and the Plioplatecarpidse. The 
 Mosasauridse were the predominant 
 type of sea-saurians during the Cre- 
 tacic period in North America, 
 and they were common in Europe 
 and in New Zealand, and some 
 species have been found in Brazil. 
 The Plioplatecarpidse 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 Ophidia include the follow- 
 ing superfamilies : 
 
 A. Supratemporal intercalated 
 in the cranial walls. {Angio- 
 stomata.) 
 
 a, No ectopterygoid ; pala- 
 tines bounding choanse 
 posteriorly ; ethmotur- 
 binal forming part of 
 roof of mouth ; rudi- 
 ments of a pelvis. {Sco- 
 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 
 choanse posteriorly. 
 Maxillary bone free, horizontal ; (3) 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. 
 
 innx 
 
 Car 
 
 Fie. 28. — Xenopeltis unicolor Reinwt, from Siam; skull, natural size. Original. 
 (Asinea.) 
 
VERTEBEATA 
 
 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 ; Stenostomidse. 
 
 Epanodonta ; Typhlopidse. 
 
 Tortricina ; Tortricidse, Uropeltidfe. 
 
 Colubroidea ; (Peropoda) Pythonidse, Boidse (Fig. 29), Char- 
 
 inidse, Ungualiidse; (Aglyphodonta), Xenopeltidse (Fig. 
 
 28), Acrochordidse, Nothopidse, Colubridse ; (Opistho- 
 
 glypha) Dipsadidse ; (Proteroglypha), Elopida;, Naji- 
 
 dse, Dendraspididse, Hydrophidse. 
 Solenoglypha; Causidse, Atractaspididse, Viperidffi, Crotali- 
 
 dae. 
 
 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 Pythonidse and Boidse ; they occur in both continents. 
 
 Fie. 29.— A, Palceovhis UUoralis Cope: B, P. halidanus Cope; vertebrm 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 : 
 
 
 1 
 
 2 
 
 3 
 
 4 
 
 5 
 
 Plistocene 
 
 
 
 
 
 
 
 
 
 
 Neocene 
 
 
 
 
 
 
 
 
 Eocene 
 
 
 
 
 
 
 
 Cretacic 
 
 
 
 
 
 
 
 Jurassic 
 
 
 
 
 
 
 Triassic 
 
 
 
 
 
 
 Carbonic 
 
 
 
 
 
 
 
 Devonic 
 
 
 
 
 
 Siluric 
 
 
 
 
 
 
 Ordovicic 
 
 
 
 
 
 
 
 Cambric 
 
 
 
 
 
 Huronic 
 
 
 
 
 
 
VEETEBEATA 
 
 83 
 
 Subclass II.— AVES. 
 There are two superorders of the birds as follows : 
 
 Metacarpal and carpal bones all distinct, the 
 digits with ungues ; caudal vertebrse 
 numerous, unmodified ; clavicles united ; 
 pelvic elements distinct ; teeth present ; Saururse. 
 
 
 Via.ZO.—Archceopteryx lUhographica Wagn. From Middle Oolite of Bavaria 
 much reduced. From Dames. 
 
84 COPE 
 
 Metacarpal and carpal bones reduced in num- 
 ber, coossified ; ungues wanting or 
 single ; caudal vertebrae reduced in num- 
 ber, the terminal areas usually coossified ; Eurhipidurm ■ 
 
 The Saueur^ includes but one order, which is defined' 
 
 as follows : 
 
 Vertebrae biconcave ; feathers arranged in 
 one series on each side of the caudal ver- 
 tebrse ; teeth present ; Ornithopappi.. 
 
 To this order but one family belongs, viz., the Archse- 
 opterygidse. 
 
 This family is represented by one genus, Archseopteryx 
 from the Oolite (Middle Jurassic) of Solenhofen, Bavaria. 
 It 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 EuRHiPiDUE^E include four or five tribes which dif- 
 fer as follows : 
 Ischium free from ilium posteriorly ; * 
 
 palate dromseognathous {i. e., maxillo- 
 
 palatines articulating with vomer 
 
 which is between them, palatines not 
 
 articulating directly with sphenoid 
 
 rostrum); no teeth ; (1) Ratitse. 
 
 Ischium free from ilium posteriorly ; pal- 
 ate ? dromsegnathous ; teeth ; (2) Odontolcx. 
 
 Ischium free from ilium posteriorly ; pal- 
 ate not dromseognathous (?) ; teeth ; (3) Odontotormie. 
 
 Ischium coossified with ilium posteriorly ; 
 palate not dromseognathous ; feathers 
 
 *Witli some apparent exceptions. 
 
VEETEBRATA 
 
 85 
 
 distributed in areas, those of the wings 
 much differentiated ; (4) Euornithes. 
 
 Ischium coossified with ilium posteriorly ; 
 no teeth; feathers universally distrib- 
 uted and not differentiated on wings; (5) Impennes. 
 
 The Ratitse are the ostriches and their allies ; the Odon- 
 tolcEE and Odontotormse 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 : 
 
 
 1 
 
 2 
 
 3 
 
 i 
 
 5 
 
 Plistocene . . . 
 
 
 
 
 
 
 
 
 
 Neocene . . . 
 
 
 
 
 
 
 
 
 
 Eocene . . . 
 
 
 
 
 
 
 Cretacic .... 
 
 
 
 
 
 
 
 
 Jurassic . . . 
 
 
 
 
 
 
 Triassic . . . 
 
 
 
 
 
 
 Paleozoic .... 
 
 
 
 
 
 
 The RatiTvE includes the following orders : 
 
 Sternum without keel ; clavicles ; wings rudi- 
 
 mental ; Struthiones.. 
 
 Sternum without keel ; no clavicles ; wings 
 
 rudimental ; Apteryges.. 
 
 Sternum with keel ; clavicles ; wings rudi- 
 
 meutal ; , Gastornithes.. 
 
 Sternum with keel ; clavicles ; wings func- 
 tional ; Crypturi.. 
 
86 
 
 COPE 
 
 The families belonging to these orders are the following : 
 Struthiones ; Struthionidee (ostriches), Rheidee, Casuariidas, 
 
 DromaBidse, Dinornithidse (Fig. 31), iEpiornithidae. 
 Apteryges ; Apterygidae (kiwis). 
 Gastornithes ; Gastornithidfe. 
 Crypturi ; Crypturidse (tinamus). 
 
 Of the above families the Struthionidiu are remarkable 
 in having the pubes forming a ventral symphysis. The 
 Rheidfe are equally remarkable in having a ventral sym- 
 physis of the ischia. The geological range of these orders 
 is as follows : 
 
 Plistocene 
 
 Neocene 
 
 Eocene 
 
 Cretacic 
 
 Jurassic . 
 
 Triassic . . 
 
 Carbonic 
 
 Devonic 
 
 Siluric 
 
 Ordovicic 
 
 Cambric . 
 
 Huronic 
 
 Struthiones 
 
 Apteryges 
 
 
 
 1 
 
 _ 1 
 
 
 
 
 
 
 
 
 
 
 -= 
 
 
 
 Gastornithes 
 
 Crypturi 
 
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. 
 edwardsil 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. 
 
 Fig. Z\.—Dinornu parvus Owen. From Plistocene of Xew Zealand. From 
 Owen. 11, ilium; is, ischium; pp, pubis; st, sternum; B, tarsonietatarsus. 
 
 The tribe Odoxtolc/E includes but one order : 
 
 Teeth in a groove; sternum without keel; 
 wings rudimeutal ; pelvic bones free 
 posteriorly ; vertebr£E saddle-shaped ; Drommopappi. 
 
 The Drom/EOPAPPi has but one family, the Hesperorni- 
 thidse, 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 Cecomorphfe). 
 
 To the Odontotorm^ one order only is referred. It is 
 thus characterized : 
 
 Teeth in sockets ; sternum keeled ; vi^ings 
 well developed ; ischium and pubis free 
 posteriorly ; vertebrte biconcave ; Pteropappi. 
 
 The family of the Ichthyornithidse is the only one 
 known to belong to the Pteropappi. Its members are 
 known from the Upper Cretacic of North America and the 
 Lower Cretacic of England. 
 
 The EuoENiTHES include numerous suborders, wliich are 
 defined as follows : 
 
 1. Maxillopalatines united across the middle of the 
 palate. (Desmognathse). 
 
 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 ; prsecocial ; (3) Chenomorphas. 
 
 No basipterygoid processes ; bill and legs 
 slender ; toes generally free ; altri- 
 cial ; (4) Herodii. 
 
 Bill and claws hooked ; toes free ; ver- 
 tebrae saddle-shaped ; altricial ; (5) Accipitres. 
 
 Bill hooked ; toes free ; vertebrae opis- 
 thocoelous ; rostrum movably ar- 
 
 *Except Caeulidse and Rhamphastidse, which are zygodactyloiis. 
 
VEETEBRATA 89 
 
 ticulated with skull ; basiptery- 
 
 goids ; (6) HctcroxpondyM. 
 
 Toes free ; vertebrte saddle-shaped ; ros- 
 trum fixed ; (7) Coccygomorphie. 
 
 AAA. First and fourth toes directed backwards 
 (zygodactylous). 
 
 Rostrum freely articulated with the 
 
 skull ; vertebrae opisthocoelous ; (8) Psittaci. 
 
 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- 
 g)tathfe.) 
 
 A. Toes three forwards (anisodactylous). 
 Schizorhinal ; toes webbed; (10) Cccomorplnr. 
 
 Toes free ; legs long; feathers with after- 
 shaft : prsecoces ; (11) Grallse. 
 
 No basipterygoids ; lachrymal bones co- 
 ossified with rostrum ; toes free ; (12) Opisthocomi. 
 
 Toes free ; hallux rudimental ; (13) Gallhix. 
 
 Toes free ; hallux well developed ; two 
 
 carotid arteries ; (14) PuUastrx. 
 
 Toes free ; hallux well developed ; one 
 
 carotid artery ; basipterygoids ; (15) Micropodioulci* 
 
 AA. First and fourth toes directed backward 
 (zygodactylous). 
 
 * Family Trochilicl:i'. 
 
90 COPE 
 
 No caeca coli ; no interclavicle ; one 
 
 carotid artery; (16) Picoidei. 
 
 III. Maxillopalatines not united on median line ; 
 vomer single, truncate, and excavated in front. 
 {JiJgithognathae) 
 
 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 caeca coli ; one carotid 
 artery ; (17) Passeres. 
 
 AA. Four toes directed forwards (pampro- 
 dactylous). 
 Toes free ; no basipterygoids ; sternum 
 entire posteriorly ; tensor patigii 
 brevis muscle attached to a tendon 
 which extends to the manus ; no 
 csecacoli ; (15) Micropodioidei* 
 
 The cormorants, pelicans, and boobies of the Stegano- 
 podes appear in the Eocene. The Eocene representatives 
 of the Chenomorphee are primitive flamingoes, the true 
 ducks and geese not appearing before the Neocene. The 
 Eocene Accipitres are Falconidfe, the owls not appearing 
 before the Neocene. The kingfishers are the earliest repre- 
 sentatives of the Coccygomorphse, appearing in the Eocene; 
 the remaining families are not known prior to the Neocene. 
 The Phasianidse of the Gallinse 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 Cypselid*. 
 
VERTEBRATA 
 
 91 
 
 
 o 
 
 to" 
 
 3 
 
 1 
 Ordovieir 
 
 
 o 
 
 Q 
 
 3- 
 
 o 
 o' 
 
 
 c 
 
 3 
 
 o' 
 
 p 
 
 o 
 o 
 
 B 
 
 r 
 
 o 
 
 2 
 
 en' 
 O 
 
 
 
 
 
 
 
 
 
 
 
 
 1 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 to 
 
 
 
 
 
 
 
 
 
 
 
 
 CO 
 
 
 
 
 
 
 i 
 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 C7I 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 a> 
 
 
 
 
 
 
 
 
 
 
 - 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 i 
 
 „ 
 
 1 
 
 
 1 
 
 t 
 
 1 
 i 
 
 1 
 
 i 
 
 
 
 (O 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 o 
 
 
 
 
 
 
 ! 
 
 1 
 
 1 
 
 
 1 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 K> 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 
 : 
 
 
 
 
 
 
 
 
 
 
 
 >(^ 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 i 
 
 o, 
 
 
 1 
 
 
 
 
 
 
 
 [ 
 
 
 1 
 
 OJ 
 
 
 
 
 
 
 
 
 
 
 _ _.. 
 
 
 
 -J 
 
!)2 
 
 (»PE 
 
 D 
 
 Fig. 32.— Diagrams exhibitins the more important modes of distribution of the deep plantar 
 tendons in Euornithes. A^ nomopelmous (schizopelmous) ; B, synpelmous (desmopelmous) ; 
 C, antipelmous ; D. lieteropelmous ; jlh, flexor longus hallucis ; fpd, flexor perforans digitorum 
 V, vinculum. 1 to XV, 1st to 4th toes. (Alter Stejneger). 
 
 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 fiexor tendons of 
 the toes are arranged differently in these different digital 
 arrangements, although not always identically in the same. 
 The arrangement usual ivi 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 
 
VERTEBRATA 
 
 93 
 
 
 ^ ^ 
 
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). 
 
 Tlie palatal structure must be consulted in attempting 
 the discrimination of the orders of birds. There are four 
 types of structure, as follows : First, the dromfeognathous 
 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 segithognathous, 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 Euornithes are as follows : 
 
 Steganopodes ; Phffitonidse, Fregatidse, Pelecanidae, Sulidse, 
 
 Phalacrocoracidse, Plotidse. 
 Colioidei ; Coliidse. 
 
 Chenomorphie ; Palamedeidae, Anatidee, Phoenicopteridffi. 
 Herodii ; Ibididee, Ciconiidse, Balsenicipitidse, Ardeidee. 
 Accipitres ; Cathartidse, Falconidse, Pandionidse, Strigidfe. 
 Heterospondyli ; Steatornithidse. 
 Coccygomorphas ; Cuculidse, Coraciidaj, Alcedinidse, Upupi- 
 
 dte, Musophagidse, Todidee, Momotidas, Bucerotidse, 
 
VERTEBRATA 95 
 
 Rhamphastidfe, Caprimulgidaj, Bucconida', Indicatori- 
 da\ 
 
 Psittaci ; Psittacidse. 
 
 Trogonoidei ; Trogonida\ 
 
 Gecomorphse ; Colymbidfe, Heliornithidse, Alcida3, LaridfE, 
 
 Procellariidse. 
 Grallee ; Chionidse, Thinocorida3, Glareolidte, Dromadida^, 
 
 Charadriidse, Otididse, Eurypygiidte, Rhinochetidas, 
 
 CariamidEe, Psophiidse, Gruidse, Rallidse. 
 Opisthocomi , Opisthocomidse. 
 Gallinse , Tetraonidte, Phasianidfe. 
 Pullastrse ; Cracidte, Megapodiidfe, Pteroclid;e, Dididffi, 
 
 Columbidce. 
 Micropodioidei , Cypselidse, Trochilidfe. 
 PicoidecX ; Picidse, Yngidte. 
 Passeres. This order is divided into five superfamilies, as 
 
 follows : 
 
 I. Tensor patagii brevis picarian ; Mnairoidei. 
 
 II. Tensor patagii brevis passerine. 
 
 A. Syrinx mesomyodian. 
 
 Tendons of flexor muscles of foot desmo- 
 
 pelmous ; Eurylsemoidei. 
 
 Tendons of foot schizopelmous; syrinx bron- 
 
 chiotrachial ; Tyran.iioidei. 
 
 Tendons of foot schizopelmous; syrinz tra- 
 
 chial ; Formicaroidei. 
 
 AA. Syrinx acromyoctian. 
 
 Tendons of foot schizopelmous ; Passeroidei. 
 
 The families of these superfamilies are the following : 
 Menuroidei ; Menuridse, Atrichornithidse. 
 Eurylxmoidei ; EuryleemidaB. 
 
96 
 
 COPE 
 
 Tyrannoidei ; Xenicidse, Philepittidse, Pittidse, Tyrannidse, 
 Cotingidse, Phytotomidse. 
 
 Formicaroidei ; Conopophagidse, Pteroptochidse, Formi- 
 cariidge. 
 
 Paftseroidei ; Alaudidse, Motacillidfe, Timaliidse, Liotrichidse, 
 Muscicapidse, Turdidse, Cinclidse, Troglodytidse, 
 Chamasidse, Hirundinidse, Artamidse, Laniidse, Paridge, 
 Paradisiidse, Corvidse, Sturnidse, Meliphagidse, Nec- 
 tariniidse, Certhiidse, Ploceidee, Tanagridae, Icteridse, 
 Fringillidse. 
 
 I.C. 
 
 Fig. 34.— Diagrams of a tracheal and a, broncliial syrinx, t.c, trachelo-clavicular 
 muscle. (After Newton. ) 
 
 Fig. 35— Lateral and dorsal views of the acrom'yodian syrinx of the raven (Corvus 
 corax), B, II, III, 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 
 
 wi;^^:^ 
 
 FlGr.36. — Diagrams of the elbow muscles of Icterus [A), showinp the passerine 
 type and NeninaC^)., showing the picarlan type. 6, biceps; emrl, extensor raeticarpi 
 radialis lon^us; h. humerus; s, shoulder; 5r, secondarv remiffes: /, triceps; tpb, 
 tensor patagil brevis; tpi, tensor patagii longus. (After Newton.) 
 
 To the Impennes but one order belongs. This is the 
 Ptilopteri : 
 
 Ilium not anehylosed with sacrum ; bones of 
 wing not foldable on each other; meta- 
 carpals not separated ; hallux directed for- 
 wards ; feathers scale-like ; vertebrte opis- 
 thocoelous ; Ptilopteri. 
 
 The Ptilopteri includes the recent family of Aptenody- 
 tidse or penguins. They first appear in the Upper Eocene 
 of New Zealand. No species, living or extinct, have been 
 found in the Northern Hemisphere. 
 

 C r iH 
 O — QJ 
 
 ■Sis 
 
 «2- 
 
 t« ^ s 
 
 I -§• 
 
 0< = 
 
VEETEBRATA 
 
 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 (J^Xurodon sasvus 
 Leidy). The mutilate type of limbs is seen in Cdotherium 
 cepAa/us 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 E, 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 cou- 
 
 /v-^ 
 
 GO) 
 
 Fig, 40. — Diagrams of tjfpes of mammalian dentition, from Osborn ; 1, haplodont ; 2, pro- 
 todont ; S, triconodont ; 4, tritubercular superior and inferior ; 5, tritubercular superior, 
 tuberculo sectorial lower; 6, quadritubercular superior, quinquetuberoulur inferior; 7, quad- 
 ritubercular, both jaws, pr, protooone ; 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, coossified 
 
 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 Pkototheria, 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 ; Multituherculata. 
 
 No teeth at maturity ; MonotremaCa. 
 
 The families are the following : 
 
 Protodonta ; Dromatheriidse. 
 
 Multituherculata; Plagiaulacidse, Chirogidse, Polymastodon- 
 tidse. Monotremaia ; Ornithorhynchidse. 
 
VEKTEBEATA 
 
 103 
 
 The geological range of these orders is shown by the 
 table below : 
 
 
 Protodonta 
 
 MuUituberculata [ Monotremata 
 
 Plistocene . 
 
 
 i 
 1 
 
 Neocene . . 
 
 
 
 
 Eocene . . 
 
 
 
 
 Cretacic . . 
 
 i 
 
 
 
 
 Jurassic . . 
 
 ! 
 
 
 
 
 Triassic . . 
 
 
 
 
 
 Carbonic . 
 
 
 
 Devonic . . 
 
 
 ■ 
 
 Siluric . . 
 
 
 
 Ordovicic . 
 
 
 
 
 Cambric . 
 
 
 
 
 Huronic . 
 
 
 
 
 Species of Protodonta 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 Multitubeeculata 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 Eocky Mountain region of 
 
104 
 
 COPE 
 
 Fig, 41. — Ptilodus medi<Bvus 
 Cope ; mandibular ramus ; a, b, 
 t; c, f. Original. FromPueroo 
 bed of New Mexico. 
 
 Fig. 42. — Ohirox plieatus Cope ; palate and molar 
 teeth, from below, 3-2 natural size. From Puerco bed 
 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 MoNOTREMATA are restricted to the Australian realm. 
 
 The EuTHERiA 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. 
 
VERTEBEATA 
 
 105 
 
 minute rudiments ; anterior limbs oar-like. 
 (MuHlata.) 
 
 Elbow joint inflexible ; carpals discoid, 
 and, with the phalanges, separated 
 by cartilage ; lower jaw without as- 
 cending ramus ; 
 
 Elbow joint flexible ; carpals and pha- 
 langes with close articulations ; 
 mandible with ascending ramus ; 
 
 (2) Cetacea. 
 
 (3) Sirenia. 
 
 AA. Posterior limbs present; ungual phalanges 
 compressed and curved on one or all the 
 feet.* ( Unguiculata.) 
 
 j3, Carpal and tarsal bones generally in linear 
 series. 
 
 Y, Teeth without enamel ; no incisors. 
 
 Limbs ambulatory ; hemispheres small ; (4) Edentata. 
 
 yy, Teeth with enamel ; incisors present. 
 
 No postglenoid process; mandibular con- 
 dyle not transverse; mastication 
 proal ; limbs not volant ; hemis- 
 pheres small ; 
 
 Anterior limbs volant; hemispheres small; 
 
 A postglenoid process ; mandibular con- 
 dyle transverse ; mastication orthal ; 
 scaphoid and lunar bones not coal- 
 esced ;t hemispheres small, smooth ; 
 
 A postglenoid process; limbs not volant, 
 with a scapholunar bone ; mastica- 
 tion orthal; hemispheres larger, 
 convoluted; (8) Garnivord. 
 
 (5) Glires. 
 (6) Chiroptera. 
 
 (7) Bunotheria. 
 
106 COPE 
 
 ^^, Carpal and tarsal bones alternating ; 
 faceted. 
 
 Anterior limbs prehensile ; mandibular 
 
 condyle and mastication transverse ; (9) Chalieotheria. 
 
 AAA. Posterior limbs present ; ungual phalanges 
 not compressed and hooked.J (Ungulata.) 
 
 /3, Carpal, and usually tarsal bones in linear 
 series.§ 
 
 Limbs ambulatory ; teeth with enamel ; (10) Taxeopoda. 
 
 /3/9, Carpal bones alternating externally ; 
 tarsals in linear series. 
 
 Limbs ambulatory, median digits 
 
 longest ; teeth with enamel ; (11) Toxodontia. 
 
 fij3^, Tarsal bones alternating ; carpals linear 
 or reversed diplarthrous. 
 
 Cuboid bone partly supporting navicu- 
 lar, not in contact with astragalus ; 
 no canine teeth ; (12) Proboscidea. 
 
 /3/9/3j8, 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 Mesonychidse, some Glires, and posterior feet of some Edentata. 
 tExcept Talpa and Erinaceus. JExcept in the Hapalidae. gExeept in Dendrohyrax. 
 IIExcept Pautolestidae. 
 
VEETEBEATA 
 
 107 
 
 ^ 
 
 
 
 
 
 
 
 
 
 
 
 
 1 1 
 
 i « 
 
 
 
 
 i 
 
 1 
 
 
 
 1 
 
 i 
 
 
 1 
 
 M 
 
 
 
 
 ■ 
 
 
 
 
 
 
 
 
 
 - 
 
 i 
 
 
 
 
 
 
 
 
 
 i 
 
 O 
 
 ! 1 
 
 
 
 
 
 \ 
 1 
 
 
 1 
 
 
 Ol 
 
 
 1 
 
 ! 
 
 
 
 
 
 
 
 
 QO 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 to 
 
 
 1 
 
 
 
 
 
 
 
 
 
 : 
 
 
 
 
 ! 
 
 
 
 
 
 
 
 
 
 j 
 
 
 i 
 
 1 
 1 
 
 i 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 1 
 
 
 
 C" 
 
 
 
 
 
 
 
 
 
 
 
 
 <i> 
 a 
 
 8 
 
 2 
 
 i 
 
 P-i 
 
 g 
 
 1 
 
 to 
 o 
 
 • 
 
 o 
 ■3 
 
 £ 
 
 
 
 .2 
 
 u 
 
 a 
 
 1-5 
 
 .2 
 H 
 
 
 1 
 
 
 
 
 
 1 
 
 
 
 
 ■3 
 
 
 
 
 
 
 a 
 
 
108 
 
 GOPE 
 
 The Marsupialia fall into two suborders, which differ as 
 follows : 
 
 Incisors |^ 
 Incisors [^ 
 
 Polyprotodontia. 
 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 j^^S^X^ <^ '^ '^ ^ h 
 (Fig. 43). Genera WZl^^^f&S^ s-^-'. 
 probably allied oc- 
 cur in the Post- 
 cretacic of North 
 America. No large 
 forms are known. ^ " ({ 
 
 The Diprotodontia /'f • ^^-P^o.I^~ f^a^^ cope ; anterior part of 
 ^ 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 Cetacea are represented by three suborders, which 
 differ as follows : 
 
 External nostrils at middle of muzzle ; 
 temporal fossse elongate, approxi- 
 mated; teeth ; Archseoceti. 
 
 External nostrils at base of muzzle ; tem- 
 poral fossse short, lateral ; teeth ; Odontoceti. 
 
VERTEBRATA 109 
 
 External nostrils at base of muzzle ; tem- 
 poral fossse short, lateral ; no teeth, 
 but transverse horny laminte on the 
 upper jaw; Mysticeli. 
 
 Fig. 44. — Zeuglodon cetoides Owen ; skull, much reduced. Prom Eocene of 
 Alabama. From Gaudry. 
 
 The Archseoceti 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 Squalodontidse, 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 Archseoceti. 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. 
 
 SiEENiA 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- 
 rastomidse 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 Halitheriidse 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 Rhytinidse all dentition has disap- 
 peared. 
 
 The Edentata have been generally restricted to the 
 Southern Hemisphere, although during the Neocene they 
 ranged as far north as the .^gean 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 Megatheriidse 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 
 Glyptodontidse were covered with an. immovable carapace 
 consisting of bony tesserse, 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. 
 
VEETEBEATA 111 
 
 The Glires 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 calcaneum ; ankle and 
 elbow not toogued 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 ; Scinromorpha. 
 
 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 coossified ; 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 
 
112 
 
 COPE 
 
 Pig. il .—Castwoides ohioensis Foster ; skull, much reduced. From Plis- 
 tooene of Ohio. Prom Hall and Wyman. 
 
 Pig. iS.—Flesiarctomys delicalissimus Leidy ; Skull. From Eocene of 
 Wyoming. Original. 
 
VEETEBKATA 
 
 113 
 
 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 thebeaver, 
 and spread into South 
 America. (Figs. 48-49.) 
 The M y o m o r ph a 
 (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 
 pocket-gophers to North 
 America. They cover 
 the earth, including a 
 few forms in Australia, 
 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 Chieopteea (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 
 
 Fig. 49. — Plesiartomys delicatissimus Leidy ; 
 a, humerus; 6, proximal ends of ulna and radius; 
 Cf d, tibia, distal end; e, /, astragalus ; all natural 
 size. From Eocene of "Wyoming. 
 
114 
 
 COPE 
 
 jaws interlocking in mastication; Animalivora. 
 
 Crowns of teeth obtuse, not interlocking in 
 
 mastication ; Frugivora. 
 
 Fig. SO.Sntoptychus crassiramis 
 Cope ; a pocket-goplier from the Middle 
 Neocene of Oregon, natural size. Origi- 
 nal, c, lower jaw from above. 
 
 Fig. 51. — Paloeolagus haydenii Leidy; 
 a rabbit from the lower Neocene of Col- 
 orado. Original ; natural size; a, ante- 
 rior half of skull, from below ; &, c, lower 
 jaw; (j, 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 BuNOTHEEiA present four suborders, which are char- 
 acterized as follows : 
 
 Molars 8 or more ; incisors several, with closed 
 roots ; 
 
 Molars 7 or less ; incisors with closed roots, 
 not enlarged ; fibula distinct ; otic bulla 
 developed ; 
 
 Pantotheria. 
 
 Creodonta. 
 
VEETEBRATA 115 
 
 Molars 7 or less ; canine generally small, in- 
 cisors generally enlarged; fibula generally 
 united with tibia ; Inseciivora. 
 
 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 
 (Hysenodon, 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- 
 ant 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, as in 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 
 
 COPK 
 
 ^ '^ 
 
 
 Fig. o2.—DeUatherium fundaminis Cope ; portions of skull, two-thirds- 
 natural size. From Puerco beds of New Mexico, a, skull profile ; 6, from 
 below : c, fragment of lower jaw, from above ; d, do., from outer aide. 
 
 Fig. 53. — Calamodon simplex Cope ; lower jaw, one-third natural size. 
 From Wasatch Eocene of Wyoming. 6, from above ; c </, an isolated molar. 
 
VERTEBEATA 
 
 117 
 
 uncommon in the Eocene of America, with one form in the 
 Eocene of Switzerland. Later than that age thej' 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 : 
 
 Plistocene 
 
 Neocene 
 
 Eocene 
 
 Cretacic 
 
 Jurassic 
 
 Triassic 
 
 Carbonic 
 
 Devonic 
 
 Siluric 
 
 Pantotbeiia 
 
 CreoiloMta 
 
 lusectivora Tillodoote 
 
 I 
 
 Ordovicic . 
 
 Cambric 
 
 Huronic . . . . 
 
 True Caknivora 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 Miacidse. 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 hyeenas in 
 the Upper Miocene, in the Northern Hemisphere. In the 
 Southern Hemisphere cats do not appear until the Plisto- 
 cene (Fig. 55), and hysenas do not occur in the Western 
 
 Fio. 54. — JVimravus gomphodus Cepe, two-fifths natural size ; left side of 
 skull. From Miocene of Oregon. 3-4, premolars ; 1-2, true molars. 
 
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 Taxeopoda 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 calcaneum ; 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. 
 
B 
 
 8 
 S 
 
 a< 
 I 
 
 2 
 
VERTEBEATA 121 
 
 Incisors growing from persistent pulps ; 
 
 anapophyses ; Daubentonioidea.* 
 
 Incisors with closed roots ; anapophyses ; Quodrumana. 
 
 Incisors with closed roots ; no anapo- 
 physes ; Anthropomorpha. 
 
 The Condylarthra are characteristic of the Postcre- 
 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- 
 dont, 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 coyerings of the terminal phalanges of all but the first digit in the 
 only known genus, are claw-like. 
 
122 
 
 COPE 
 
 Fig. Sl.—Necrolemur antiquusYilh; a lemur from the Upper Eocene of Fraicp, 
 natural size. From Filliol, 
 
 the Upper Neocene of ludia, 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 maybe expressed as follows : 
 
 
 1 
 
 2 
 
 3 
 
 4 
 
 5 
 
 6 
 
 Plistocene 
 
 
 
 
 
 
 
 
 
 
 Neocene- 
 
 
 
 
 
 
 
 Eocene . 
 
 
 
 
 
 
 
 
 
 
 Cretacic 
 
 
 
 
 
 
 
 
 Jurassic 
 
 
 
 
 
 
 
 Triassic . 
 
 
 
 
 
 
 
 Carbonic 
 
 
 
 
 
 
 
 Devonic 
 
 
 
 
 
 
 Siluric . 
 
 
 
 
 
 
 
 Ordovicic 
 
 
 
 
 
 
 
 Cambric 
 
 
 
 
 
 
 
 Huronic 
 
 
 
 
 
 
 
VERTEBRATA 
 
 123 
 
 The ToxoDONTiA 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 
 lophodout, and early began to be prismatic, and to grow 
 
 Fig. 68. — Dryopitheeus 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 ; 
 Femur without third trochanter; no clavicle ; 
 
 Typotheria. 
 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. 
 
 Fig. 59. — Cm'ypho'don elephaniopus Cope ; skull; from below, 2-9 natural size. 
 Original. Frona the Wasatcli 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. 
 
VERTEBEATA 125 
 
 Astragalus without head ; no third trochanter 
 
 nor superior incisors ; Dinocerata. 
 
 The Taligrada are confined to the Postcretacic epoch ; 
 the Pantodonda to the Lower, and the Dinocerata to the 
 Upper Eocene. 
 
 The order Proboscidia 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 (Ele- 
 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 epehs over 
 Xojth America and Europe. In America their remains are 
 
 a 
 
 Fig. 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 DiPLARTHRA iucludc 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 
 
4 
 
128 
 
 COPE 
 
 dimensions ; distal end of astragalus not 
 
 forming a ginglymus ; Perissodadyla. 
 
 Feet with the digits 3 and 4 predominating 
 and subequal ; astragalus with a distal 
 ginglymus ; Ariiodactyla. 
 
 The Peeissodactyla exhibit various series of forms in 
 which the toes diminish in number, from four in front and 
 
 Fig. 63. — Aphelops meyalodus Cope, one-sixtb natural size; B, inferior 
 view of cranium, Original. Hornless rhinoceros from the Loup Fork Neo- 
 cene, Colorado. 
 
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 (Suoidea) 
 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 (Booidea). Some types of the former and all of the 
 latter lack superior incisor teeth, and have the cuboid and 
 navicular bones coossified. The Booidea culminate in forms 
 with horns or bony processes of. the skull, which may be 
 permanent (Bovidse, or annually shed (Cervidse). 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 : 
 
VERTEBRATA 
 
 131 
 
 
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 IS 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 n 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 rt 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 V 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 ea 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 O 
 
 
 
 
 
 
 
 
 
 
 
 
 
 at 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 S 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 JS 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 03 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 3) 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 s 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 O 
 
 
 
 _ 
 
 
 
 
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 133 
 
 
 
 
 
 
 
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 0) 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 « 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 T) 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 — ^ 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 O 
 
 
 
 
 1 
 
 
 
 
 
 
 
 
 
 
 <-« 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 S 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 "o 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 fl 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 si 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 a« 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 cd 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 V 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 s 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 <: 
 
 1 
 
 
 
 
 
 
 
 
 
 
 
 
 
 ij 
 
 ,<? 
 
 
 
 
 
 
 
 
 
 
 
 
 
 >< 
 
 w 
 
 
 
 
 
 
 
 
 
 
 
 
 
 H 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 u 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 <! 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 O 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 ea 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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 « 
 
 3 
 
 
 
 
 
 
 
 
 
 
 
 
 
 H 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 Ph 
 
 u 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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132 
 
 COPE 
 
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 V- 
 
 Of the superfamilies which continue to the present day 
 the existing species are few in number, with the exception, 
 of the Booidea. All except the Booidea are now restricted 
 to tropical and subtropical regions, and while the majority 
 of Booidea have the same distribution, there are many 
 species m 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 in, 
 the Old World. This and the Booidea only appear in South, 
 America m the late Neocene. 
 
 As in the Perissodactyla there is a reduction of the digits. 
 
YERTEBRATA 
 
 133 
 
 ill most of the lines of the Artiodactyla, which reaches its 
 extreme in the Cameloidea, and in the Booidea. 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.) 
 
 Fig 6ii.— Anterior feet of Artiodactyla, with both series of carpals, except in 
 No. 4. Frnm Kowalevsky. No. 1, Hippopotamus ; 2, Hyopotamus ; 3, Dorcathe- 
 rhim : 4, Gelocus ; 5, Cervus. 
 
 Tiie families embraced in the orders of Eutherian Mam- 
 malia aie the following : 
 
 Marsupialia; (Polyproiodontia) ; Triconodontidse, Amphi- 
 theriidffi, Myrmecobiidse, Dasyuridss, Didelphidse, Per- 
 amelidffi; (Diprotodoidia) ; Phascolomyidse, Phalangist- 
 idse, Tarsipedida3, Diprotodontidje, Macropidse, Thyla- 
 coleonidge. 
 
 Cetacea; (Archieoceii); Zeuglodontidse ; {Oduidoceti) ; Squ&l- 
 odoiitidee, Platanistidse, Physeteridse, Deli>liiind£e,(il/^s- 
 tacoceti) ; Baltenida:. 
 
 Sirexia; Prorastomidse, Halitheriidie, Manatidte, Halicori- 
 da;, Rhvtinidffi. 
 
134 COPE 
 
 Edentata ; Orycteropodidse, Manidte, Bradypodidse, Mega- 
 theriidse, Myrmecophagidse, Dasypodidse, Glyptodon- 
 tidse. 
 
 Gliees ; (^Hystricomorphse) ; Paradoxomyidse, Hystricidse, 
 Echinomyidse, Octodoiitidse, Capromyidee, Caviidse, 
 Chiuchillidse ; [Sciuromorpha) ; Sciuridse; (Myomorpha) ; 
 Dipodidse, Muridse, Myoxidse, Saccomyidse, Microtidte, 
 Lophiomyidse, Bathyergidse; (Lagomorpha) ; Leporidse. 
 
 Chiroptera ; (^mmaHi;ora) ; Phyllostomidse, Desmodontidge, 
 Rhinolophidse, Noctilionidse, Vespertilionidse, Embal- 
 lonuridse ; (Frugivora) ; Pteropidse. 
 
 BuNOTHERiA ; {Pantotherio) ; Aniblytheriidse ; (Creodonta) ; 
 INfesonychidge, Esthonychidse, Arctocyonidse, Miacidae, 
 Hyseiiodonlidae ; (Insedivora) ; Leptictidse, Centetidae ; 
 Galeopithecidse, Tupseidse, Solenodontidse, Macrosceli- 
 didse, Talpidse, Adapisoricidaj, Mj'thomyidae, Scalopidse, 
 Clirysochloridse, Erinaceidge, Myogalidae, Soricidae ; 
 {Tillodonta) ; Tillotheriidie; (Tseniodonia) ; Ectoganidae, 
 Stylodontidae. 
 
 Carnivora; {Fissipedia) ; Cercoleptidae, Procyonidte, iElii- 
 ridae, Canidae, Bassarididae, MustelidsB, Protelidae, Arc- 
 ticidae, Viverridffi, Cynictidae, Suricatidae, Cryptoproc- 
 tidae, Nimravidae, Felidae, Hyaenidae ; [Pinnipedia) ; 
 Phocidse, Otariidae, Odobsenidae. 
 
 Chalicotheria ; Chalicotheriidae. 
 
 Taxeopoda ; (Condylarthra) ; Phenacodontidae, Pleuraspido- 
 theriidae, Meniscotheriidae ; (Litopterna) ; Proteiotherii- 
 dae, Macraucheniidaj, Astrapotheriidae ; (Hyracoidea) ; 
 Hyracidae ; (Daubentonioidea) , Chiromyidae, Mixodec- 
 tidae; (Quadrumana) ; Adapidse, Anaptomorphidae, Tar- 
 siidae, Lemuridae, Hapalidae, Cebidae, Cercopithecidae ; 
 (Anthropomorpho!) ; Simiidae, Hominidae. 
 
 Toxodonita; [Typotheria) , Atryptheriidae, Interatheriid®, 
 
VEETEBRATA 135 
 
 Protoxodontida?, Mesotheriidse ; {Barytheria) ; Xotodon- 
 tidfe, Toxodontidse. 
 
 Proboscidia ; Dinotheriidse, Elephantidse. 
 
 Xmblypod A : (Taligrada) ; Periptj^chidse, Pantolambdidse ; 
 (Pantodonia) ; Coryphodontidee ; (Dinocerata) ; Uinta- 
 theriida?. 
 
 DiPLAETHRA; {Perissodactylo) ; Lophiodontidse, Triplopodi- 
 dse, Ctenopidse, Hyracodontidse, Rhinoceridse, Tapiridse, 
 Lambdotheriidse, Menodontidee, Palseotheriidse, Equi- 
 dtp ; {Aiiiodadyla;) ; Trigonolestidse, Eurytheriidse, An- 
 oplotheriidse, Dichobunidse, Csenotheriidse, Anthraco- 
 theriidte, Xiphodontidse, Suidse, Hippopotamidse, Mery- 
 copotamidEe, Dichodontidse, Oreodoatidse, Poebrotherii- 
 dse, Protolabididse, Camelidse, Eschatiidee, Tragulidse, 
 Moschidfe, Bovidse, Cervidse. 
 
ERRATA. 
 
 Page 95, 5th line from bottom, for acromyoctian read 
 acromyodian. 
 
 Page 1 24, 8th line from bottom, for Fig. 60 read Fig. 62. 
 
 Page 134, under Bunotheria, transfer Leptictidse and 
 . Centetidse from Insedivora to Creodonta. 
 
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