i aii a aie ae eran aCe ene EY er ea, we gnisa # roe Peter nnte ty Ae Ki} ean) Het ean i 4 yy Cin pes Bec aren pea ares Asa een iii i fehseahse ete aS a ahs Bb oO nn ee ah | u | fi i i ; ete ehh Pee a aio te i a aes 2 “ oe See eat et : + : I ty y 4 Pies Ht U a si: fae) er a + r ' A i f Mea? bat id ; How) Hi io 3 Py i Mi aaep ‘ i ; pte f © i Hl + ey oat ; Nett tae ‘ ih ‘a ae aa ! rtd a Yi i fi i i H Lie zi i r i aa a ER oe ye, f Springs iy i Bat : Paya ont] i ‘i A f H ; rae A iy d ; / } re ts i rt ti s i 9 ! 7 3s | maine VarT) tN HPO Bee dt ie i i} ba F f ig heat iyi eae er a 5 al a4 Prieee token og ¢ 4 i i (Pitiy ee teat H ' 1 ( i tee itd fy } Vf me) PLUMB EDD i f MOR revit z A aA urues iat ane ERAT PUB ! { MEV TAC TET PATE NA f i ; ne Aare AP AON EMRE E THLE UENO Lives Attn ia HTP easton are} ‘ marie ed tes eeen an Mb Rane Uae Aa i Mt SH dhagldsds Maat LRP eE aes ru RRA '' ‘3 hag . i a an i i) t S te Saree ''Table of Contents A memoir upon Loxolophodon and Uintatherium The horse, past and present Paleontology. Progress of the past decade (1911-1921) Craniometry of the Equidae Tetraplasy, the law of the four inseparable factors of evolution '' '' '' '' ''es % Ae Swen COG oe ''a , CONTRIBUTIONS | FROM THE EM. MUSEUM OF GEOLOGY AND ARCHAOLOGY OF THE COLLEGE OF NEW JERSEY. VOLUME I, PRINCETON, NEW JERSEY. '' '' CONTRIBUTIONS FROM THE E. M. MUSEUM OF GEOLOGY AND ARCHZOLOGY OF THE COLLEGE OF NEW JERSEY. The E. M. Museum of Geology and Archeology at Princeton College con- tains not only an ample collection of specimens for college instruction, but also a large amount of material for more advanced study in the shape of fossils new to science, mainly collected by the college scientific expeditions of 1877 and 1878. It has seemed desirable for the advancement of science, as well as in justice to the palzontologists and biologists connected with the Museum, that the results of their studies should be placed in a permanent and suita- ble form before the scientific public. It is, therefore, no small gratification to the Director of the Museum to be enabled to enlarge the system of publication begun by the bulletins in 1878, under the auspices of the Museum. The mode of publication will be the same as that adopted by most of the Scientific Institutions, viz.: bulletins in octavo for advanced work and minor notices, and memoirs in quarto for later and more mature researches. The contributions will be based chiefly upon the study of the palzontologi- cal and geological collections made by the past, and those which may be made by the future scientific expeditions from the college. The other portions of the collections, however, contain a considerable amount of new material, both in fossil botany and zoology, for study and publication. Biological and Geological Memoirs, as well as communications on the Era of Pre-historic man, will be embraced in the field of research covered by the publications of the Museum. A: GUYOF. Director of the E. M. Museum. PRINCETON, NEW JERSEY, July 10, 1881. J 767488 fi '' ''CONTRIBUTIONS FROM THE E. M. MUSEUM OF GEOLOGY AND ARCHEOLOGY OF THE COLLEGE OF NEW JERSEY. \ Vou. i, No. & A MEMOIR UPON LOXOLOPHODON ann UINTATHERIUM Two GENERA OF THE SuB-ORDER DINOCERATA BY HENRY F. OSBORN, Sc.D. Lecturer on Biology. ACCOMPANIED BY A STRATIGRAPHICAL REPORT OF THE BripGER BEDs IN THE WasHakie Basin BY JOHN BACH McMASTER, C.E. Instructor in Civil Engineering. PRINCETON, NEW JERSEY, 1881. '' ''INTRODUCTORY, THE study of two genera or the Dinocerata forms the first of a series of Palzontological Memoirs that will be published by the E. M. Museum of Prince- ton College. The Museum already contains a very interesting collection of Tertiary fossils, chiefly procured by parties going out from the college. The first of these field-parties was formed in the summer of 1877. After four weeks in Colorado they camped for the remainder of the season in the country lying near Fort Bridger, Wyoming Ter. In 1878 a second party, smaller but better equipped for collecting fossils, returned to the same field, extending their jour- ney into the Loxolophodon beds of the Bitter Creek country, and making their way back to Twin Buttes, and thence to Fort Bridger, after a short trip up Ham’s Fork. During both seasons the principal collections were made in the Bridger Beds (Upper Eocene), so that the fauna of this group is finely repre- sented in the Museum. In the first Bulletin the collections made during the summer of 1877 were described. The writers there stated that they were fully aware they were upon ground already quite familiar to science. The same thought occurs in some measure in connection with this Memoir. Two distinguished palzontologists in this country, with large collections at their command, have been contributing to a general knowledge of the Bridger fauna for years. But many features that have not been brought to light are discovered among our collections, so these studies are offered, in the belief that they con- tain much that is original. At the outset, the writer acknowledges with pleasure his indebtedness to the works of these authors, to whom full credit is accorded in the text The horizon of the genera which are the subject of the present Memoir is the Bridger group, Upper Eocene. They had quite an extensive range in the Tertiary basin in which these beds were formed, and were the largest in size of the many varieties of animals which roamed on the borders of the great Eocene lakes. The sub-order of Dinocerata is of more interest from a consideration of the many varieties which it includes than from a phylogenetic point of view; for its genera possess many characters by which we must consider them as form- ''oO: t ; INTRODUCTORY. eo aaee iag!an aberrant group which became extinct long before the close of the Ter- tiary. The indirect relations of these genera with the elephant tribe were apparent from the first, but there is no foundation for a belief that they came in the line of forms ancestral to the Proboscidia. The Stratigraphical Report by J. B. McMaster, leader of the expedition of 1878, made from his notes and sections of the strata of the Bridger beds as ex- posed in the Washakie basin, points chiefly to the present condition of the strata as they are left by erosion and denudation. As it indicates the level at which the fossils described in the text were discovered, and embraces an outline of the principal topographical features of the country it is published in connection with this memoir. The thanks of the writer are gladly expressed to Dr. Arnold Guyot, Director of the Museum, for frequent advice and encouragement, and to Dr. William B. Scott for several suggestions of value. Dr. Franklin C. Hill also rendered much kind assistance in the Museum. i. FO. PRINCETON, Fly 1881. ''Loxolophodon and Uintatherium BY HENRY F. OSBORN. '' '' GEOLOGICAL INTRODUCTION. THE testimony of the rocks is nowhere more marvellous than in the cen- tral deserts of Wyoming Territory. Even the view obtained in passing rapidly through this barren country cannot fail to arouse interest; for to the thoughtful traveller it seems that Nature here tells her story with peculiar sim- plicity. The great elevation of the whole desert above the sea, the noble and grotesque forms which centuries of wind and rain have carved out of the sandstone bluffs, the brilliant colors which enable the observer to follow the strata as they appear and disappear in bold outlying buttes and receding curves, are all highly unique characters. The name Mauvaises Terres conveys an impression of the singularly forbidding aspect of this country—of the vege- tation, sparse from the protracted summer droughts, and of the naked strata over which, in the rare atmosphere, the eye easily ranges from mile to mile. Add to this the discovery that in these rocks are buried many generations of animals whose genealogy can often be clearly followed, and it is unde- niable that here, if anywhere, the records of an ancient land are written in char- acters which cannot be mistaken. When the relations of the desert plains to the high mountains sur- rounding them at all quarters became known, and the relative ages of the mountains and the plains were ascertained from the fossils they contained, it was a natural conclusion that these sedimentary beds were formed in inland waters by erosion from the hills freshly emerged from the Secondary seas. The earnest labors of Hayden, King, and Powell have thrown light upon the different aspects of this geological problem. To the early studies and ex- plorations of Dr. Leidy and the persevering efforts of Professors Cope and Marsh are due most of our knowledge of the animals which these rocks entombed. With these researches at hand, and those of Prof. Lesquereux, who has devoted many years of study to the ancient flora, it follows that we now have a considerable knowledge of the condition of animate and inanimate nature of Eocene times in this region. The ancient geography and ancient fauna should be placed and studied together even more intimately than has been done hitherto, in order to afford the mind a comprehensive view of the whole. For facts of the structural pecu- ''10 £. M. MUSEUM MEMOIRS. liarities of species and genera are of diminished value when isolated from an investigation of their natural and physical environment. We cannot grasp the meaning of the peculiar dentition, of the anomalous features of the skull, or, what is more important, of the modifications of the ankle-joint and feet; in short, of the adaptability of the skeleton to certain functions, unless we first inquire into the nature of the moist and semi-tropical country which formed the habitat of these Eocene animals. However vague and unsatisfactory this inquiry be, it affords a collection of additional facts upon which to advance. Clarence King in his recent work’ has given an original and interesting account of the early geological history of this region. And this memoir upon the Dinocerata cannot be more appropriately introduced than by a résumé of his narrative, accompanied by several observations of our own. At the close of the Cretaceous period, the country lying between the pres- ent Rocky Mountain system and the Wahsatch range lay open to the sea on all sides. By the elevation of the land which now forms the summit of these ranges, a great area became enclosed on the East and West, and the convergence of the ranges somewhat in crescent shape formed a northern shore. To the South the waters extended into Colorado and New Mexico. The discoveries of Hayden and Marsh have confirmed and increased this southerly extension. Its exact limits are not yet ascertained, nor do they, although interesting, im- mediately belong to the present subject. Thus, at the beginning of the Eocene an inland sea was formed, almost as broad as Lake Erie, enclosed on the North by the spurs of the great system of hills which formed the eastern and western barriers. These spurs are repre- sented by the Bear River, Wind River, and Sweet Water mountain systems of to- day. On the southwestern shore was a great island, the Uinta chain, also of Secondary rocks, lying in an east and west direction. This inland lake therefore filled the basin which is at present drained by the head-waters of the Green and Colorado river systems, and the Green River now cuts through the eastern base of the Uinta range, marking the central point of southern outlet probably from the earliest enclosure of the basin. These waters, which King has named the Ute Lake, gradually lost their saltness by the drainage from the hills sur- rounding them, while the mouth, or southern outlet, was slowly losing its direct communication with the sea. This change from marine to lacustrine con- ditions was preceded by the extinction of many of the reptilian types of the Cretaceous. Underlying the waters of the lake, somewhat tilted by the upheavals which formed the shores of the basin, were the upper members of the Cretaceous series, the Laramie beds. Here and there also were islands of earlier Second- 1U. S. Geol. Explor. of Fortieth Parallel. Vol. I. Systematic Geology. ''LOXOLOPHODON AND UINTATHERIUM. II ary rock; these had not shared in the general submergence which preceded the deposition of the Laramie. On the flanks of the Wahsatch hills were Mesozoic and Carboniferous beds, and the same strata were exposed on the Uinta slopes. The degradation of these older strata, if we can judge from the investigations of Major Powell’ in the Uinta range, taking place farz passu with upheaval, was on a grand scale. Vast quantities of water were borne by the winds from the Pacific Ocean, as yet uninterrupted by the Sierra range, and were condensed on the Uinta slopes. The sediments thus collected formed the Vermillion Creek group of rocks—so named from their prevailing red color. They lie unconformably over the Laramie strata, and are highly fossiliferous. The most characteristic mammal at this period was Coryphodon, five-toed and with a full dentition. The presence of this with Hyracotherium, Hyopsodus, Opisthotomus, Amblyctonus, Clastes, and others, has led Prof. Cope’ to indicate the close paral- lelism between these beds and the Suessonian or Orthrocene of Europe, in which these genera also abounded. The middlemost member of the Eocene series, the Green River group, was deposited over a still larger area; for after the Vermillion Creek beds were formed, the western, and in some degree the eastern, borders of the Ute Lake subsided, enlarging the basin to nearly double its former size. This second lake, named the Gosiute Lake, in which the Green River beds were slowly deposited, was an almost uninterrupted sheet of water, extending about three hundred miles on the fortieth parallel, and one hundred and fifty north and south. While the close of the Vermillion Creek deposition was marked by an expansion of the basin borders, the close of the Green River deposition was marked by contraction; so that the third member of the Eocene series, the Bridger beds, was deposited in a smaller basin. According to Powell, a dry- land period also intervened. It followed that when the limits of the third lake were defined, they were on all but the southern side within the area of the Vermillion Creek beds. The oscillations of level which preceded it and the contour of the present exposures of the beds, formed in this third or Washakie Lake, leave considerable doubt whether it was a continuous sheet of water enclosed in a single basin or several smaller basins, with a common southerly outlet. This question will be resumed further on. A liberal estimate of the total thickness of these three groups of Eocene rocks is 9500 feet. Powell estimates them at 6000 feet. They form the lower, mid- ! Geology of the Uinta Mountains. Washington, 1876. * Ext. Bull. U. S. Geol. and Geog. Survey, Feb. 28th, 1879. The Relations of Horizons of Extinct Vertebrata of Europe and North America, ''12 £. M. MUSEUM MEMOIRS. \ dle, and upper Eocene. Just south of the Uinta range is another small group of rocks, of about 500 feet in thickness, which caps the Eocene and forms an approach to the Miocene. The age of these beds has been determined by their fauna. Their position and relation to the Uinta chain, as well as that of the other groups, is made clear in the accompanying sketch-map. The history of these Tertiary lakes and the present confines of the various beds can be followed in this map, which affords an admirable idea, also, of the upheavals which formed the great rim of the basin. It will be kept in mind that the country circumscribed has been elevated since Eocene times to 6000 and 7000 feet above the sea by the general uplifting of the continent, and that the mountain ranges of to-day rise to twice this level. The total thickness of the Bridger beds is about 1500 feet, presented in two or three comparatively contracted exposures. These overspread two main tracts: one in the Bridger basin, lying wholly east of the Green River; a second to the west of the river, situated in the Washakie basin; a third, smaller tract lies southeast of Vermillion Creek. That the Bridger beds were deposited in a much smaller sheet of water than the Green River beds is evident at once from the nature of the rock, and quite apart from a consideration of their extent. The Green River beds are largely calcareous, fine fissile shales, some- times containing carbonaceous matter. The lower members are limestones. The uppermost strata of the Uinta were Mesozoic rocks; and King suggests that the earlier erosion (forming the Vermillion Creek strata) must have worked these off, exposing the calcareous strata of the Carboniferous rocks, about the inception of the formation of the Green River beds. The presence of lime in these beds is thus ingeniously accounted for. The shales are often of delicate texture, and rival the Solenhofen slates in the exquisite impressions of leaves, fishes, and crustacea which they contain. They indicate that the streams feeding the Gosiute Lake were of a rather sluggish nature, and heavily charged with fine silt. There are occasional calcareous strata among the Bridger beds, but they are essentially a sand formation. The reader is here referred to the stratigraphical section accompanying this memoir. Alternating with the fine clayey and sandy beds are coarse gravelly beds, indicating streams of con- siderable size and rapidity which were suited to carry such heavy materials far into the centre of the lake. Rarely they contain remnants of leaves, and sel- dom or never, the remains of fishes. In the coarser beds we usually found only the remains of larger vertebrates, and those considerably dissociated and scattered by violent aqueous and other agencies. There is for the above reasons no room for doubt that the Bridger beds were formed in a much smaller basin than the Green River. The question remains, were the later beds formed in a single basin or in a number of separate basins '' LOXOLOPHODON AND UINTATHERIUM. 13 which the waters occupied as they slowly retired? King leaves the matter in doubt, expressing however, the opinion that it is improbable that erosion could have totally removed all traces of the Bridger rocks from the country lying between the Washakie and Bridger basins. He therefore rather favors the view that the waters in which the Bridger beds were formed were not con- tinuous. This view, it appears to the writer, may be sustained on two grounds. First, the nature of the rocks in the Washakie and Bridger basins is somewhat different ; secondly, the character of the fauna is more dissimilar than we should expect to find it at opposite ends of a continuous shore. The prevail. ing colors among the Bridger Basin strata are light grays and drabs, with an occasional green. In the Washakie beds the colors are much more brilliant; many of the strata are a very brilliant green, others have a brownish-red hue. The manner of weathering is dissimilar ; in the Washakie basin the occasional beds of harder rock form bold cappings for the softer strata beneath, result- ing in mushroom-like forms, which are rarely seen in the Bridger basin. More important is the evidence derived from the character of the fauna, found in the deposits of the two basins. During the summer of 1877 the Princeton party collected wholly in the Bridger-basin area. The dominant species were Palzosyops and Hyrachyus. The remains of Uintatherium (Dinoceras) were quite abundant; and two species of the Carnivora, a small rodent, and numerous quadrumanous forms were obtained. A doubtful Artiodactyle form was secured, referred to the camel tribe. A second tour through these beds, in the following summer, resulted in the collection of other individuals belonging to the same orders. In the summer of 1878, the party spent six weeks in the Washakie Basin beds. Here the number of Loxolophodon remains was sur- prising, especially in the upper strata. Not a single specimen of the allied genus, Uintatherium, was obtained. Prof.Cope has, however, reported a single species of Uintatherium from these beds, on a rather doubtful determination. He found here several species of Loxolophodon. It certainly appears that the latter genus ranged exclusively in this region, while it is possible that some species of Uintatherium found their way here, but not in great numbers. In the lower Washakie-basin strata, in a matrix quite different from anything found in the Bridger, we discovered the skeleton of an Achenodon, one of the Bunodont Artiodactyla, and an unspecialized member of the Rhinoceros family. Palgosyops was rarely found. Hyrachyus has been obtained by Cope in these beds. It is unfortunate that the localities in which the Bridger- group fossils have been discovered have not been more frequently published, as data of this description would greatly aid in determining the extent of the third Eocene lake, as well as giving a clue to the migrations of these forms. Our own discoveries point to the distribution of several genera over the region ''14 £. M. MUSEUM MEMOIRS. of both basins, also to the fact that several genera which are common in one basin are rarely or never found in the other. The three exposures of the Bridger (Br.) beds are indicated in the map. First, that of the Washakie basin, which has an east and west extension of about twenty-five miles by about sixteen miles north and south. It is sur- rounded by a narrow band of Green River (Gr.) beds. Passing to the south across Vermillion Creek, is another small exposure of much more contracted dimensions. Much larger than either are the beds lying in the Bridger basin, extending east and west about sixty miles with an expansion to the north which has not been surveyed. These beds are largely surrounded by those of the Green River period. At some points, however, they overlap the Vermillion Creek (V.) beds. On the flanks of the Uinta Mountains they come in contact with the still older Mesozoic rocks. If the view advocated above be adopted, they represent two large sheets of water lying east and west of the present bed of the Green River and draining to the south. A low line of hills made up of Vermillion Creek and Green River rocks separated them, and was suf- ficient to account for the variations of the fauna observed in the two basins. It is well to add that there may have been some slight differences in the character and date of their deposition which would attract one group of animals rather than another. The comparison of the Suessonian of Europe with the Vermillion Creek or Wahsatch of the American Eocene has been extended by Prof. Cope, with more restrictions, into a further comparison of the Bridger with the Parisian beds. They possess many genera in common—chief among which are Hyeno- don, Adapis, Plesiarctomys, Hyrachyus, Tapirulus, and Anaptomorphus. The Paris- ian beds have, moreover, a large number of Selenodont Artiodactyla of primitive type, but are distinguished by the absence of the Z7//odonta and Dinocerata which are so characteristic of the Bridger epoch. Wandering on the shores of the Bridger or Washakie lakes were great herds of Palgosyops, the American Palgotherium, and other perissodactyle ani- mals more closely allied to the tapir. There were also primitive horses of the size of the modern fox. Many lemurs and small rodents testify to the presence of wooded vegetation. The Carnivora were represented by several genera of formidable proportions. Dominant in size were the huge Dinoce- rata, Loxolophodon, and Uintatherium, standing a little lower than the elephant, but equally long in the body. Crocodiles and many smaller saurians were abun- dant on the shores, and turtles as large as the modern loggerhead floated on the water, and there were many varieties of land and aquatic birds of small size. The fauna was that of a semi-tropical country, and the climate was very moist; for, alternating with the finer clays, are coarse gravelly beds indicating periods ''LOXOLOPHODON AND UINTATHERIUM. 15 of floods when the heavier materials were transported far into the lakes. All these conditions are in striking contrast with the meteorological phenomena of this desert land at the present day. The flora of the upper Cretaceous period in this region presents an inter- esting mingling of hardy northern plants with those of a more tropical character. The plants of the Laramie are well known. to resemble those of the European Tertiary formations It is not improbable that they survived the changes accompanying the upheaval of the land and clothed the borders of these lakes, although the manner of sedimentary deposition during the Bridger epoch did not often admit of the preservation even of the littoral plants. Here and there, however, are found the fragmentary remains of reeds and other water-plants. The position of some of these is indicated in the strati- graphical section accompanying this memoir. The abundance of polydactyle forms, or spreading feet, among the mammals adds probability to the natural supposition that the lakes were bordered by wide-stretching marshes. As to the causes of extinction there is much evidence to show that they were slow and natural, very much such causes as we should observe on the borders of similar lakes in Southern Africa or elsewhere. The manner in which the parts of a skeleton are found widely scattered, the occasional evi- dence from the position of the limbs in the rock that the animal has been mired, the absence of large numbers of bones in any one locality, all indicate a long period in which the struggle for existence was not intensified by rapid geologi- cal changes. The great variety of species which succeed one another and are closely allied in structure afford further proof of a long undisturbed period in which these gradual modifications could arise. The prevalence of weapons of defence, such as the great tusks and cranial protuberances of Loxolophodon, the sharp dentition and powerful muscles of Palgosyops, indicated by the skull, the tusks of Achenodon, and the full dental series of a primitive Rhinoceros, all go to show that the greatest contest was between the animals of differ- ent families rather than against unusually violent forces of nature. Enough has been written to clearly introduce the short study of the Dinocerata which follows. They were a class of animals which present fea- tures of great interest, and so numerous and characteristic of this geological period that one writer has chosen to name the whole series of Bridger rocks after them. The two species which will be described represent two of the principal genera of the sub-order; namely, Uintatherium and Loxolophodon. No explanation is needed of the length of this geological sketch with its accom- panying maps, further than to say that it is hoped it will also serve to assist the understanding of some of the other Bridger Eocene fossils. ''THE DINOCERATA. The Dinocerata form a Sub-order of the Amblypoda, an Order which Cope’ has defined as follows: “Mammalia with small cerebral hemispheres which leave the olfactory lobes and cerebellum exposed. The feet short and plantigrade, with numerous (in the known genera, five) digits, terminating in flat, hoof-bearing, ungual phalanges. The seven bones of the carpus distinct, the unciform articulating with both lunar and cuneiform. The astragalus flat, without trochlear surface, and attached to the tibia with very little freedom of movement; its distal extremity divided into two facets, one for the navicular and the other more or less for the cuboid. Molars inserted with enamel, with wide crowns and transverse crests. A post-glenoid process.” The Order Amblypoda falls into two Sub-orders: I. A third trochanter on the femur, and a fossa for the round ligament; no alisphenoid canal; superior incisors present-————— __ PANTODONTA. Il. No third trochanter, nor fossa for the round ligaments; an alisphenoid canal; no superior incisors DINOCERATA. The sub-order Pantodonta includes the genera Coryphodon and Bathmodon. The former is the most characteristic fossil of the Wahsatch group, in the lower Eocene. It has, as the ordinal name indicates, a complete dentition, and is also marked by a very unmodified type of foot. According to Huxley and others, these unspecialized characters and its geological position combine to place it at the bottom of the Perissodactyle scale. The sub-order Dinocerata includes at present several genera which widely vary in size and seem to have been peculiar to different portions of the great Eocene Rocky Mountain basin in which they lived. A distinctive character common to all is the possession of a number of paired protuberances upon the upper surface of the head. Another general feature is found in the downward projections from the rami of the lower jaw which were opposite the huge upper canines. These projections, or flanges, underwent various modifications. The nasal bones also supported a small pair of knobs. It is chiefly upon the position of the cranial protuberances, the conformation of the lower jaw, the develop- 1Cope. Wheeler’s Survey, Vol. IV. pp. 178 e7 seg. '' LOXOLOPHODON AND UINTATHERIUM. 17 ment of the nasal bones, and the relations of the bones forming the hard palate that the following genera have been determined. SYNOPSIS OF THE GENERA OF THE DINOCERATA, A. Lower jaw with powerful downward flange, wide diastema between lower canine and incisor series. Three premolars on lower jaw. Premaxil- laries send in narrow plate to form forward portion of hard palate. Nasal tuberosities do not overhang nasal tips. Median protuberances anterior to orbit Ointatherium. B. Lower jaw has a slight, narrow, downward convexity opposite upper canine, wide diastema between lower canine and incisor series. Three premo- lars in lower jaw. Premaxillaries send in a broad plate to form the forward portion of the hard palate. Nasal tuberosities overhang nasal tips. Median protuberances above or anterior to orbit Loxolophodon. C. Ramus of lower jaw is arched downwards from the symphysis to the angle. The incisors, canine, and anterior premolar form a continuous series. Between the anterior premolar and the remainder of premolar series is a dias- tema. There are four premolars in the lower jaw————_____ Bathyopsis.' The first of the above genera was Uintathertum, discovered and described by Dr. Leidy in the Bridger Beds, in the exposures referred to above as lying wholly in the Bridger basin or west of the Green River of Wyoming. Prof. Marsh, who reached this field soon afterwards, rapidly obtained a large collec- tion of the remains of these huge mammals, the greater part of which he referred to a distinct genus, Dinoceras. This is considered in this report as synonymous with Uintatherium, as we find the generic characters nowhere distinctly stated. The second genus, Loxolophoden, was discovered by Prof. Cope, in 1872, in the Washakie exposures of the Bridger beds. He gave the generic characters soon after the discovery.’ The individuality of the genus was strengthened in 1878 by the fortunate discovery of the lower jaw by the Princeton party. The third genus, Bathyopsis, has been quite recently obtained in the country surrounding the head-waters of the Wind River. Cope is doubtful whether to place these beds among the Wahsatch or the Bridger Eocene groups. Some of the genera they contain are found in both the lower and the higher groups. At all events, the discovery of this genus gives the sub-order a still wider geographical distribution. A fourth genus, Hodasz/eus, has been based by Cope on some very short cervical vertebrae. This seems of rather uncertain position. 1Cope. Bull. U. S. Geol. Survey, Vol. VI. pp. 194, 195. 2 Proceedings American Philosophical Society, 1872, p. 580, ''18 £. M. MUSEUM MEMOIRS. UINTATHERIUM. A synopsis of the numerous species of Uintatherium (Dinoceras) will not be attempted here. From numerous specimens in our collection the following species has been positively identified : A. No tubercle at the entrance of the valley between the lobes of the last upper molar. Nasals divided by a deep groove; slender zygomatic arch. Temporal fossa not continued far behind posterior protuber- ances U. Leidianum. These characters distinguish it from U. robustum' (Leidy). This individual is figured in Plate II. In the first Bulletin from this Museum was described another species of Uintatherium, which we called U. princeps. This species is no longer recog- nized as being clearly distinct. In the multiplicity of varieties that have been found by different parties, there are at present no means of ascertaining how far the characters upon which species are constantly based are the outcome of differences of age and sex. The development and proportions of the pro- tuberances especially belong to the class of cranial modifications which may have been wholly subject to differences in sex. The following synopsis of the species of Loxolophodon is therefore offered provisionally. LOXOLOPHODON. Nasals deeply cleft in front and much produced beyond nasal tips. Frontal protuberances large and placed directly above the orbit. Foramen incisivum widely cleft in front. (Occiput unknown.) L. cornutus. Nasals wide and less cleft. Frontal protuberances above orbit. Occiput low and broad L. galeatus. Nasals uncleft and barely produced beyond nasal tips. Frontal protuber- ances small and anterior to orbit. Occiput high and narrow. L. Speirianum, sp. nov? The two unusually fine skulls of Uintatherium and Loxolophodon in our museum have afforded an admirable opportunity for a comparison of these two well-determined genera of the Dinocerata. The distinguishing features of 1Leidy. Proc. Ac. Nat. Sc., 1872, p. 169.—Cont. Ext. Vert. Faun. of Western Territories, p. 93. 3 Dedicated to the discoverer of the head, Mr. Francis Speir, Jr., a member of the two expeditions. ''LOXOLOPHODON AND UINTATHERIUM. 19 the Sub-order have already been pointed out, also the characters distinguishing the genera known to the writer. In this memoir a careful description of the skull of Loxolophodon Spetrianum will be given, supplemented here and there by details from Uzntatherium Letdianum wherever the study of the former is interrupted by imperfect preservation. The U. Letdianum skull will not be here described in detail, as a careful description of it has already been given in a former publication from this museum. This skull was found by Mr. Speir, a member of the party of 1877. The basi-occipital and sphenoidal regions had been exposed to the weather for some time, and were completely broken away; in other respects it is perfect. As figured in Plate II., the outline of the occiput is restored from the posterior half of another skull which we have assigned to Uintatherium. The latter was procured two years later. It happily fills the gap left in the occipital region of the U. Leidianum skull, and admits of a close study of the bones and foramina of the two posterior segments of the skull, including the periotic portion. It is regretted that we have not as yet been able to obtain transverse sections of this specimen to study the cavity of the brain and the petrous portion of the periotic. The L. Speirianum skull, represented in Plate I., was obtained in the summer of 1878. Itis wonderfully preserved, and has been put together with great skill by the Curator of the Princeton museum. Unfortunately there is a wide break at the base, between the palatines and the forward lower margin of the tem- porals. The pterygoid bones, which form a deep backward continuation of the posterior nares, are thus wanting in all our specimens. All the remaining bones of the skull are represented in the different specimens. COMPARISON OF THE SKULLS OF LOXOLOPHODON AND UINTATHERIUM. When placed beside each other, the skulls of the two genera offer a wide contrast even under a rapid glance. Seen from above, the latter is one third longer, while it is considerably narrower at the broadest point. This chiefly arises from the forward extension of the skull in front of the orbit, the distance between the orbit and the occipital condyles being approximately the same in both genera. The broadest portion of both skulls is between the parietal and median protuberances. In Uintatherium the lateral expansion above the orbits overhangs the zygomatic arches, while the Loxolophodon skull is gently rounded at this point and the zygomatic arches project widely. The Uin- tatherium nasals are shorter, and their processes are knobbed and project more upwards. The fossa above the brain-case is deeper in Uiutatherium. But by far the most conspicuous feature of the skull is the huge size of both pairs of protuberances—they are so out of proportion to the skull as to give it a most ''20 £. M. MUSEUM MEMOIRS. grotesque appearance, which must have been heightened, if, according to the questionable theory, they bore horns. The Loxolophodon protuberances are of more modest proportions, but still out of keeping with anything at present found in nature. In side view the Uzntatherium canines are seen to project more backwards, and the premaxillaries are shorter. It is at once noticed that the eye of this genus must have appeared smaller and more sunken from the excessively developed supraorbital ridges. The condyles of Loxolophodon are more closely attached; the occiput overhangs them slightly in both genera. The malar and jugal portions of the zygoma unite by suture in Loxolophodon, by smooth articular facets in the allied genus. Many of these minor characters are undoubtedly due to variations which would not extend through all the species. At the base of the skull we find characters which seem to be more permanent. The premaxillaries send in a very narrow strip to form the anterior portion of the hard palate in Uzntatherium. The foramen incisivum is unenclosed, but towards the extremity of the tips two small processes, partly broken in our specimen, show that in the living state this foramen was partly enclosed. In Loxolophodon broad plates grow well forwards on the inner sides of the pre- maxillaries, extending the hard palate much further forwards. It is extended further back also, in L. cornutus to a point slightly behind the last molar. In U. Leidianum the horizontal palatine and maxillary plates are somewhat shorter, and the posterior nares open opposite the penultimate molar. LOXOLOPHODON SPEIRIANUM, sp. nov. General Features of the Skull. The skull is long and narrow, with a high occiput, surmounted by a slight ridge. The median protuberances project well outwards, and are placed considerably anterior to the orbits. In front of them the nasals narrow gradually into the snout. The zygoma arches well outwards. The upper surface of the skull between the frontal and occipital protuberances is gently rounded upwards—although the upper outline is somewhat exaggerated | in our specimen from lateral crushing. When compared with other species the features just detailed come out very prominently ; in fact, this skull breaks down many of the characters which have hitherto been assigned as distinguishing the whole genus. i Comparison with L. cornutus. The median protuberances in L. Speirianum are smaller, and their forward position relatively shortens the snout, and gives greater distance ‘between them and the posterior protuberances. The total length of the skull is the same in both species. The nasals are not cleft in front, and the knobs, misnamed protuberances, which they bear, project more upwards. The zygomatic arches are closer to the skull in ZL. cornutus, the ''LOXOLOPHODON. AND UINTATHERIUM. 21 post-glenoid processes are longer, the occipital condyles less prominent, canines set wider apart and the canine alveoli more compressed from side to side; the premaxillaries are shorter, and the horizontal plates between them _pro- duced further forwards. The hard palate is produced backwards by the horizontal plate of the palatine bone slightly behind the last molar in L. cornu- tus, while it is opposite the last molar in L. Spetrtzanum. With the exception of the last features, the base of the skull closely resembles that of Z. cornutus. Comparison with L. galeatus—The shape of the occiput principally dis- tinguishes this species from LZ. galeatus. Judging from a figure of the latter which Prof. Cope has kindly lent us, the occiput of Z. galeatus was about thirteen inches broad and twelve inches high, while the occiput of ZL. Speirianum is eleven inches broad by thirteen high. SKULL OF LOXOLOPHODON SPEIRIANUM. Description in detail. (see Plate 1.). The general contour of the head has already been described. The Wasals, which are unusually long, form the upper and all of the anterior portion of the snout, and are produced upwards upon the inside bases of the median protuberances; behind this they pass back beyond the line of the orbits to form a V-shaped suture with the frontals, terminating in a point about half-way between: the median and posterior protuberances. The snout, rounded above, narrows gradually. It is not constricted immediately in front of the protuberances as in L. cornutus, nor does it taper so rapidly as in Uintatherium. At its extremity it is broad and shovel-shaped, and not so deeply notched as in L. cornutus. The Premaxillaries form the downward projecting tips. They are wholly edentulous, with horizontal palatine plates between them produc- ing the hard palate forwards to within an inch of their extremities. This notch was probably, in the living state, still further produced by a membrane which supported a callous pad, opposing the sharp lower incisors. The pre- maxillaries could not be described as “deeply furcate,” as in the case of Z. cornutus. Their lines of junction with the nasals above and maxillaries behind are indicated in the plate. The Maxillaries rise well on the sides of the snout anteriorly, contributing all of the outer and upper moieties of the forward protuberances, and forming a considerable portion of the zygomatic arch. They contain the wide alveoli for powerful canines; these are wanting in our specimen. The alveoli indicate that they were much compressed later- ally. The protuberances are about six inches long, placed upon the sides of the skull, slightly recurved, and with an obliquely placed oval in transverse section. A peculiar feature of the maxillaries is their long suture with the lachrymals and short frontal articulation. Projecting back beneath the malar bones they form the lower anterior third of the zygoma, articulating beneath the orbit with the orbitosphenoids, and on their lower sides forming the greater ''22 £. M. MUSEUM MEMOIRS. extent of the hard palate. The infra-orbital foramina are obscured by fracture in each skull. Cope speaks of them as of small size. The hard palate is very long and narrow. The anterior three inches are formed, as stated above, by the horizontal plates of the premaxillaries. The maxillo-premaxillary suture is marked by the anterior palatine foramina, which are quite small. The palatine groove is found about half-way back on the maxillary plates. Here the palate is arched upwards from side to side with a somewhat decided median ridge. The inward shelf of the palatine bones forms the last inch and a half, notched in front by the posterior palatine foramina, and with a small median projection beneath the opening of the posterior nares behind. This opening is slightly anterior to the last molar. Behind the nasals the Frontals extend a short distance backwards, taking a comparatively small share in the upper walls of the cranium. Their con- nection with the parietals is indistinct. At this point the skull is more rounded than in U. Leidianum, where the upper surface is nearly flat, except for the prominent supra-orbital ridges. A broad swelling on each side, just above the orbits of LZ. Sperrianum, represents the supra-orbital ridges of Uintatherium. At their sides the frontals unite for some distance with the alisphenoids, the line of junction with these bones, as with the lachrymals, being about one-third way down the temporal fossa. The Farietals, uniting broadly with the squamosals and, by a narrow junc- tion, with the alisphenoids, form all the upper posterior portion of the tem- poral fossa. Above, they form narrow marginal ridges, just behind their articulation with the frontals, which arch rapidly upwards and backwards, bounding the deep supra-cerebral fossa between. The marginal ridges rise into the great parietal protuberances, behind which they slope away for a short distance, then rise again to form the occipital crest. The protuberances are somewhat smaller than in U. Leidianum. They are directed outwards, widely expanded at the top, forming in section a fore-and-aft oval at their bases and a transverse oval at their summits. They are admirably represented in the drawing. The occipital crest arches upwards in the centre, thickening into a slightly prominent rim which is most marked at the sides. The Occipitals are comparatively narrow just above the condyles, forming a junction at the sides of the lower third with the mastoid portion of the periotic, if the sutures are correctly interpreted. The occiput, narrow below, rises thirteen inches and spreads to a width of eleven inches in the upper third; it is inclined obliquely backwards over the line of the condyles. The surface is slightly concave from side to side, with a central vertical ridge dividing it. It is quite different from the occiput of L. galeatus, which is much lower and broader. It conveys a remote likeness to a half-opened fan. The ''LOXOLOPHODON AND UINTATHERIUM. 23 rim and central ridge are slightly rugose for muscular attachment. The con- dyles are directed obliquely downwards and set closely to the skull; they are perforated at their bases by the condylar foramina. The Basz-occipitals are bounded at the sides by the mastoid portion of the periotic, and narrow forwards, joining the long narrow Basz-sphenoids. The mastoid portion of the periotic is rough and prominent. The external auditory meatus is directed sharply upwards and inwards. The post-glenoid process of the Sguamosal is very large and projects below the mastoid, while the glenoid cavity is broad and rather shallow. The squamosal portion of the arch springs well up above the level of the orbits. The JJ/a/ar is rather small, forming a sutural connection with the squamosal in Loxolophodon, as distinguished from the smooth facet in Uintatherium. They overlap the maxillaries, but do not quite extend to the orbit. The zygoma, as a whole, is arched well upwards and somewhat outwards. The A/isphenoids are perforated opposite the post- glenoid process by the foramen ovale. Their posterior limits are defined by the foramen lacerum medium. The study of the skull is here interrupted by a wide break which extends as far forwards as the palatines. The Loxolophodon skull in the basi-occipital region is in fair preservation, but the base of another skull belonging to U. mzraéile affords an opportunity for still clearer interpretation of this difficult portion. The two skulls placed beside each other are found to beara close resemblance in this region; the only dis- tinctions which arise are in sizeand proportion. In the lateral walls of the skull, behind the orbit, the skull of U. Letdianum (Plate II.) offers the best opportunity for study. The description which has been given of ZL. Speirianum will accord- ingly be continued and supplemented in greater detail by a study of the two above-mentioned species of Uintatherium, and those portions only will be described which are imperfect or wanting in the Loxrolophodon skull. UINTATHERIUM LEIDIANUM. A full description of the skull of U. Leidianum was given in the Bulletin from this Museum published in 1878. It possesses many characters in common with Loxolophodon, and those which distinguish it have been outlined in the com- parisons given above. They are chiefly in the palatinal region, in the propor- tions of the nasals, and size and relations of the horn-cores. It is not necessary to detail the lesser variations which can be found in the skull. Many of them can be traced in the excellent figure given in Plate II. The topography of the lower side walls is one of the most difficult problems met with in the study of these skulls. This is owing in part to the age of the individuals, which has caused most of the sutures to become obliterated. The U. ''24 £. M, MUSEUM MEMOIRS, Letdianum skull is in fine preservation, but even here some of the most impor- tant points are concealed by age and fracture. Many of the sutures of the bones can, however, be clearly traced, Partial Description of Skull. The Sguamosals form a long suture with the parietals, and the latter have only a short connection with the alisphenoids. The alisphenotds in turn unite for some distance with the frontals, and form also a short connection with the unusually large /achrymals, thus shutting out the orbito-sphenoids from articulation with the frontals. It is difficult to ascertain whether the ordcto-sphenoids are distinct. Their position is judged of by the fora- mina. The line between the ali- and orbito-sphenoids seems to be marked by a wide fissure, the foramen lacerum anterius, and just above this is a groove extending obliquely downwards and backwards across the alisphenoid and ter- minating ina foramen. It is not clear whether this is the Soramen rotundum or not. The exact position of the foramen opticum is difficult to ascertain, owing to a slight displacement of the bones by lateral pressure; it seems to be above and slightly anterior to the sphenoidal fissure. The /achrymals are perforated on the inner side of the orbit by two foramina; on the outer side, just above the maxillary suture, they are marked by a deep pit. UINTATHERIUM MIRABILE. The skull which has, upon conjecture, been referred to Uintatherium (Dino- ceras) mirabile (Marsh) is quite distinct in contour from the corresponding por- tions of the skull of either U. Leidianum or L. galeatus. It is represented by the posterior half, the break having occurred just in front of the posterior protuber- ances and extending vertically downwards. The distinctions arise first from the smaller size; it may be the head of a younger animal, but, as the teeth are wholly wanting, there is no means of ascertaining this. A second and more striking variation is in the shape of the posterior protuberances, which are well pre- served; they are low and tuberous at the summits, barely rising two inches above the marginal ridge, and presenting the usual transverse oval in section. The marginal ridge of the parietals rises high behind them to a point about level with their summits, so they do not form a conspicuous feature of the skull as in U. Leidianum. The backward prolongation of the temporal fossa is also marked. The occiput resembles that of L. Speirianum on a smaller scale, except that it is less narrowed above the condyles; it spreads widely above, and is directed obliquely backwards. The surface is slightly concave from side to side, with a median vertical crest for attachment of the Agamentum nuche, which is less pronounced than in Loxolophodon. Partial Description of Skull—The base of the skull is in beautiful preservation, ''LOXOLOPHODON AND UINTATHERIUM. 25 affording, as remarked above, an admirable opportunity for the study of the bones and foramina of the base. The condyles projecting downwards and slightly back- wards are raised upon a short neck. This is perforated by the condylar foramen. Just above the upper level of the condyles and perforating the rim of the occipital crest is the stylo-mastoid foramen. Immediately in front of the con- dyles, bounded anteriorly by the post-glenoid process and externally by the peri- otic mass, is a deep space. Into this projects the paroccipital process, which does not extend below the level of the basi-occipitals. Marking the postero- internal angle of this space is the foramen-lacerum posterius. On the antero- internal side is the foramen-lacerum medius, and just external to this is the upward-directed external auditory meatus. This has a wide open entrance behind the post-glenoid process, and narrows quickly as it enters the skull. The low paroccipital process is immediately behind this. A deep groove leads forwards on the inner side of the post-glenoid pro- cess, terminating in the foramen ovale. Immediately in front and internal to this is the posterior opening of the short alisphenoid canal. This canal pierces the base of the pterygoids; it is one of the characters common to all the Dino- cerata, and distinguishing the sub-order from the Pantodonta. It transmits the external carotid artery for part of its course. It is one of the characters which the Dinocerata possess in common with the Proboscidia and Perrissodactyla. If the sutures and foramina have been correctly interpreted, the A/isphenoids extend well around on the inner side of the glenoid fossa, forming a short sutural connection with the dasv-occipitals. The limits of the Presphenoids cannot be posi- tively determined ; they are very narrow forwards. The lower portions of the pterygoids are wanting in all our specimens. According to Cope, they form a high backward continuation of the posterior nares. The Exocczpitals, if the suture has been discovered, are found to extend about an inch forwards in front of the condyles. Here is an obscurely marked suture showing their junction with the mastoid portion of the periotic. This is about an inch and a half broad, with a considerable upward extension on the side of the head. Here it is perforated by the stylo-mastoid foramen, which is placed high. This surface is extremely rough for muscular attachments. The éasv- occipttals narrow rapidly forwards and, terminating opposite the post-glenoid processes, are convex from side to side; they form a narrow suture with the dasv- sphenoids, which are obscurely marked off from the presphenoids. ''26 £. M. MUSEUM MEMOIRS. COMPARATIVE MEASUREMENTS OF SKULLS, Lox, Uinta M. M Head, length over all, (Lox. 374 inches), estimated for Uintathevium..cccccccccccccccces QI 76 Plead breadth Of aboveiOtbits 7c. ec dice cess fesse. see. oda. We lalers ots odin ails vires & sLObe* 222 Bony palate: dengthe es. sok eee isn Ss ey on Becare ven S dap csi Sea ce sven cane ee ees 22 Meal UES MCDM ce se oR Ne ae i eh ok Soar 458 35 BIR OING ME CAMUNE Cea cee te ce usa ee legos ee bos col eecas cc Goes .105 4.52 Median protuberances, distance between, at summits.............ceccceccecccccececcces 37 33 Distance between median and posterior protuberances. .........0.ccccccceccccccccccece -45 237 Distance between anterior protuberances and nasal CIPS see ces ce thes oe eile oe eek 35 +25 a ee PEt CE es ik bike 5 bees Coos Lb wee cose ce ecu idol ccc ls 132 30 soistance trom apple of mares to tips of premaxillaries: 2.2.5.6 6.2 6 osede o5ocs eo ig e. 12 -II4 Distance from angle of nares to tips of nasals...........-ccecceeccs Befeiee oe mee sese 18 .162 mewon plouperdnce, Hewitt 110M base 6600... a occ, ol .16 75 Posterior protuberance, height from supra-cerebral fossa, ......:..+sceccccscceecccccces 25 | PocohmiecRcoueteDIOadest DOIN. t5. os ss oe lols ee co es eet ocue oe oe ease ee 28 Occipital crest, suciout trom base of skulls 2. oo. 6 oes s bcck cee ceiceeccis desc 2 eckbeee 31 oe Comparison with Proboscidian Skull. Some of the points which are sug- gested by a comparison of the skull of Loxolophodon with that of the elephant are of considerable value as showing the common origin of the two forms. Others point to a wide divergence and perhaps predominate over those indicat- ing an approximation of structure. These may be placed with the comparisons of the same character which are inserted with the description of the skeleton which follows, before conjecture can be made as to the degree of divergence. The features of agreement are: (1) Both possess an alisphenoid canal; (2) The anterior portion of the zygomatic arch is made up by the maxillaries; (3) The frontals are narrow and do not join the premaxillaries; (4) There are extensive air-cavities in the skull. In the following respects the Dinocerata differ from the Proboscidia: (1) The periotic bones have.a considerable exposure at the side of the skull; (2) There is a condylar foramen; (3) There are post- glenoid processes; (4) There are no postorbital processes; (5) There are no upper incisors. Comparison with Rhinocerus. The skulls of the Dznocerata agree with those of the Rhinocerotide in the following particulars: (1) They have no postor- bital processes; (2) They possess an alisphenoid canal; (3) The nasals are produced forwards. The points of disagreement in the Rhznocerotide are: (1) The maxillaries, as in all ungulates, form a small portion only of the zygo- matic arch; (2) The external auditory meatus has a more posterior position; (3) The mastoid portion of the temporal bone is not exposed. ''LOXOLOPHODON AND UINTATHERIUM. a7 TEETH—UPPER JAW. Dental formula: J 3 C , Pm e . The narrow premaxillaries are eden- tulous in all the known genera of the Dinocerata. Behind the maxillo-premax- illary sutures are the strong recurved canines which form a striking feature of these animals. These are not preserved in either of our skulls, although with one of the lower jaws, figured in Plate III., Fig. 1, the tip of a tusk was found and has been figured in the same plate, Fig. 6. It is lanceolate in outline, with a median ridge worn by friction with the pendent process of the lower jaw. This belongs to Loxolophodon. Cope describes that of LZ. Cornutus as a tusk of compressed form strongly recurved, and with anterior and posterior cutting edges. According to Marsh, the tusk of Utntathertum (Dinoceras) mtrabile has a decidedly lance-shaped extremity. The Museum contains no remains of the canines of Uintatherium. In this genus they were protected by the pendent flange of the lower jaw, but projected widely below the short corresponding processes in Loxolophodon. The molar series are peculiar for their small size in proportion to the skull. UprpER MOLAR-PREMOLAR SERIES. The upper molar-premolar series are poorly preserved in Loxolophodon, pre- senting much-worn crowns and broken edges. They agree in essential charac- ters, as far as can be judged from our specimens, with those of Uintatherium, a description of which (Plate III., Fig. 5) follows. In the table of measurements the difference in size is seen to be in keeping with the greater dimensions of the Loxolophodon head, which is nearly a third larger. Premolars. The first premolar is not preserved in our specimen. The sec- ond and third are subequal and of about the same conformation, implanted by three fangs, one internal and two external. A large basal ridge completely sur- rounds the sub-circular crown. The crown, somewhat trihedral above, is sur- mounted by two transverse grinding edges; of these, the anterior is the lower and crescent shaped, while the posterior is straight. Together they form an irregular V opening outwards. The anterior ridges throughout the series are much the most worn. Molars. The true molars increase in size regularly backwards. The first, worn almost to the basal ridge, is larger than the last premolar, and shows, more- over, an accessory tubercle on the inner side of the posterior ridge. The extremities of the ridges rise into points and their summits are transversely con- cave. In the last molar there are but two fangs, which are long and wide, ''28 £. M. MUSEUM MEMOJRS. extending the whole breadth of the crown. This tooth is much larger and less worn than the preceding, with a strong, slightly serrated posterior basal ridge, which recalls the corresponding ridge in the last lower molar. The crescentic anterior ridge rises into high points at the extremities, giving a tripodal charac- ter to the crown; it projects beyond the line of the straight posterior ridge. The accessory tubercle is large and shows a tendency to become double. In this respect the tooth agrees with that of U. rodustum, but there is no tubercle occu- pying the entrance of the triangular valley between the lobes. As a whole the series diverge rapidly, but the distance between the first premolars and posterior molars is about the same, owing to the great breadth of the latter. The molars are distinguished from the premolars by the accessory tubercle. The entire series present the outward opening V, the posterior ridge in all being straight, the anterior curved forwards. MEASUREMENTS AND RELATIONS OF UPPER TEETH. Lox. Uinta, M. M Distance between canine alveoli, inner margins...... aerate. cscs ue cere Ee Ge oe TL -08 I i 5 cc nd ow se dede bavecdeccscccc oc peice cs +09 -072 MP Pe Ce mIO TAUEVEINE CIASICICE. Fos ei. bo os dec tc scadecccescicece asses: -04 +039 EN ie ee vied creeds vicceescccs oe. -065 -060 Were re erine 16 Tips of premazillarics, ..... o.oo cence ce cceecs ce ceecce.. -070 +09 oie vec iwc es eevee ee ee - 169 -148 NN yk ots dire wc ne ok cance s Oacdeviessecthuc ls ok -074 -062 feet rte Oe HISE PREMIOIAE, os Sa ee eck se vee bocce bloc cecyedus. 025 Bie aeeno aineiperr Ol Urst PLEMOIML |... oko. ccs cee oad seo ncgs ods e cide Ca O19 Morerauccam diameter or FECOnd premolar... cession nc ee ree voce lol oeddecce ccs. See -022 Meee’ Wie tictCrO! SCCONG PremOlats .. ssw icici ewieie cs noel e sek oe oe .023 Bite Anal atmotor Of third Premalat. . 6.5 6. eseGo9 ''Ry vs as x Bs Es '' maf , Aa eer — AXT AX sine TOO, XTX” ie 700." se AVAL — XVI a. - XAT + x IX. Wa 100. VT ae Ve. f= Yy ona. 7a ake LT + . IT 200! te ae ae tere op —f~- 250. e i i “et a e > . Cio ° S ° Go TTT ater TT RRRTTORRRTTTT RTT TOTAL OLITTOTTD TOOPIDTHDI VIII PTOVUD VTA ETD DART IPODS TIT TIVIN Dy a1 a TTT JHE DRAUGHT HORSE The draught horse similarly is a very careful study based upon the skeleton of the draught horse in action. It represents the opposite extreme of the ““Sysonby” stride since three out of the four limbs are on — the ground and the fourth, the left hind leg, is just being raised in the forward step. The head is extended forward as far as possible so as to balance the weight, because the horse is pushing and also leaning his entire weight against the collar so as to assist the muscles as much as possible. [ 20 ] '' MODEL OF *SYSONBY 7 TWO PHASES OF [ 21 HE GALLOP ''THE FOSSIL SERIES The horse from the very earliest geologic times, roughly estimated at two and a half million years, all the period since the birth of the Rocky Mountain system, has been the aristocrat among quadrupeds in point of speed and delicacy and beauty of construction. This statement is borne out by the comparison in the American Museum exhibitions of the little coursing hound, the whippet, and the original four-toed horse, in which the proportions of the different segments of the limbs are seen to be strikingly similar; in fact, the Hohippus probably had a little more speed, indicated in the elongate structure of its hind feet, than the whippet. SKULL OF MODERN HORSE AND MODEL OF FOHIPPUS It is very difficult to realize the multiple structure of the foot and the diminutive size of these very ancient horses until a life-size model of one is placed beside the skull of a modern draught horse, when it is [22 | ''observed that Hohippus and the skull are of about the same length; also that one feature of equine evolution is a continuous increase in size. This principle of continuous increase in size is graphically displayed in the wonderful SERIES EOCENE TO OLIGOCENE representing the first five or six stages in the evolution of the horse, where three principles are at once apparent: first, increase in size; second, increase in length and delicacy of limb; third, elongation of the limb below the knee joint and hock joint; fourth, disappearance of the outer hoofs, and concentration on the median hoof which now begins to rapidly increase in size. SKELETONS OF WHIPPET?T AND OF POHTPPUS These steps are wonderfully displayed in the series of horses begin- ning with Hohippus on the left and ending with Mesohippus on the right, representing a transformation which occupied perhaps a period of eight hundred thousand to one million years, through natural proc- esses of breeding and the increasingly severe competition of these 23 | ''animals with many carnivorous enemies. The Mesohippus is already a superb mechanism, more delicate in its proportions than any modern race horse and probably equal in fleetness for short distances. nia isi llr ates FOUR TOED HORSE , LOWER EOCENE SERIES EOCENE TO OLIGOCENE Remains of hundreds of these animals are found in the beds of old watercourses which traversed the region now politically divided into South Dakota, Nebraska, Colorado, and Montana. The recapture of a complete skeleton from these ancient watercourses and floodplain basins is a very rare event. The rocks have, however, yielded to the persistent search of the very able corps of explorers engaged in the work, most of whom are natives of our Western States. Especially we may mention James W. Gidley of South Dakota, who was in charge of the Whitney explorations for three years, and Barnum Brown of Kansas. Recently Mr. Walter Granger of Vermont has with great success taken up the search in the Rockies for the oldest American horses. These remains are generally found in a very fragmentary con- dition; they have been repaired and set up by Mr. Adam Hermann, head preparator, and his assistants. [ 24 | ''Nature has produced even more distinct breeds than those pro- duced by man, or rather greater extremes of structure and of habit. Thus very early in equine history among the race of Mesohippus the so-called Forest Horse appeared. These horses browsed on shrubs and soft plants rather than grazed, and seeking this kind of food in soft and swampy ground are distinguished by broad, spreading feet with three hoofs, and by short-crowned teeth resembling those of a tapir. These animals lived for hundreds of thousands of years and found their way even into western Asia. REE TOED HORSE UPPER OLIGOCENE TSE THREE TOLD H MIDDLE OLIGOCENE | ee Seema) SERIES EOCENE TO OLIGOCENE In the other extreme is the high speed mechanism of the grazing or DESERT HORSE which has limbs as finely drawn as those of the existing Virginia deer and was undoubtedly an animal capable of very high speed. This type is represented by Neohipparion whitneyi, or “Whitney’s New Hipparion,” the name having been given in honor of the late Mr. William C. Whitney. This is the most perfect skeleton of a fossil horse ever discovered, so perfect in preservation that even the cartilages of the ribs are fossilized and preserved as well as all the [ 25 | | ''delicate vertebre to the very tip of the tail. It was found near the Rosebud Indian Agency by Mr. H. F. Wells of the Whitney expedition sent out by the Museum in 1902, and was one of the finest products of the whole series of explorations conducted under this fund. The skeleton is that of a mare, as indicated by the small size of the tusks. With the mare in the sandy deposit were found the skeletons SKELETON AND RESTORATION OF THE FOREST HORSE HYPORITPPUS [ 26 ] ''of four younger animals, probably colts which had sought refuge from a sand or electric storm or cloudburst with the mare and were killed and buried at the same time. The head is exceptionally large, the teeth are long and highly effective for the grazing habit, while the limbs are excessively light and delicate in proportion. ioe LI Cy SKELETON AND RESTORATION OF THE DESERT HORSE NHEOHIPPARION WHITNEYI 27 | ''Neither of these types, the Forest, the Desert type, or the H ipparion, are known to be directly ancestral to the true modern horse Equus, and one of the gaps still remaining for our exploration is to discover the immediate ancestors of the true horse. It has long been known that wild horses of great variety covered our country long before the period of the Spaniards and probably long before the period of the first appear- ance of man. The natural causes of the extinction of these splendid native races are still unknown. Not improbably these animals were swept away by an epidemic. HQUUS SCOTTI AND EOHIPPUS First and last stages in the Evolution of the Horse in America Up to the time of our exploration only fragments of these native horses had been found, together with a single fragmentary skull. Thus one of the most important discoveries made in the whole twenty-two years of exploration was the finding of remains of a herd of true horses near Rock Creek, Briscoe County, Texas, by James W. Gidley, of the Museum expedition of 1899. The herd consisted of seven skeletons, [28 | ''most of which were nearly complete. Other skeletons have recently been found as a remnant of the same herd. No other such complete single find has ever been made in all the exploration, covering fifty years, of our Western States and Territories. SIDE VIEW OF HOCK JOINT Showing conversion of Lateral Toes into Splints The animal known as Scott’s horse or Equus scotti, represents the last stage in the evolution of the horse of North America just before it became extinct in this country. It is in every respect a horse, although a badly proportioned one, the head being large and the hip girdle short and clumsy. It differs from the domestic horse in the heavy, deep, zebra-like skull, compact body and smaller legs and feet. Like the modern horse it has only a single hoof on the fore and hind feet, while the side toes are represented by the “splints.” [ 29 ] ''THE WILD HORSES, ASSES, ZEBRAS IN THE ZOOLOGICAL PARK The presence of a great Zoological Park in New York, under the direction of the New York Zoological Society, will render possible in future years the completion of the H1isrory oF THE Horse through the exhibition and study of all the wild living types. A complete list of the wild equines now or very recently shown in the Park, and the principal geographic range of each, is as follows: Wiup Horses. PRZEWALSKY Horsss, Hquus przewalskii. DESERT OF GOBI, CENTRAL ASIA. Wixtp ASsEs. Persian Wixtp Ass, Equus hemippus. DESERTS OF §8. PERSIA, AND ARABIA. Krane, Hquus hemionus. N. Asta; TRANS-BAIKAL REGION. ZEBRAS. GREVY ZEBRA, Equus grevyt. ABYSSINIA AND Br. E. AFRICA. GRANT’s ZEBRA, Equus granti. British East AFRICA. CHAPMAN Zesra, Equus burchelli chapmani. CENTRAL SouTH AFRICA. Mountain Zesra, Equus zebra. Carr Couony, 8. AFRICA. The Przewalsky Wild Horse, otherwise known as the Steppe horse, and nearest relative of the domestic horse, is readily distinguished from all modern domesticated breeds by the entire absence of the forelock and by the fact that the mane rises along the neck like a crest exactly as in the zebras and asses, and does not fall over on one side, as in the domestic horse. The large head, rather short and truly horselike [ 30 | ''ears, small and inexpressive eyes, and light buff-colored muzzle are well shown in Fig. 1. The body is uniformly colored, with a dark brown dorsal stripe. Sometimes there are faint horizontal stripings on the legs. Another very distinctive feature, well shown in Fig. 1, is the short, stiff hair on the upper portion of the tail, of buff or dun color, traversed by the vertical stripe. There is a vast difference between the short, smooth and rather handsome coat of these animals in summer and the rough, shaggy coat of the winter, when a long beard appears be- neath the jaws. Fic. 1. HERD OF PRZEWALSKY WILD HORSES IN ZOOLOGICAL PARK The original stallion and mare to the left. The small colt, born June 8, 1912, is of uniform buff color with a woolly coat. Photographed June 20, 1912 These animals were formerly widely spread over Europe, between twenty and twenty-five thousand years ago. During the Ice Age, they were among the favorite subjects of the cave men, who represented them with extraordinary fidelity as to all the features we have men- tioned, on the walls of the caves of the Pyrenees, and of Dorgogne and northwestern Spain. Not one of these drawings shows a forelock, [31 | ''and it is remarkable how those prehistoric artists portrayed the rather dull eyes in con- trast with the fierce expres- sion they gave the eyes of the bison. The general dun or light- brownish color of the Prze- walsky horses conforms to their semi-desert environ- ment, rendering them less : : Hie. 2. THE KIANG, OR WILD ASS conspicuous, like the now OF THIBET extinct quagga of the Zebra From photograph by the Duchess of Bedford. family, whieh formerly made in Woburn Park roamed the open plains south of the Limpopo River in the Transvaal, South Africa. But the closest imitation of the wild horse is in the wild ass (Fig. 2) from the Trans- Baikal of Asia, known as the Kiang, a specimen of which was presented to the Society by His Grace the Duke of Bedford. The light under- color of the belly of the wild horse is also seen in the wild ass of Southern Asia (Fig. 3) Equus hemippus which has a much _ lighter Uniform Isabella, or fawn-color, with dark dorsal stripe, light colored and slender limbs, light under color and dark erect mane. This Przewalsky. Its limbs are animal differs from the Abyssinian ass, the progenitor of the domesticated asses, in the absence of the shoulder stripes. dark. It shows, too, the dark, [ 32 | Fic. 3. THE PERSIAN WILD ASS color scheme than that of the also light instead of being '' whe Ca AAA PML LLG itabel lal peetcheba hahaa ela Fie. 4. THE PRINCE OF THE ZEBRA FAMILY, Equus grevyt From photograph by Sanborn, in the Zoological Park erect mane and black stripe down the back. In fact, this black stripe down the back so well shown also in the back view of the Grevy Zebra (Fig. 5), is the most universal of all the color markings in the family of horses. It is difficult to conjecture what advantage this dark brown or black line brings to the animal. In all the accompanying photographs it appears to shade off into the background. The very brilliant dark-brown stripes of the Grevy zebra, shown in Fig. 4, certainly tend to make the animal very conspicuous as seen in its yard; but from certain points of view, such as that of Fig. 4, where the sunshine glances off the glistening hair, the white and brown stripes on certain regions of the body entirely disappear. Those who strongly believe in the color protection theory truly point out that in certain sur- roundings this most brilliantly marked of all the mammalia almost Be | ''disappears from human vision. I myself have seen a small herd of Grevy Zebras standing under a tree in the Duke of Bedford’s Park, Woburn Abbey, with the sunshine glistening down on them against a light background, become almost invisible. The vanishing effect is only transitory, however, and from other points of view they again be- come conspicuous. The Grevy is readily distinguished as the largest of the zebras. It is characterized by delicate striping, a very long head, and very large, rounded ears, like those of many other forest-loving ani- mals. Its narrow striping contrasts very strongly with the broad and brilliant stripes of the Grant zebra, which, as shown in Fig. 6, so com- pletely surround the body that they unite with a black line extending along the under surface of the belly. Grant’s zebra, like the Grevy, has a very conspicuous set of hori- zontal stripes extending down Hig. 5. THE GREVY ZEBRA, FROM : ABYSSINIA the legs to the hoofs, and is Distinguished by sharply defined and very thus readily distinguished numerous narrow white and dark chocolate from the Chapman zebra in stripes, and by a very heavy dorsal stripe : : which is continued down the center of the tail. Which the lower portion of the leg is quite pale. The Grant Zebra is typical of a very large group entirely distinct from the Grevy and Mountain zebras. It is broadly known as the Burchell group, the type of which was the zebra found and described by the English explorer Burchell north of the Orange River, which roamed north of that stream as the Quagga roamed to the south. In the typical Burchell zebra (EL. burchelli, now believed to be almost extinct) [ 34 | '' Dg Os Ae a 2S Bape @> 4 ales», Mi ae Fie. 6. THE ACCLIMATIZATION OF THE GRANT ZEBRA, Hquus grant Mare, and foal born July 17,1911. The mare shows the black muzzle, dia- mond-shaped pattern of the star on the forehead, black, erect mane, which extends back into the thin dorsal stripe and broad gridiron over the hips. The slender limbs of the zebra colt have nearly the same length as the limbs of the mother, although the body is very much shorter. This enables the colt to keep pace with its mother in escaping the attacks of the lion, the chief enemy of the Grant zebra. From photograph by Sanborn, in the Zoological Park the entire legs are devoid of stripes, so that the zebras of the Burchell group from the Grant zebra on the extreme north of British East Africa to the extinct Quagga of the Cape of Good Hope region, once presented a complete color transition from the universal striping in the North to striping confined to the shoulders and anterior portion of the trunk in the Quagga of the South. This fading out of the stripes, which affords a color transition between these brilliantly marked animals and the apparently monotonous color of the Przewalsky horse, affords strong ground for believing that all the horses were originally [ 35 J '' Fie. 7. THE EXTREMELY RARE MOUNTAIN ZEBRA, Hquus zebra striped. This belief is strengthened by the fact that reversional striping occurs in all the dun colored horses on the face, the limbs, and the shoulders, while the medium back stripe is found in the duns, bays and browns among the horses. The Mountain Zebra (Fig. 7) is the rarest animal in our entire col- lection, because it is now extinct throughout a large part of its former range and is carefully protected by the South African government in its remaining mountain fastnesses. Like the Grant zebra, its color bands are very broad and comparatively few in number, but it possesses a broad gridiron of transverse stripes over the hips, which is only partially developed in the Grant. Other characteristic features are its short head, very long ears, the distinct lap or loose fold in the under [ 36] ''skin of the neck, and the very short, heavily-built limbs which adapt it to its mountain habitat. The call of the Mountain Zebra is between that of the horse and the ass, and usually consists of three short, barking whinnies in quick suc- cession. The note is uttered with great gusto, and the position as- sumed during the call is more like that of a horse than of the ass, which while braying stands quietly with the head up and the ears pricked forward. The disposition of the Mountain Zebra is generally vicious, whereas the Grant zebra is much more docile and capable of domesti- cation. It is interesting to note that although the zebras were well known to the Romans, this true or Mountain Zebra was the first of this group to be described by Linnaeus, as Equus zebra, from the figure in Edward’s “Gleanings of Natural History.” The Grevy zebra on the other hand, occupying the heart of Abyssinia, was the last of this great group to be discovered, not having been made known to science until 1882, when a specimen was presented to President Grevy of the French Republic, in whose honor the new species was named. FINIS [ 37 | '' '' '' '' a EN RY FAI RFI ELD OSBO RN Honorary Curator of t the ‘Denarnicat of Vertebrate - Paleontology, ‘American Museum of Natural History, since 1910, Vertebrate Palzontologist of the United States Geological Survey since 1900; author of the Age of Mammals, Men of the Old Stone Age, The Origin and Evolution of Life, of American Museum _ Bulletins and Memuirs, 1892-1922, and of the United States Geological Survey Monographs, The Titano- theres and The Sauropoda, unpublished With the cooperation of CHartes D. Watcort, Secretary of the Smithsonian - Institution, also of WittiaM Ditter MatrHew, Wiiram Kino Grecory and Marjorie O’Connett, American Museum of Natural History REPRINTED FROM THE TWELFTH EDITION (Eleventh Edition and Three New Volumes) OF THE EN CYCLOPADIA BRITANNICA | - THROUGH THE COURTESY OF THE PUBLISHERS ; FOR LIMITED DISTRIBUTION 1922 '' ''PALAEONTOLOGY Prats I. Middle-Cambrian invertebrate fossils, showing the diversity of the animal life of that period and the similarity of many of the types to recent forms. The specimens from which the photographs were taken are in the U.S. National Museum. (Illustrations reproduced by permission of the Secretary of the Smithsonian Institution, Washington, D.C.) Fic. 1.—Choia carteri Walcott, a silicious sponge. Fic. 2.—Ottoia prolifica Walcott, a gephyrean annelid. Fic. 3.—Ottoia minor Walcott, another gephyrean annelid. Fig. Fic. Kre. 4.—A ysheaia pedunculata Walcott, a Tomopteris-like annelid. 5.—Canadia spinosa Walcott, a polychaete annelid. 6.—Amiskwia sagittiformis Walcott, a chaetognath. 7,—Waptia fieldensis Walcott, a Mysis-like crustacean. 8.—Opabinia regalis Walcott, a Branchipus-like crustacean. 9.—Another specimen of Opabinia regalis Walcott. . 10.—Burgessia bella Walcott, an Apus-like crustacean. 11.—Marrella splendens Walcott, a simple trilobite. 12.—Naraoia compacta Walcott, a curious crustacean. ''PLATE II. Notable vertebrate fossils, complete remains of which have been discovered during the last decade. The mounts from which the photographs were taken are in the American Museum of Natural History, New York. (Illustrations reproduced by permission of the President of the American Museum of Natural History.) Fic. 1.—Deinodon or Gorgosaurus, a mid-Cretaceous carnivorous dinosaur from Alberta, Canada, mounted in running position. Fic. 2, 2a.—Struthiomimus, the “ostrich mimic,’ a mid-Creta- ceous browsing dinosaur, from Alberta, Canada, a toothless offshoot from the carnivorous dinosaur stock. Fig. 2 shows the complete ci PALAEONTOLOGY skeleton in rigor mortis, while fig. 2a represents the same skeleton partly restored from fig. 2. Fic. 3.—Diatryma, a gigantic mollusc-eating bird, from the Lower Eocene of Wyoming. : Fic. 4.—Moropus, an okapi-like herbivore, from the Lower Mio- cene of Dakota, related to the chalicotheres of Europe and Asia. Fic. 5.—Pliohippus, direct one-toed ancestor of the modern horse, from the Lower Pliocene of Nebraska. ; Fic. 6.—Trilophodon, direct descendant of Mastodon angustidens of Europe and North Africa, Lower Pliocene of northern Texas. All the figures are on the same scale. ''PALAEONTOLOGY 9 _roysky, om the other hand, have gone to old Russia/for, their. mo- tives: Stelletsky is the purist of the group, his reconstructions of Russian mediaeval life. being based upon minute archaeological study. of .ikons,, service books. and, similar. sources, Nicolas Roerich has departed from strictly documentary methods in seek- ing to reconstruct, primeval and prehistoric. Russia in his fantas- tic flat decorations based on Russian legends, and thereby joins hands,.with the group-represented, by. Vasnetzoy.. Rather apart is. Boris Anrep (working in1924) in. England), who, studied Byzantine art and the ikon,.not in,an archaeological spirit, but as;exemplifying a means, for, the expression of human emotion, His work is. principally. in. mosaic, submission, to, whose, lim; itations; he holds, makes for the. simplicity, and . directness which are often lost amid the technical possibilities of oil paint, The. close connexion.of modern) Russian art with the theatre is another important. characteristic, which has. grown directly out of the, decorative, reaction against, realism, ..LLeon. Bakst repre- sents one, side.of this... Originally, associated with the Petrograd historical. group, he came into touch with Serge Diaghilev. and became. one of the chief designers,.of settings for the Russian Ballet... His use,of line and colour, relates him to the East; but, like Benois and Somoy, his outlook and method.are those of the West. ..Distinct.in character is the art of Nathali Gontcharova and | M,, Larionoy..Using..the methods, of the Petrograd. group; they,took their material from Russian peasant art, as represented in,the decoration. of articles in. daily use and in the.“ lubok,”’ the Russian. equivalent,of the ‘‘ images d’Epinal,”’, which, gives. their earlier work notable simplicity and directness: , The West was not to.be,denied, however, and Gontcharova’s.setting for, the 1914 production, .of; the ,‘‘\Cog,d’Qr,” ,and) Larionoy’s, “,Les Contes Russes ”” o£, rors. mark.;the invasion .of .the theatre by.cubist ideas, , The, colour scheme,.was. still that of Russian; peasant,art; but the design was based on abstract forms, and aimed at a rhythm-in harmony with the music and the dances.-To this development the name of rayonnisme has been given. : Muchof the criticism levelled at the modern moyement, like that one “directed against impressionism,'is merely a violent statement of personal preference. Weightier arguments’ point: out that-theemphasis given inthe modern movement to the third dimensionomerely exalts one! element:in natural appearance, and urge that ultimately design must be based on the play of contour and_.shapes on the picture plane...Also, itis said, modern methods of simplification: and, distortion tend, to become formulas, which prevent; sincere. and: spontaneous .expression no,less than older conyentions. . But contention chiefly centres round the question ef representation. Itis argued,that,a purely abstract art, which takes no.account of the ideas and emotions conveyed: by the ob- | jects represented, is a limited and empty affair. Rhythm-in the plastic arts, no less than in literature, must ‘emphasize some meaning; and form takes on, a.significance by association, if not with specific objects, yet with general ideas of, mass, space and movements foscta: cr seccls 5-8 a See also:,..Maurice Denis, Theories1890-1910..(1912); W., H.. Wright, Modern Painting (1916); R. Fry; Vision and Design (1920) ; A. Salmon, L£’Art Vivant’ (1920); G.* Coquiot, Les ‘Indépendants | (1920); P) Westheim, Die Welt als Vorstellung (1918); Fritz Birger) Cézanne und Hodler (1913).; Ambroise Vollard; Paul Cézanne; (1914)5 Charles Morice;,Paul Gauguin ,(1919);, Vincent .van.Gogh, , Lettres a Emile Bernard (1912); Kandinsky, Phe Art of Spiritual Harmony; A. Gleizes, Du Cubisme (1920). CW CP Unirep Srates.—Between. 1910 and, 1921 many of the paint- ers mentioned inthe earlier article (20.518), had passed away, and some of their younger contemporaries had also laid down their brushes: Ryder,.; Bunce, Blakelock, Duveneck,. Alexander, Smedley, Millet, Cox, Beckwith, Alden Weir. Abbey, who. died in.1911, left no followers, but La Farge and Chase wielded great influence over.ahost.of pupils... With the development of Ameri- can art-schools and the increasing number of capable instructors, the trend towards European art-centres had by 1921 grown less,. There was already, promise of a school. with, distinctly American characteristics. ‘This:was to beseen most clearly among the paint- ers of landscape. Twachtman and Robinson, among the older men who were trained abroad, brought back some of the light of the so-called. Impressionist. School, and. their example in: raising, the colour-pitch was of great benefit. Crane, a pupil of Wyant, and such'men as Tryon, Murphy and Ben Foster, ably carried on the tradition they received from their American masters. Dew- ing, Metcalf and Childe Hassam, developed individual ways of looking at their subjects. Carlsen, Dougherty and Waugh found the sea an ever-changing theme for their brushes, and they pro- duced canyases not behind those of the landscape men. noe” With the passing. of the Society of American. Artists, the men who made'this organization a force were merged with the mem- bers of the older National Academy and became conservatives in theit turn. Thayer, Brush, Blashfield, Tarbell, Mowbray, Melchers .and, Simmons. were still. in 1920 painting pictures which showed their sound technical training and their artistic point of view: Some of the later men who developed original ways of doing things were Robert Henri, Jonas Lie, William Glackens,' Rockwell. Kent, John.Sloane, George B. Luks, C.\C. Cooper; A. B, Davies; Jerome Myers, George. Bellows, Gardner Symons, Everett. Shinn,,W. E. Schofield and Randall Davey.., Abundant manifestations of vorticism.and cubism came to be seen, in. American painting... The followers of Cézanne, Matisse, Gauguin, Van, Gogh, and, Picasso..were-many, but. chiefly the younger, men. whose. work. was. still, in the experimental stage. | There, was.a steady. advance.in mural painting. Sargent added to the, decorations: forthe Boston, Public Library, and the exam- ple set. there.and. elsewhere was followed in many, of, the larger, cities, in state capitols, municipal, courts, churches and. theatres. | Pittsburgh, .Harrisburg, Baltimore, St.Paul. and Minneapolis | have important-buildings, decorated by; such mural. painters. as La.Farge;, Blashfield, Alexander,-and.others.,.There has been remarkable, growth:at.the art museums, especially at, the Metro- politan Museum of New York, the Boston Museum.of Fine Arts; the Carnegie Institute of Pittsburgh (whose internationgl exhi- bitions draw many exhibits from overseas); and the Chicago Art Institute...No less remarkable has been the formation of impor- tant-collections in cities whose size would often afford no reason for expecting their. presence. Worcester, Providence, Cleveland ‘and Minneapolis have, excellent museums. .Washington now possesses three collections. of paintings—the Corcoran. Gallery, the National Gallery and the Freer Collection:: Moreover, private collections of importance have increased in number and quality,.and native artists are often given there.the,.high place they.deserve.,.Some of the universities offer courses.in the His- | tory, ofArt.and in, the elements of design, In, time. this should | produce.a, body of intelligent criticism which should stall further stimulate, artistic. effort in: America... (Tn Go VAB Ide). PALAEONTOLOGY. (sce20.579).-During the period 1910-25 | the science of) extinct forms of, life made rematkable ;progress, _ especially in North America, where explorations.and studies were less interrupted by the World War... The contact of palaeontology _ with: other, sciences—even those-apparently remote, like, as- tronomy, physics and chemistry, less| remote like comparative | anatomy, orvery intimate like geology-——was; one of the out- standing features of the synthetic work accomplished. . Of tran- scendent interest, however, was the contact between mammalian palaeontology..and anthropology, especially. through the, re- searches: of; William’ K., Gregory -of the American) Museum of Natural History, and also of, G.Elliot Smith, of London Univer- sity: to whom is due the article,on ANTHROPOLOGY in. these New. Volumes. Principal Synthetic Works of toro-21.-Chief among’ the syntheti¢ works in’ pure palaeontology are those of the Austrian palaeon= tologist Othenio: Abel, Grundziige.der. Palaeobiologie der Wurbeltiene (1912), Die, Stimme der. Wirbeltiere (1919), and Lehrbuch der, Pa- leozoologie (1920), which give masterly reviews of the whole fossil history of the vertebrates, especially in analogous and’ convergent adaptation: In invertebrate palaeontology the readeriis referred to Amadeus. Grabau’s. Principles of Stratigraphy (1913) and Textbook of Geology (1920-1),,in which are summed, up the principles derived | from the teachings of Waagen and Neumayr,in Germany, of Hyatt and Beecher in’ America, in ‘pure’ palaeontology’ and in application to'geology.’ A broad syntheticitreatment of climate and time in rela+ tion tothe evolution,of life isthat of the late Joseph Barrell, (1917) in his Rhythms and, the Measurements, of Geologic Time. The best ''TO synthetic treatment of climate, time and geologic change in relation to the geologic origin and the migration of the différent .vertebrate groups is William Diller Matthew’s,.Climate and Evolution (1915). The subdivisions of geologic time and the successions of faunas and climates are broadly reviewed in the Textbook of Geology by Louis V. Pirsson and Charles’ Schuchert (1915; revised edition, vol. 1; 1921). ‘The latest summary of the geology, past physiography and palaeontology of the world is found in the French edition of the great work of Eduard Suess, Das A nitlitz der Erde, translated and annotated by Emmanuel de Margerie as La Face de la Terre (1902, 1918). The comparative evolution of the mammalia of the eastern hemi- sphere and of North America is broadly treated in Henry Fairfield Osborn’s Age of Mammals (1910); while the mammals of North and South America are compared in W, B. Scott’s History of Land Mammals in the Western Hemisphere (1913). A broad treatment of the whole subject of invertebrate and vertebrate evolution is given in Richard S. Lull’s Organic’ Evolution (1917) and a synthetic review of the earth’s history, from, its solar beginnings to the Age of Man, in Osborn’s Origin and Evolution of Life, Life Epochs of Geologic Time.—The time scale in the accom- panying table is taken from the work of Pirsson and Schuchert of 1915, modified by the substitution of ‘geologic time units for years. There is a growing indisposition to reckon’ past time in terms of years, and a growing tendency to substitute a relative term like time units, because of the enormously wide discrepancy between the older estimates of geologists, based’on sedimentation and the thickness of the various assemblages of rocks, which, taken together, make up the whole geologic time scale, and the estimates of physicists, based on the slow liberation’ of radium from radioactive minerals. The radium estimates of the age of the earth range as high as 1,400,000,000 years for the oldest known rocks, according to Barrell, who has adopted the calcula- tions of Rutherford and others based on the “ rate of disintegra- tion ” of radioactive minerals. The contrast between the two methods is exemplified in the following table: PALAEONTOLOGY by palaeontologists. In the same discussion W. J. Sollas com- ments on the expansion of time estimates proposed by physicists: “The age of the earth was thus increased from a mere score of millions to a thousand millions and more, and the geologist who had before been bankrupt in time ‘now found himself suddenly transformed into a capitalist with more millions in the bank than he knew how to dispose of.” In this connexion we may recall the fact that as early as 18 590 Charles Darwin pointed out that the high degree of evolution and specialization seen in the invertebrate fossils at the base of the Palaeozoic, namely, the Cambrian, proved that Precambrian evolution occupied a period as long as, or even longer than, that of Cambrian to Recent time (see Table I on p. 11): Poulton, the leading disciple of Darwin in England (1896); declared that 400,000,000 years was none too long for the whole life ‘evolution period; this would allow 200,000,000 years for Precambrian time and another 200,000,000 years from Cambrian to Recent time, Walcott’s Revelation of Precambrian and Cambrian Life— Charles D. Walcott (1899, 1914) has discovered the remains of life in the Precambrian (Proterozoic) rocks of North America and has been able to give us a fragmentary picture of the fauna and flora of that very ancient period. In Montana at a depth of nearly 10,000 ft. below the earliest Palaeozoic rocks (Cambrian) he found evidence of ancient reef deposits of calcareous algae, which ranged upward through 2,000 ft. of strata.‘ Above these reefs are 3,000 ft. of shales containing worm trails and the frag- mentary remains of large crustacean-like organisms. From rocks of approximately the same age in Ontario, Canada, he has de- scribed sponge-like forms (Atikokania) which are of such general- ized structure that it is difficult to decide whether they should bé regarded as sponges or as archaic corals. These few plant and animal remains are all that are known from remotely metamor- ~ Walcott (1893) Years Barrell (1917) Years Age of Man and of Mammals—Cenozoic WWSe.Or RepiilecsWlesozZ0iG nF anpauades stadt - Age of Amphibians, Fishes, Invertebrates—Palaeozoic Minimum Total f 3 J J a ® Maximum Total Precambrian Time—Evolution of Invertebrates and of Unicellular Life’ 3,000,000 55,000,000— 65,000,000 9,000,000 140,000,000—180,000,000 18,000,000 360,000,000-540,000,000 30,000,000 600,000,000—800,000,000 1,200,000,000 1,400,000,000 90,000,000 (Geikie, 1899) 400,000,000 (Geikie, 1899) ~The most original part of Barrell’s contribution was the measurement of time from the base of the Palaeozoic to Recent time by new palaeophysiographic methods, taking into account particularly the rhythms or cycles of dry and moist’¢limates and of elevations and depressions, theories which were’ originally interpreted by T. C. Chamberlin and popularly treated by Ellsworth Huntington, the physiographer of Yale University. A few decades ago the physicists and mathematicians, ’es- pecially Kelvin and Tait; insisted that the earth could not be more than 10,000,000 to 20,000,000 years old; now the physicists are extending the age of the life period to 1,400,000,000 years, as estimated by Barrell (1917). The most recent determination by physicists, as reviewed by Lord Rayleigh (1921), takes into consideration the transmutation of chemical ‘elements, for example, in the bréggerite of the Precambrian rocks at Moss, Norway: “ Taking the lead as all produced by uranium at the rate above given, we get an age of 925 million years. Some minerals from other archaean rocks in Norway give ‘a rather longer age. . . The helium method is applicable in some cases to materials found in the younger formations, and proves that the ages even of these are to be reckoned in millions of years. Thus the helium in an Eocene iron ore indicated. 30,000,000 years at least. . . The upshot is that radioactive methods of research indicate a moderate multiple of 1,000 million years as the dura- tion of the earth’s crust as suitable for the habitation of living beings, and that no other considerations from the side of pure physics or astronomy afford any definite presumption against this estimate.” Applying this estimate to the evolution of a familiar mammal like the horse, it might be said that the four- toed horse (Eohippus) existed 30,000,000 years ago; a somewhat larger estimate of the life period of the horse than that demanded phosed rocks of Precambrian time, but the existence of annelids and possible arthropods marks a break into the hitherto unknown Precambrian. Walcott’s most surprising discovery in Precam- brian time is a monad or bacterium attributed to Micrococcus sp. indet. from the Algonkian of Montana, but probably related rather to the existing Nitrosomonas, one of the prototrophic or primitive-feeding bacteria, which derives its nitrogen from ammonium salts. In 1910 Walcott discovered in the Cambrian (Burgess) shales of Alberta, Canada, a marvellously rich fauna whose preservation is so perfect that the setae of the worms, the jointed appendages of the trilobites, the impressions of soft-bodied medusae and holothurians, and even the alimentary tract and stomach of certain of the crustaceans can be seen on the shale surfaces almost as clearly as in living forms (Plate I.).\ This discovery fairly revolutionizes our knowledge of the anatomy of the delicately organized as well as the chitinous-armoured forms, like the trilobites. Including the new forms contained within these Albertan shales, the Cambrian marine fauna is now known to be far more abundant than even imagined by Darwin, comprising some 1,500 species, 1,200 of which occur in North America. From Lower as well as Middle Cambrian (Burgess) faunas, it appears that the Precambrian invertebrates had probably be- come completely adapted to all the life zones of the continental and oceanic waters, excepting possibly the abyssal. All the principal phyla—the jointed arthropoda (including the trilobites among the crustaceans and the merostomes among the arachnids), segmented worms (Annelida), echinoderms, molluscs (including pelecypods, gastropods and primitive cephalopods), brachiopods, medusae and other coelenterates, and sponges—were presumably established in Precambrian times. ''a PALAEONTOLOGY TABLE I.—PROGRESS IN PALAEONTOLOGY. - MILLIONS ge AGE OF MAN e QUATERNARY _ — 35 eae S TERTIARY Time 45 Se OF = zZ wee MAMMALS) | & UNITS .j a ‘ee ‘e tee lw UPPER 545 : Ig O CRETACEOUS Wy 5 at O LOWER fr ulo AGE N CRETACEOUS bods oO OF fe) (COMANCHEAN) > : a 0 . en ~) JURASSIC tig u 104 2 bu Ss Ww 2 a TRIASSIC zru/ Oqz JWQw PERMIAN no 405 3 | PENNSYLVAN- a IAN 18> wae aap & (UPPER ; ] 0G 0” AMPHIBIANS = CARBONIFEROUS 4 tow E MISSISSIPPIAN rox zZ ali (LOWER \ ae 3) O CARBONIFEROUS J=Z2N Wl Oj,e it AGE S | DEVONIAN BOr Bi n 2 OF N 8 45 8 K FISHES v Ss JZ bP oO. a SILURIAN Sriai.o oe 17> wi =| jE 3/8 a 2a. O Oo AGE 2 | ORDOVICIAN ea OF x JOON WZ 128 INVERTE- @. bwiot! BRATES z x J 9 2 a | CAMBRIAN 4 x 2 30 MILLIONS 4 ‘a KEWEENAWAN OF dank ° fF rape L Nz Ae O41 oe O82 UNITS Lie i — [YS] ANIMIKIAN £0 O jz ‘354200 Evoctution |N Jes Og 4) OF O}S |HURONIAN | O78 sd INVERTE- | ; a o BRATES [yj] ' oO 4 Pyare st - S | ALGOMIAN Oot] w O Ss ] ae > Y i oO So 404 yZu us o & 7a. 3 }/SUDBURIAN oul O = J="~ufo Jousye wa} 0 4 We w Oo = Ons] O Ze abt Siok O q{ o 1 r2? (09) WW LAURENTIAN {O4uT: I Zhi . 4 a>? Zz O Feu) ¢ aw aWq0}—- < > ere | We 2S 504 0 zW) Mm] EVOLUTION | $309 5 | UNICELLULAR 0 J05 LIFE Jqug| < Oo wir 0 N | Bro ? O dept a WW GRENVILLE S5-l ont . L (KEEWATIN) | pest (COUTCHICHING) 4 a jxuz O JOS OG Y i xe 4 n 60}, TABLE |: Life Epochs: and Geologic Time Units of Europe and North rayeries (After Pirsson and Schuchert, 1915; Issued by Osborn in [91 II The, Cambrian fauna hasbeen. made known to us in large measure through the field discoveries and monographic studies of Philip Lake (1906) for Great Britain, of Walcott (1909-21) for North America, and of Cowper Reed (1915) for India, The great variety and high specialization of the Cambrian marine forms, including representatives of all the known marine in- vertebrate phyla, is in harmony with the trend of discovery among the vertebrates, which is to put the origin of existing families, very far back into the Age of Mammals and even into the Age of Reptiles (Mesozoic). In fact, the antiquity and per- sistence of modern types, as distinguished from modern genera and species, is an illustration of a very far-reaching principle, namely, that the most stable form of energy im matter known 1s that of the heredity chromatin on which this extraordinary pres- ervation of the main features of the ancestral type depends. Next to the stability of the properties of the chemical.elements, which are now known to pass into,each other by transmutation, the most stable physicochemical properties are those which form the heredity basis of life. Freshwater and Terrestrial Origins —The eurypterids appear as contemporaries.of the Cambrian trilobites and traces of them are found ‘in: Precambrian rocks; they attain to their ‘acme in Silurian time and develop into the eight-foot giants of the fauna of the Devonian of Scotland and eastern North America, suffering extinction at the close of the Palaeozoic. In 1916 appeared Marjorie. O’Connell’s: memoir, entitled: The Habitat of . the Eurypterida, giving as the summation of her studies that through- out their entire phylogenetic history the eurypterids lived in the rivers,.a.conclusion, accepted in the main by Schuchert (1916), with the modification that they also appeared to have lived at times in the brackish waters of more’or less large’ bays and possi- bly in limited numbers even in the seas. Many other origins formerly traced:to:the sea. have more recently been traced to fresh water. ‘T..C. Chamberlin (1900) proposed the hypothesis of a prevailing freshwater origin both for the ancestral backboned animals known as chordates as well as for the much more ancient arthropods, the eurypterids, His strong influence was needed.to overcome the widespread notion that all forms of life originated in the sea; and, one after another, theories of freshwater and terrestrial origin have replaced the theory of marine origin. Early in 1916 Barrell pointed out the influence, of Silurian- Devonian climates on the rise of air-breathing vertebrates and freshwater origin in Devonian time under seasonal rainfall. Schuchert continues that the probable freshwater life of the eurypterids opens a vista into continental life as far back as the Upper Cambrian. Other, merostomes related to the eurypterids radiated out from the fluviatile faunas of Cambrian, Ordovician and Silurian time, while in the Devonian rivers ‘dwelt great spider-like eutypterids together with forms so. similar to. scor- pions that, they. might be called river scorpions, and others that were active swimmers. O’Connell’s argument regarding the freshwater eurypterids applies equally to Limulus, the horse- shoe crab... In brief, the existence of freshwater faunas no less varied than the marine faunas is beginning to be traced back to Lower Cambrian time. O’Connell shows that the entire phylogeny of the eurypterids, which includes about 160 species from the Precambrian to the end. of the Palaeozoic, dis- tributed in 78 geologic horizons throughout the world, points to migrations like those of fishes from the headwaters of inter- lacing river systems, and, taken with other evidence, strongly supports the theory of Predevonian river life as opposed to the general assumption.of marine. life, of, all early faunas. It now appears that beginning in Precambrian time the trilobites, by wide adaptive radiation, reached the acme of their development in the Cambrian, displaying a high degree of articu- lation and specialization of appendages, suffered a marked decline after the Silurian, and became extinct at the end of the Palaeozoic. James Perrin Smith, who has made’ avery ex- haustive analysis of cephalopod’ evolution and especially of the Triassic ammonites, observes that the evolution of form con- tinues uninterruptedly éven where there is no evidence whatever of environmental change. ''{2 Principal Literature, Cambrian to Pleistocene:—A few of the:major contributions to, our: knowledge of, the life of thé. Palaeozoic are: Cambrian Geology and Palaeontology,(1910) and,Cambrian Brachiop- oda. (1912), by Charles D,,. Walcott;, Cambrian. Fossils of Spit (1915) and other papers on the, Palaeozoic of India by Cowper Reed; A Monograph of British Cambrian Trilobites (1906) by Philip Lake, and =A: Monograph of British Graptolites (1901); by: Gettrude Elles and Ethel Wood....The foraminifera have been, treated by E. Schell- wien, Monographie der. Fusulinen. (1908-12); the bryozoa by R. S. Bassler, Early Paleozoic Bryozoa of the Baltic Provinces (1911) and G.'W. Lee, British Carboniferous Trepostomata (1912); the echino- derms by Rs T. Jackson in his memoir on the: Phylogeny. of the Echint with a, Revision of Palaeogoic,Species. (1912); and: by. Frank Springer in his monograph Crinoidea’ Flexibilia (1920) and in numerous shorter contributions. The ancient arthropods, including, besides the trilobites, merostomés and other arachnids’ and also insects, have been described: by J. M. Clarke and R. Ruedemann in their memoir on. The Eurypterida of New York.(1912), by Alexander Petrunkevitch, A Monograph of the Terrestrial Palaeozoic Arachnida of North America (1913), by R. I. Pocock, A Monograph of the Ter- restrial Carboniferous Arachnida of Great Britain (1911), and by F. Meunier, Nouvelles recherches sur quelques insectes du\térrain houiller de Commentry (Allier) (1906-12). The literature: on the ; Mesozoic contains more references. to ammonites than to.other groups, ,be- cause of their abundance and palaeontological importance. ‘The ammonite faunas of the Triassic have been ‘described by James P, Smith, The Middle Triassic: Marine Invertebrate, Faunas of North America’ (1914) and by Carl Diener, The) Trias of the Himalayas (1912), Japanische Triasfaunen (i915), and other papers on the Triassic of the Himalayas and southern Europe (1915). -For the Jurassic there are the classic volumes by 'S. S. Buckman, Yorkshire Type Ammonites (1909-19) ‘continued in the Type Aim- momites' (1920) and the memoir by ©. Burckhardt, Faunes. Juras- siques et. Cretaciques de San, Pedro del Gallo (1912) for Mexico, » The studies on Cretaceous ammonites have been more local in character and include: E. Stolley’s Bettrdge zur Kenntniss der Cephalopoden der norddeutschen unteren Kreide (1911-2), D. N2 Sokolov’s Zur Am- moniten Fauna des. Petschoraschen, Jura, (Russian) (1912), H.-Yabe and S. Shimizu’s, Notes on Some Cretaceous Ammonites from Japan and California (1921), and numerous papers. by A. de Grossouvre, W. Kilian and E. Haug for France and the Mediterranean region. The silicious sponges which are so well represented in the’ Mesozoic have received the most careful microscopic study by the studerits and followers of Zittel.. Pioneer work was done in England; by the late George Jennings Hinde, A Monograph of the. British Fossil Sponges (1887-1912), and this work was followed in Germany by A. Schrammen’s’ Kieselspongien der oberen Kreide von \Nordwest- deutschland (1910) and R. Kolb’s Die Kieselspongien des schwibischen weissen Jura (1911). Special works on other groups are: A ,Mono- graph of the Cretaceous Lamellibranchia of England (1899-1912) by Henry Woods, Synopsis des Spirobranches (Brachiopodes) Surassiques Celto-Souabes (1915-9) by the Swiss palaeontologist Louis Rollier, and Clarke and Twitchell’s The Mesozoic and Cenozoic Echinodermata of the. United States (1915). Among the major con- tributions to Mesozoic stratigraphy and entire faunas or. floras may be mentioned: Victor Uhlig’s The Fauna of the Spitt Shales (1903), Carl Renz’'s Die mesozoischen Faunen Griechenlands (1911), G. R. Wieland’s ‘American. Fossil Cycads | (1906-16), and E; W. Berry’s | The Upper Cretaceous Floras. of the World (1916). For the Tertiary life especial reference should be made to. the con- tributions on different groups made by Thomas Wayland Vaughan (corals), E. W. Berry (plants), J. A. Cushman (foraminifera), R. T. Jackson (echinoderms), Mary Rathbun (crustaceans), A. Pilsbry. (cirripedia),, and others in Contributions to the Geology and | Paleontology of the Canal Zone, Panama, and Geologically Related | Areas in Central America and the West Indies (1919). The bryozoa | have been carefully described and beautifully illustrated by Ferdi- | nand Canu and Ray S. Bassler, North American Early. Tertiary | Bryozoa (1920), while the foraminifera have been described in equal | detail by Joseph A. Cushman in numerous contributions, and by H. | Yabe (1921) and H. Douvillé (1911). For other groups we may note: J. Lambert’s Description des Echinides des terrains néogénes du bassin du Rhone (1911-6), F. W. Harmer’s The Pliocene Mollusca (1914-20), and papers by W, H. Dall on the mollusca, .A-general résumé of the | Pleistocene vertebrate and invertebrate Jife is embodied in F. C. Baker’s The Life of the Pleistocene or Glacial Period (1920). Stimulat- ing general reviews of the progress of invertebrate palaeontology are the presidential addresses by F. A. Bather, Fossils and Life, British, Association (1920), by Ruedemann, The Palaeontology of 'Ar- rested Evolution (i916), and by Clarke, The Philosophy. of Geology and the Order of the State (1917). PROGRESS IN VERTEBRATE. PALAEONTOLOGY Personnel: Advent of the Fourth Generation——The principal feature of the decade has been the advent of a new generation of explorers and workers in vertebrate palaeontology who, ina sense, constitute a fourth or ‘‘ 20th century”? group. Beginning | of Stuttgart. with Cuvier (1769-1832) as founder of. the science and leader | PALAEONTOLOGY of the first grotip, thé sécond group etibraced thé British anato- mists “Richard 'Owen (1804+1892) and. Thomas Henry, Huxley (1825-1895), the French leader Albert Gaudry (1827-1908), the Swiss palaeontologist Ludwig Riitimeyer (1825-1895), and the three~-great~Americans, namely,-Joseph Leidy (18234801), Edward. Drinker Cope (1840-1897), and Qthniel. C. Marsh (1834=1800). “These men marshalled the first positive-proofs of vertebrate evolution in Europe and America; they ‘worked more or less independently as pioneers and laid the entire foundation of the: modern classification of the Vertebrata. The‘léader of the third group was the Russian; Waldemar Kovalevsky (1842+ 1883), who instituted intensive investigation of mechanical adap- tation in relation to natural selection. Still productive: members of the same* period are Arthur Smith Woodward. (b. 1864) and Charles W. Andrews (b. 1866) in England, Marcellin Boule (b. 186r).and, Charles Depéret (b. 1854) in France; Louis Dollo (b..1857) in Belgium, Max Schlosser in Germany, Giovanni Ca- pellini (1833—') in Italy, and in America William B. Scott (b. 1858) and Henry: Fairfield Osborn (b. 1857). This group‘includes also Samuel Wendell. Williston recently deceased (1852-1918), and Ramsay H, Traquair (1840-1912). Scott treated chiefly mammals, Williston chiefly reptiles and amphibians, Osborn both mammals and.reptiles..The principal accomplishment of this’third school has been (1), to conduct world-wide exploration, (2) to'correct, co- ordinate and firmly establish the great classifications proposed by the second school and (3) to fill out the details and-principles of phylogeny ‘or lines of reptilian, avian and mammalian descent. The leading explorer of this period was John Bell’ Hatcher (1861- 1904), who brought together a large part of the materials for two great monographs of the United States Geological; Survey, Os- born’s Titanotheres and the Hatchér-Bull Ceratopsia; he“also made the wonderful collection of South American fossils'which forms the basis of Scott’s monumental mémoirs'of the Princeton. University Expeditions to Patagonia during the years 1896-9. Osborn’s monograph: Lhe Titanotheres (an Eocene-Oligocene family of mammals), twenty-one years in preparation, has been completed but not published; others of his memoirs are the Egquidae of the Oligocene; Miocene, and Pliocene of North America (1918) and Camarasaurus, Amphicoelias, and other Sauropods of Cope (1921). Williston’s monographs are chiefly on the Cretaceous mosasaurs and the archaic Reptilia of the Perm-Trias, to which he made-most-notable contributions. Of this period were Flor- entino Ameghino (1854-1911), the distifiguished vertebrate. palaeontologist of Argentina, atid Eberhard Fraas, (1862-1915) { Oliver P. Hay (b. 1846) is also of this group, author ofthe monograph of the Fossil Turtles of North America (1908) and of the invaluable Bibliography and Catalogue of the | Fossil Vertebrata of North America (1902). _ To the fourth group of vertebrate palaeontologists belongs the school trained by Professor Osborn in the American Museum of Natural History, of which the senior is William Diller Matthew (b. 1871), Walter Granger (b. 1872), Barnum Brown (b. 1873), William K. Gregory (b. 1876), Richard S. Lull (b..1867), of Yale University, Lawrence M. Lambe (1863-1919), late of the Cana- dian Survey,,-and C. Forster-Cooper, Cambridge University. The chief intensive work of Matthew and Granger-has been on | the American Eocene mammalian faunas and in aiding Osborn to | establish sixteen Eocené-Oligocene life zones of: North America very closely codrdinated with corresponding life zones of west- ern Europe. Brown’s explorations have added greatly to our knowledge of Cretaceous dinosaurs. © Of the same group are the pupils of Williston, of whom the leader is Ermine C. Case (b. 1871), who has contributed treatises on Permian life. At the same time John C. Merriam (b. 1869) has led explorations on the Pacific coast of America and inspired a school of younger workers both in vertebrate and. invertebrate palaeontology. In Great Britain D. M. S. Watson (b. 1886) has taken up the work of Owen and Huxley in primary groups of fishes, amphibians and reptiles; in Austria Othenio Abel, a pupil of Dollo, is the great exponent of vertebrate evolution; in Germany Friedrich von Huene and Ferdinand Broili are leaders in sauropsidan palaeontology, other notable palaeontologists of recent years being Franz Dre- ''| PALAEONTOLOGY ‘primary groups of fishes and thé first steps towards the frame of wermanh, Ernst Strémer (b. 1871) and Otto Jaekel (bi 1863). At ‘Upsala’ in Sweden Carl Wiman has inspired''a ‘rémarkably ‘pro- ‘gressive group of workers, while in Switzerland Hans Georg Stehlin (b. 1870) has continued in the great field of Riitimeyer. ~ For the’ principal contributions by ‘palaéontologists of ‘the third and fourth groups above described) the'reader is referred moirs of the Société’ Paléontologique Suisse. Yyesearches’ of these workers in field’ and Jaboratory that’ the tothe Memoits and Bulletins of the American’ Museum of Natural History, of ‘the university of California, of the Carnegie Institution of Washington, to “the Contributions from the ‘Palaeontological Laboratory (Peabody Museum) of Yale Uni- WVersity, to the Memoirs’ and’ Catalogues of the. British Museum (Natural History), to the Palaeontographica, and to the Me- It is upon the great synthetic volumes referred to earlier are chiefly founded, and that the following generalizations’ of modern vertebrate palaeontology are chiefly due. ORIGINS OF THE GREAT VERTEBRATE STOCK AND Irs BRANCHES Origin of Chordates—No ‘discovery has thus fat léssenied the gap between the modern Protochordates (Amphioxus, tunicates, etc.) and any of the known phyla of invertebrates. Sone of the “eephalaspid ostracoderms have been cited by Patten as favouring the view that the chordates have been derived from certain arthropods, but.such resemblances are ascribed, to, convergence by Dollo and many others. The earliest ostracoderm remnant ‘actually known is a dermal plate of a genus named A siraspis from the Upper Ordovician near Canyon City, Colorado; this represents a new family Astraspidae allied to thé Psammosteidae of the Silurian and Devonian (C. R! Eastman, 1917): These chord- ‘ates, heavily shielded and hence known as 6stracoderms, were dom- inant in the Upper Silurian, radiating into’six families and many genera, abundant in thé Lowér Devonian, diminishing in the ‘Middle Devonian and becoming extinct in the Upper Devonian. Origin of Fishes —The earliest fish remnant actually known is the fin-spined Onchus from the Upper Silurian of Scotland, which appears to représent’ the group of acanthodian sharks, covered with fine quadrate scales like those of ganoids and with 4 skull structure distinctly elasmobranch. The elasmobranchs (shark and ray types) are still the oldest known gnathostomes or true jaw-bearing Vertebrates, constituting (a) one of the four primary gnathostome groups, i.e. jawed groups, the others being (b) the fringe-finned ganoids (Crossopterygii), (c) the ray-finned ganoids and teleosts collectively known as Actinopterygii and (d) thé lungfishés (Dipnoi). ‘Thé fossil ancestors of the fringe- finned ganoids have not yet been discovered; so these animals are theoretically traced to unknown C¢artilaginous fishes of Silurian times. The oldest Crossoptérygian actually known is the Osteolepis macrolepidotus of the Middle Old Red Sandstone of Scotland. There were two principal périods of adaptive radia- tion among the Crossopterygii, the ‘first in Middle and Upper Devonian times, the second in Mesozoic times which produced the family Coelacanthidae, from which may have sprung the existing fishes Polypterus and Calamoichthys as degenerate off- shoots. From the earlier Devonian radiation of the Crossoptery- gians is traced the theoretic origin of the Dipnoi or lungfishes, on the one hand, and of the oldest known amphibians on the other. The Devonian Crossoptérygian skull and fins appear to be “archetypal,” to the lungfish type, on the one hand, and to the amphibian type on the other. Cope’s genius in separating the Actinopterygii is sustained, for there is as yet no fossil evi- dence of the connexion of this group with the Crossopterygii, other than the supposed community of origin in Silurian times. Here the reader should consult the writings of Smith Woodward, Joseph F. Whiteaves, Bashford Dean, William K. Gregory and the synthetic reviews of Osborn (1918) and Lull (7017). Origin of Amphibians and First Tetrapods—In this epoch- making transition from the fringé-finned fish type to the tetrap- odal amphibian and terrestrial type, the prophecies of Huxley, Cope and Baur and other great anatomists of the second and third groups of palaeontologists appear to be fulfilled. ‘The Silurian period marked the parting of the ways among the great 18 the terrestrial amphibians. Not: until the Upper Devonian 6f Pennsylvania do'we find a footprint (Thinopus antiquus Matsh), which may be referred to an amphibian -tetrapodi The. first known actual skeletons occurred in the Coal Measures (Uppér Carboniferous) of Europe and America and represented. four widely radiating groups. The structurdl. gap: separating the earliest tetrapod amphibians and fishes is perhaps the greatést known in the whole range of vertebrate evolution, but:all modern authorities agree that the amphibians weré probably derived from a Silurian or early Devonian type of fringe-finned fish. Even as far back as the Upper Carboniferous and even! in the Lower Carboniferous the Amphibia were adaptively. radiating into several orders and numerous families comprising highly. special- ized forms. During thé Carboniferous we find numerous indé- pendent phyla of eel-like or burrowing,. and: of compressed, swimming, ‘as well as of large-bodied, predatory’ forms... The latter culminate in the gigantic labyrinthodonts of the Triassic. The exact connexion of any of these forms with ‘the: modern ‘Amphibia (urodeles and Anura) is doubtful. The Anura first appear in the Jurassic, and atthe present time they retain! many characters reminiscent of such: Palaeozoic Amphibia) as: the branchiosaurs and the Eryops group. The urodeles ate first known in the genus H ylaeobatrachus of the Lower Cretaceous of Europe. Both groups, especially the Anura, appear to:have gonle through a wide adaptive radiation during the Tertiary! The connexion of the ‘modern caecilians with ‘the ancient types is obscure. The reader is referred especially to the contributions of Williston, Case, Watson, Gregory; Broili, and: the synthetic re- views of Osborn (1918) and Lull (1917) Origin of Reptiles ~The oldest-known reptiles, solid-headed Cotylosauria of Cope, are regarded. as amphibians which had eliminated the aquatic stages in development, the oldest reptile actually known being the genus Hosauravus from the Coal Measures of Ohio. In other words, the cotylosaur reptiles are traceable to solid-headed stegocephalian amphibians, which, in turn, are traceable to solid-headed unknown Crossopterygians of Silurian times. The oldest and most primitivé reptiles (Coty- losauria) occurring in the Upper Carboniferous and Permian; are thus structurally very close to certain contemporary stégocé- phalian amphibians. The first great adaptive radiation of ithe reptiles into the two grand divisions, the solid-headed (Coty- losauria) and the temporal-arched (Pelycosauria), began in the Upper Carboniferous and still more widely diverged in Permian times. As early as the Permian, occurs a mammalian-like series of reptiles which exhibits an extensive adaptive radiation and gives off one branch, the Cynodontia, which, in turn, survivés into Triassic times and clearly approaches the mammalian grade of organization. From the primary temporal-arched also appear ‘the forerunners of the Mesozoic reptiles, the plesiosaurs, ichthyo- saurs, dinosaurs and pterosaurs, widely separated from each other in the Triassic and thus’ having their branches deep down in the Permian and Carboniferous, each grand division giving rise to an adaptive radiation of itsown. These have been traced in detail by such authorities as Andrews, Dollo, Abel, von Huene, Williston and Osborn. Here the reader is referred to the writings of Williston, Hatcher, Osborn, Merriam, Lambe, Lull, and especially during the past decade to those of Charles W. Gilmore of the United States National Museum, Washington, and of Dr. Robert Broom of South Africa, as well as to the synthetic reviews of Osborn (1918) and Lull (z917). The two greatest achievements of the decade are the clearing up of the relationships of the primitive South African terrestrial Reptilia of the Perm-Trias, beginning with the solid-headed types (pareiasaurs) and ending in their highest expression, the mammal-like types known as Cynodonts and -Theriodonts. The field explorations of Robert Broom and the profound compar- ative researches of D. M.S. Watson and of William K. Gregory have given us a clear comprehension of the habits and relation- ships of this first terrestrial radiation group. Williston and Case have covered the same great period in America. ''a4 Origin. of : Birds.—Palacontologists still agree in endorsing ‘Huxley’s opinion that birds are “‘ glorified reptiles.” ‘The origin of birds, according to recent reviewers such as Osborn and Gregory, brings’ us close to the two-temporal-arched | (i.e. Diapsida) reptiles, namely, to the stem which also gave off the dinosaurs, the pterosaurs and the smaller parasuchians (Eu- parkeria): Fossil bird remains are extremely rare. The earliest bird known is the famous Archaeopteryx of the Jurassic of Soln- hofen, Germany. This is largely a bird, excepting in the tail, the simplicity of the feather arrangement and the possession of teeth. According to the four-winged hypothesis of origin ad- vocated by Beebe, we should:some day discover a bird with parachute-like action in both fore and hind limbs. Recent con- tributions of note on this subject are those of Gerhard Heilmann (1913) and of William Beebe (1915), and the synthetic reviews of Osborn (1918) and ‘Lull (1917). Origin of Mammals.—Evidence has been accumulating rapidly in favour of the theory that’the origin.of the mammals should be traced to the more progressive terrestrial mammal-like reptiles (the Cynodontia) of the Permian and Triassic of South Africa and Europe, as described in the studies of Broom, Watson, Haughton, Osborn and Gregory. Structurally related to these Cynodonts are ‘the so-called Protodonts of Osborn, e.g. Droma- therium and Microconodon of the Triassic of North Carolina. But of equal antiquity are the multituberculates, e.g. Plagiaulax and Microlestes, widely spread over Europe and North America. No substantial additions have been made during the decade to our knowledge of this vague period; readers are referred: to the reviews of Osborn (1918) and Lull (1917), also to the recent works of Gregory, The Orders of Mammals (1910) and The Origin and Evolution of the Human Dentition (1921). Origin of Primates and of Man.—Combined palaeontological and anatomical evidence indicates that the source of the Primates is to be looked for among tree-living insectivorous mammals more or less closely similar to the modern tree shrews (Tupatidae) of Africa. This view advanced with ‘ability by Gregory is in general accord with the opinion that during the phase of arboreal life many of the psychic and anatomical characters of the Pri- mates were acquired. It was not until the Lower Eocene of North America and of Europe that there appeared :undisputed Primates of lemuroid affinity, e.g. notharctids and tarsioids in -America, adapids and tarsioids'in Europe. At this time the zoological relation of the two continents was close and it would appear that while the primitive horses were acquiring their cursorial characters on the ground, these primitive lemuroids were acquiring their distinctive characteristics in the trees. Actual ancestors of the existing Tursius of Madagascar have been found in France (Pseudoloris). ‘The attempt of Ameghino to trace the higher Primates to South American types, e.g. Homun- culus, appearing in the Lower Miocene of Patagonia, is not sup- ported, because these animals from the first are the true broad- nosed, 7.e. platyrrhine, type still characteristic of South America. The Old World division of the catarrhines or narrow-nosed true Primates has been traced to the Parapithecus, described by Max Schlosser from the Lower Oligocene of Egypt. | Propliopithecus is possibly ancestral to the true anthropoid apes and thus possibly related to the ancestors of man himself: Darwin’s broad con- clusion that: man was derived from “‘ some ancient member of the anthropomorphous subgroup of Old World Primates ”’ is fully sustained by anatomical evidence, but the precise. lines of descent are still in dispute.. Some hold that'the human line came from Middle Tertiary anthropoid apes allied to, Dryopithecus of France and Sivapithecus of India, while others (including the present writer) regard the Hominidae as a widely distinct family separated especially by its upright walking gait, by the non- divergence of the great toe, arid by the retention of its tool-making thumb. A series of masterly reviews of this whole question has appeared in the American Museum publications from Gregory, whose recent memoir On the. Structure and’ Relations of Noth- arctus, an American Eocene Primate (1920) sums up our present knowledge. of this whole subject. (See also ANTHROPOLOGY.) The Dinosaur Fauna of Alberta, Canada.—The greatest new PALAEONTOLOGY achievement in exploration is the revelation of the dinosaur fauna of Alberta in|the fossil beds extending along the Red Deer river, which were first made known. to science by explorers of the Canadian Geological Survey in 1897, 1898, 1901. The first general review of this wonderful fauna was that of Osborn and Lambe, On Vetebratra of the Mid-Cretaceous of the North West ° Territory (1902), based chiefly on the collections in the Ottawa Museum. The American Museum explorations under Barnum Brown, which extended over ten years, have resulted in the discovery of the entire fauna of the middle portion of Upper Cretaceous time, a complete revelation especially of the dinosaur world as it approached the height of its adaptive radiation into herbivorous and carnivorous, armoured’ and defenceless, swift- moving and slow-moving types, which severally imitate more or less fully the long subsequent adaptive radiation of the mammals. In 1914 the Canadians renewed exploration, so that at. present the Ottawa and Toronto Museums have rich collections, part of which has been described by the late Lawrence M. Lambe, while Osborn, Barnum Brown and W. A. Parks have also made known.a part of this wonderful fauna. Two. of the greatest extremes of adaptation, namely, Deinodon or Gorgosaurus and Struthiomimus, are figured in the accompanying Plate II. In the same plate appear some of the outstanding American dis- coveries of the decade. : New Discoveries AMONG FossiL VERTEBRATES Fossil Fishes-—Dr. A. Smith Woodward’s Fossil Fishes of the English Wealden.and Purbeck (1915-8) is a beautifully illustrated memoir of the most thorough, systematic type, well sustaining the. traditions, set by Traquair and by the author himself in earlier works. The period dealt with affords an interesting cross- section of the stream_of piscine evolution, at a time when many of the old. Mesozoic ganoids were dying out and the teleost fishes were beginning \their remarkable expansion. Other. important systematic memoirs are those by Stolley on the ganoids of the German Muschelkalk (1920),.and by Stensié (1921) on. Triassic fishes, from, Spitzbergen. The latter memoirs. contain, a wealth of material of great morphological interest concerning the early stages in the eyolution of/the skull of the fringe-finned and ray- finned ganoid fishes;.this discussion also throws light on the origin of certain elements in the.skull of higher vertebrates. In this connexion should be mentioned. the brief but highly important paper on. Eusthenopteron by. W. L. Bryant.(1919). This fringe- finned ganoid is of particular interest because the construction. of its. skull/and paired limbs approaches the type which may be expected in'the piscine ancestors of the land-living vertebrates. ‘The arrangement of the elements on.the under side of the skull of this fish raises, morphological questions of wide general interest. Papers by Watson and Day (1916) and by Gregory. (1915, 1920) deal with the. ancestral relations of these fringe-finned ganoids with the land-living vertebrates (tetrapods). ° The swarming fauna of, Devonian arthrodires, ptyctodonts, cladodonts and) other archaic fossil fishes from. the vicinity of Buffalo, N.Y., is ably described by Bryant and. Hussakof in their Catalog of the Fossil Fishes in the. Museum of the Buffalo Society.of Natural: Sciences, (1918). A serious difficulty encoun- tered by all students of recent and fossil fishes is the getting in contact with the vast and scattered literature of the subject. The great Bibliography of Fishes by Bashford Dean and his associates Eastman and. E. W. Gudger (1917). will undoubtedly stimulate research in this field. Fossil, Amphibians.—The outstanding: publications, in. this field are The Coal Measures Amphibia of North America by R.L. Moodie (1916).and a memoir,on The Structure, Evolution. and Origin.of the Amphibia by.D: M.S. Watson (1919). »Moodie’s memoir is a valuable description and compilation of the extensive and varied fauna of swamp-living amphibians of the American Coal. Measures. Watson’s memoir is a brilliant and highly original contribution. to. the classification and phylogeny of the labyrinthodonts.. Much detailed work jon fossil amphibians ap- pears in papers by, von Huene, Broom, Williston, van Hoepen, Haughton and others. ''PALAEONTOLOGY Stem Reptiles.—In the field of the oldest reptiles, those of the Carboniferous and Permian, perhaps the most important con- tributions are those by S. W. Williston and D. M.S. Watson. The former, in his paper on The Phylogeny and Classification of the Reptiles (1917), traces the rise of the common amphibian- reptilian stock through the ‘‘ Protopoda,” which are so far known only from certain footprints of Upper Devonian age. According to Williston, who built on Osborn’s system of 1903-4, the primi- tive reptilian stock early divided into the following series:— Anapsida (Cotylosauria and their specialized descendants, the modern tortoises and turtles). Synapsida (Theromorpha or pelycosaurs, etc.; Therapsida, or mammal-like reptiles, the latter giving rise to the mammals; plesio- eapsidé (reptiles with two temporal arches, such as crocodiles, dinosaurs, rhynchocephalians; this stock gave rise to birds). Parapsida (including the proganosaurs, ichthyosaurs, lizards, mosasaurs, snakes). Watson (1917), in his Sketch Classification of the Pre-Jurassic Letrapod Vertebrates, assigns a high value in classification to the characters of the brain-case. A general and conservative classifi- cation of the early reptiles is given by W. K: Gregory (1920). The most primitive known reptile, Seymouria, from the Permo- Carboniferous of Texas, almost bridges the gap between the Amphibia and the Reptilia. Watson (1919) gives a morphological description of this reptile, accompanied by valuable figures and reconstructions of the skull and skeleton. The habits and environments of the teeming reptilian and amphibian faunas of the Permo-Carboniferous of North America are intensively considered ina memoir by E. C. Case (1919), which also deals with the stratigraphy, climatology and palaeogeog- raphy of the late Palaeozoic. Mammal-like Reptiles.—In no other field of fossil reptiles has the progress of discovery been more satisfactory than in that of the mammal-like reptiles of South Africa, as set forth in numer- ous papers, especially by Watson (1913-4), Haughton (1918), Broom (1913-4), van Hoepen and others. The relationships of these animals with other reptiles and with the mammals have been reviewed by W. K. Gregory (1920-1). Marine Reptiles —These have always been of great interest on account of their secondary adaptations to aquatic life which have been ably discussed by Abel (1912, 1919). One of the outstanding contributions of new material in this field is the British Museum Catalogue of Marine Reptiles of the Oxford Clay by C. W. Andrews (1910-3). The origin and relationships of the plesiosaurs and their allies are treated by von Huene (1921). Dinosaurs.—The Triassic dinosaurs of Europe are of particular interest because some of them tend to connect the very diverse carnivorous and herbivorous saurischian dinosaurs of later ages. Here the leading author is F. von Huene, in a long series of papers and memoirs. Plateosaurus, perhaps the most primitive of these reptiles, has been fully described both by von Huene and by Jaekel (1913-6). Primitive dinosaurs from the summit of the Karroo series in South Africa (Gryponyx, M assospondylus, etc.) are described by Broom and Haughton. During the long ages of the Jurassic the gigantic sauropodous dinosaurs attained their maximum in size and specialization. The leading feature in this field is the description of the strange and monstrous dinosaurs of the Tendaguru fauna of East Africa‘in the collections of the Berlin Museum, by Janensch (1914). One of the most remarkable of the North American sauropods is the genus Camarasaurus, which has been intensively described by Osbornand Mook (1921). Barosaurus, a gigantic relative of Diplodocus, with a tremendously heavy neck, has been described by R. S. Lull (1919). Tyranno- saurus, the greatest carnivorous reptile of all time, and Struthi- omimus, a contemporaneous ostrich-like dinosaur, have been described by Osborn (1912-9). The highly varied and grotesque armoured dinosaurs, namely, the Ceratopsia and related groups, have been the subject of numerous papers by Gilmore, Brown, Lambe and others. Pterosaurs.—The pterosaurs, or flying reptiles, have continued to excite the interest of students of flight, such as Abel (1912), Watson and Hankin (1914), Arthaber (1921). The greatest 15 flying reptile, Pteranodon, is the subject of a memoir by Eaton (1910) of Yale University. MI Chelonians.—An important memoir by O. ‘Jaekel (1913-6) describes the skull, skeleton, carapace and plastron of Triasso- chelys dux from the Upper Triassic of Germany. Although widely differentiated from all other orders this reptile was the most primitive of all known chelonians. Of even greater interest is the Eunotosaurus from the Permian of South Africa which Watson (1914) describes as a veritable “‘ Archichelone.” Fossil Birds.—Diatryma, a gigantic ground bird from the Lower Eocene of Wyoming, has been described by W. D. Matthew and W. Granger (1917) from a nearly complete skeleton, which is a most rare and valuable fossil. This bird, which has no near relatives, was about seven feet high and of massive proportions, with an enormous head and great compressed beak. The wings were vestigial. This high degree of specialization at such an early epoch indicates that the modernized groups of birds were differ- entiated during the latter part of the Age of Reptiles. Monographs on Special Groups of Tertiary Mammals.—The fossil mammals of the basal and Lower Eocene of the western United States are represented in the American Museum of Natural History by collections numbering many thousands of specimens which are being described jointly by Matthew and Granger (1915). Besides describing many new or little known forms these authors also deal with the relationships and mor- phology of the various groups of early placental mammals. In the paper dealing with the edentates and their relatives, the ““palaeanodonts,’”’’ Matthew (1918) advances the view that the modern Pholidota (Pangolins) are an offshoot of the primitive “palaeanodonts ” of the Lower Eocene. Other papers of the same series cover the Creodonts, Insectivores, Primates and Condylarths. Several mid-Tertiary mammalian groups, such as chalicotheres, entelodonts and the diceratheres, have been revised in the publica- tions of the Carnegie Museum, Pittsburgh, by W. J. Holland and by O. A. Peterson. Baluchitherium, perhaps the most gigantic land mammal of all time, has been described by C. Forster-Cooper (1913) froma huge atlas, astragalus, cervical vertebrae and limb bones from the Upper Oligocene deposits of the Bugti Hills of Baluchistan. The evolution of the Sirenia is treated by Abel (1921) and by Depéret (1920); that of the Cetacea by Abel (1919) and by Winge (1918-21). The phylogeny and evolution of the Pro- boscidea are considered in the researches by Schlesinger (1917), Matsumoto (1915) and Osborn (1918-21). The Eocene and Oligocene titanotheres have been dealt with in numerous papers by Osborn in preparation for his monograph on these extinct animals. The revision of the mid-Tertiary Equidae by Osborn (1918) affords an exceptionally full document on the exact course of evolution in the multitudinous phyla of a typical mammalian family. A most valuable expansion of our knowledge of the anthropoid apes of the mid-Tertiary is found in the work of Pilgrim (1915) on the fossil apes of India of the genera Dryopi- thecus and Sivapithecus. South American Fossil Mammals.—The strange offshoots of the ungulate and edentate orders which swarmed in Patagonia during the mid-Tertiary and Pleistocene times are treated in the excellent memoirs of the Princeton University Patagonian expedi- tions by W. B. Scott. Herluf Winge has admirably monographed the fossil and recent edentates of Brazil. The mammalian fauna of the Deseado formations is described by F. B. Loomis of Am- herst College. These and other investigations are correcting the erroneous correlations by Ameghino, in which the older mammal-. bearing horizons of Patagonia were assigned to the Cretaceous. This more modern work indicates that the Pyrotherium beds are not older than Upper Eocene and that the Santa Cruz formation is of Lower Miocene Age. The Pleistocene fauna of Tarija, Bolivia, is the subject of a beautiful memoir by Boule and Thévenin (1920), in which the anatomy and relationships of ‘“‘ Mastodon’ andium and of the highly specialized horses Hippidium and Onohippidium are treated. ''* 16 Pleistocene Mammalian Faunas (North America, Europe).— The Pleistocene represents the climax of the Age of Mammals in point of differentiation and,richness‘of mammalian faunas. In Europe the Pleistocene faunas:have been, the subject of memoits by, Boule, Schoetensack and many others. In North America we have the teeming fauna of the Rancho, La /Brea; California, de- scribed by Merriam, Stock and 'their:colleagues of the university of California. The correlation of the American Pleistocene faunas has been treated especially by Osborn.and by Hay. Inthe preparation of; this article the writer is especially indebted for,the entire invertebrate section to the codperation of Miss Mar- jorie O’Connell, who has summarized the chief discoveries in Pre- and Postcambrian time and given a review of the outstanding literature ‘in the invertebrate field: He is also indebted to Charles D: Walcott; chief authority: on Cambrian and Precambrian life of the world, for the type figures assembled in Plate 1.; to Curators Matthew and Gregory of the American Museum for a revision of the text relating to the evolution of the vertebrates; and to the President | and Trustees of the American’ Museum for ‘permission to reproduce the photographs which are assembled toithe same scale on Plate II. ey. at (H. F. PALESTINE (see 20.600):—During the earlier years: of the decade:1gr1~21 little’ of importance occurred in: that country. | Afflicted: by ‘the: economic stagnation and ‘financial strain which affected the whole Ottoman Empire in consequence of the war with Italy (1911~2);,and the war with the Balkan States (1912-3), Palestine was unable to) develop herself in any way before the outbreak: of the. World: War in) ro14. Yet to a-section of her population the decision of the Palestinian Jews, in the autumn of 1913,'to reject!German and insist upon Hebrew as the language | of instruction and to-secede from the, Hilfsverein .and:set up | schools» of their own, was momentous: | The outbreak-of: the World: War; besides leading to’a renewed blockade of the coast, | and fresh military requisitions, also involved the expulsion or | interhment ‘of mtmerous ecclesiastics: -and: laymen of -Entente nationalities: and ‘the deportation of numbers! of Jews. >It was followed ‘at the beginning of 2915 by:one of the most destructive | visitations of locusts recorded for a generation. Thereafter until the! arrival of thé British army in the autumn of: 1917 the pros- | perity of the whole country slowly withered under the crushing burden of the war. ‘ jise «At the time of the British occupation of Jerusalemin:Dec.'1917 the'economic’situation of southern ‘Palestine was bad. Not ‘only hadithe Turks requisitioned far.and wide without-repayment, or against inadequate payment; but they had cut down:numbers of olives and: revenue-producing trees and carried: offsthe greater part ofthe agricultural and \draught) animals.: ‘The paper. cur- rency had:depreciated some 84% and was no longer! accepted by the producing classes-mostly outlying Moslem: peasants--who | would only) discover their concealed stores ofngrain for» gold, | ‘Fhe civil population: of: Jerusalem; dependent ordinarily ‘upon | the pilgrim traffic) or upon: the offerings of pious Jews) for its livelihood) was emaciated, and reduced: by starvation.’.The' only | | ton any longer to combine 0.E.T.A. with the work of the political products which ‘Jerusalem: had. to sell were designed» for | the pilgrim trade. and | were; unmarketable; consequently :at»the — beginning of the occupation many shops were able ‘to ‘offer:only cigatettes, picture-postcards and wild radishes for sale.’ In view of'this it was urgently necessaryito provide food for the exhausted: inhabitants: of Jerusalem and: Palestine, to !provide work for the purpose’of enabling them to earm money with which to: pay for the food, and to re-start' trade in order that the mer- cantile:community should: have something: to: barter:against the gold hoarded. by the péasantry> and thus: make’ it worth ‘the peasant’s while to: cultivate and:market his produce as he had for some time past tealized that his:gold' was unable to buy the trade goods he’ required: But there were grave difficulties—the | single line of railway by: which alone food or trade goods could be brought ‘from \Egypt was very fully occupied with the para- mount needs of the army. The daily tonnage:of supplies alone-— not including ‘munitions or transport of men or guns—varied | from! 800 to 2,300: Ammunition was often’ 2go:tons |per day. The civilian population: was) unaccustomed to the Egyptian currency and more than suspicious of paper money, and Egyptian | silver put into circulation was at once hoarded against the prox- modation thus :becamexavailable for other merchandise. PALESTINE imate return of the Turks, which was confidently predicted’ by enemy sympathizers who further assured every one that the Egyptian paper pound at ‘par was worth no more than the de- preciated Turkish paper lira and offered to. prove it: by readily exchanging Turkish for Egyptian pounds whenever possible— at a profit to themselves of 17s..7d.' on each; deal.. Yet without money the civilians could not buy food; without food they could scarcely walk from weakness, and there was every prospect of the establishment of a vicious circle. Brig.-Gen.. G. F. Clayton, (afterwards, Sir Gilbert, Clayton), chief political officer to Gen. Allenby, was appointed chief ad- ministrator and began to construct’ such a form of government as is provided for in “The Laws and Usages of War” laid down by international agreements embodied in the Hague Conyention. Transport for a few truck-loads of foodstuffs per week was secured fromthe military ‘railway, and lorries brought ‘it to Jerusalem until the army was able to reopen the narrow-gauge line from Ludd to the Holy City...Then a small consignment of trade goods came up from Egypt and-merchants were permitted to import small quantities from Egypt indépendently of. the over-burthened railway. The labour corps employed numerous civilians, paying them at first daily in Egyptian silver and paper, and then weekly in'cash or kind at the choicé:of the labourer. In this way the new currency came to its own, helped by the stringent measures taken by the military administration to suppress trafficking in or artificial’ depreciation of Egyptian paper. With the arrival-of trade goods in the towns the peasants began to spend their gold and sell:their produce so freely that it became unnecessary to import so much food:and more accom- But even so, goo tons of cereals had to be importéd monthly for the use of refugees alone. -Gen.'Clayton took other steps to restore public confidence and -reéstablish the amenities ‘of civilization: Bazaar gossip and rumour ‘which for some weeks| was hostile to the British was’ counteracted: by the publication of Arabic ‘and Hebrew’ editions of the newspaper, Lhe, Palestine News, which had:been:started, by the army: in March 1015, and intet- course with the greater-part of the world was rendered. possible by ‘the:restoratiom of-the postal service, for which’ special stamps for the use of the civilian’ population began to be issued on Feb. 16 z018; Steps were taken to reassure the Moslems; who were much alarmed at:reports’sedulously propagated:by the enemy, that all land» was to. be:given to the Jews, and:resident, British officers ‘were appointed to administer the various kazas.of the old Turkish regime. Thus military:governors were established.at Gaza, with ‘a deputy ‘at: Mejdel;. at) Jaffa, with a deputy at Ramleh}. at Beersheba;, at Hebron, with «a! deputy at: Deir. Aban; ‘and at Jerusalem, with depitties. at Bethlehem, Jericho and Ramalla. At first the Occupied Enemy Territory Administration-(“ O.E+ T.A.’’)) was'at Bir Salem, near Ramleh, the general headquarters of the army, but later when it became impossible for Gen. Clay- mission, Maj:-Gen.' Sir.Arthur W. Money was appointed chief administrator in April 1918, and he rémoved the administration to the ‘imposing and convenient Empress Victoria’ Hospice ‘built by) the Germans on the Mount. of Olives just before the war. In March the country had so far recovered that it became possible to collect taxés once'more, in May public:confidence was gréatly increased by-the skilful and tactful handling of the great Moslem religious festival of the Nebi Musa pilgrimage, invented by:the Turks as an artificial check on the great Christian gathering at Jerusalem for the Orthodox Haster, with which it always: coi- cides regardless:of thé Moslem calendar; and by:the successful way in which the dangerous, and often: fatal, ceremony of the Holy Fire on the Orthodox and Gregorian Easter was conducted by Col. Storrs the military governor of Jerusalem, and Haddad Bey the Syrian chief of police. - During the summer>the administration was able to resume the payment of revenues appropriated by international agreet ment to the servicé’of the Ottoman debt, but ithe income of ‘the Moslem Waafs (pious foundations): was used) for the: benefit of Moslem beneficiaries. in ‘Palestine instead of! being drained '' '' ''CAROLINGIAN REFORM] Latin Writinc. IV.—THE CAROLINGIAN REFORM AND’ THE MepievaL MinuscuLeE Hanp It has been stated above that in the Merovingian MSS. of the 8th century there was evident progress towards a settled minu- scule book-hand which only required a master hand to fix it in a purified and calligraphic form. This was effected under Charlemagne, in whose reign the revival of learning naturally led to a reform in handwriting. An ordinance of the year 789 required the revision, of Church books; and a more correct orthography and style of writing was the consequence. The abbey of St Martin of Tours was one of the principal centres from whence the reformation of the book-hand spread. Here, from the year 796 to 804, Alcuin of York presided as abbot; and it was specially under his direction that the Carolingian minuscule writing took the simple and graceful form which was gradually adopted to the exclusion of all other hands. In carrying out this reformation we may well assume that Alcuin brought to bear the results of the training which he had received in his youth in the English school of writing, which had attained to such proficiency, and that he was also beneficially influenced by the fine examples of the Lombard school which he had seen in Italy. In the new Carolingian minuscule all the uncouthness of the later Merovingian hand disappears, and the simpler forms of many of the letters found in the old Roman -half-uncial and minuscule hands are adopted. The character of Carolingian writing through*the oth and early part. of the toth century is one of grmeral uniformity, with ‘a contrast of light and heavy strokes, the limbs. of tall letters. being clubbed or thickened at the head by pressure on the pen. As to charac- teristic letters (fig. 44) the a, following the old type, is, in the oth century, still frequently open, in the form of 4; the bews of g are open, the letter somewhat resembling the numeral 3; and there is little turning of the ends of letters, as m and n. PEP Tae me comugemn cucu, quod enim execneftecurderp ales ee. parr surcerm filam e@cuocabifnomeneauNnhm Fic. 44.—Gospels, 9th century. (accipere mariam coniugem tuam quod enim ex ea nascetur de spzritu sanctio est. autem filium et uocabis nomen eius lesum) Pariet In the roth century the clubbing of the tall letters becomes Jess pronounced, and the writing generally assumes, so to. say, a thinner appearance. But. a great change is noticeable in the writing of the 11th century. By this time the Carolingian minuscule may be said to have put off its archaic form and to develop into the more modern. character of small letter. It takes a more finished and accurate and more upright form, the individual letters being drawn with much exactness, and gener- ally on a rather larger scale than before. This style continues to improve, and is reduced to a still more exact form of calligraphy in the 12th century, which for absolute beauty of writing is unsurpassed. In England especially (fig. 45) the writing of this century is particularly fine. atof ctarumulif fnladdeure firper alareurmla ai ple ¢earnibuf & fino crema gr€ caftra Lit Pooper dit Fic. 45.—Leviticus, A.D. 1176. (—culos cum aruinulis suis adoleuit super altare uitulum cum pelle et carnibus et fimo cremamns extra castra sicut preceperat dominus) As, however, the demand for written works increased, the fine round-hand of the 12th century could not be maintained. Economy of material became necessary, and a smaller hand with more frequent contractions was the result. The larger and xX 16 PALAEQOGRAPHY RC more distinct writing of the 11th and rath centuries is now replaced by a more cramped though still distinct hand, in which the letters are more linked together by connecting strokes, and are more laterally compressed. This style of writing is characteristic of the 13th century. But, while the book-hand of this period is a great advance upon that of a hundred years earlier, there is no tendency to a cursive style. Every letter is clearly formed, and generally on the old shapes, The particular letters which show weakness are those made of a succession of vertical strokes, as m, n, u. The new method of connecting these strokes, by turning the ends and running on, made the distinction of such letters difficult, as, for example, in such a word as minimi. The ambiguity thus arising was partly obviated by the use of a small oblique stroke over the letter 1, which, to mark the double letter, had been introduced as early as therzth century. The dot on the letter came into fashion in the 14th century. Clignentyoote op placer ent ernie pocitimfi vekeans, Lerit ofif ques fererredy pees uti be] tefoprramis.andifamoreoy tn quota Yirans.Cgvaiizoni men tinen? ota Bet. Ving Hs ear. Abia nob urrelung Fic. 46.—Bible, 13th century. (Eligite hodie quod placet cui seruire potissimum debeatis. Utrum diis quibus seruierunt patres uestri in mesopotamia, an diis amoreorwm in quorum terra habitatis. Ego autem et domus mea seruiemus domino. Respon- ditque popzlus et ait, Absit a nobis ut relinquamus dominum) In MSS. of the 14th century minuscule writing becomes slacker, and the consistency of formation of letters falters. There is a tendency to write more cursively and without raising the pen, as may be seen in the form of the letter a, of which the character- istic shape at this time is @, with both bows closed, in contrast with the earlier a. In this century, however, the hand still remains fairly stiff and upright. In the 15th century it becomes very angular and more and more cursive, but is at first kept within bounds. Inthe course of the century, however, it grows more slack and deformed, and the letters become continually more cursive and misshapen. An exception, however, to this disintegration of minuscule writing in the later centuries is to be observed in church bocks. In these the old set hand of the 12th and 13th centuries was imitated and continued to be the liturgical style of writing. It is impossible to describe within limited space, and without the aid of plentiful illustrations, all the varieties of handwriting which were developed in. the different’ countries of. western Europe, where the Carolingian minuscule was finally adopted to the exclusion of the earlier national hands. In each.country, however, it_acquired, in a greater or less degree, an individual national stamp which can generally be recognized and which serves to distinguish MSS. written in different localities. A broad line of distinction may be drawn between the writing of northern and southern Europe from the 12th to the 15th century. In the earlier part of this period the MSS. of England, northern France and the Netherlands are closely connected. Indeed, in the 12th and 13th centuries it is not always easy to decide as to which of the three countries a particular MS. may belong. As a rule, perhaps, English MSS. are written with more sense. of gracefulness; those of the Netherlands in darker ink. From the latter part of the 13th century, however, national character begins to assert itself more distinctly. In southern Europe the influence of the Italian school of writing is manifest in the MSS. of the south of France in the 13th and 14th centuries, and also, though later, in those of Spain. That elegant. roundness of letter which the Italian scribes seem to have inherited from the bold characters. of the early papal chancery, and more recently from Lombardic models, was generally adopted in the book-hand of those districts. It is especially noticeable in calligraphic specimens, as in church books—the writing of Spanish MSS. in this style being distinguished by the blackness of the ink. The medieval minuscule writing of Germany stands apart. It never attained to the beauty of the hands of either the north or 3 ''578 the south which have been just noticed; and from its ruggedness and slow development German MSS. have the appearance of being older than they really are. The writing has also very commonly a certain slope in the letters which compares unfavour- ably with the upright and elegant hands of other countries. In ‘western Europe generally the minuscule hand thus nationalized ran its course down to the time of the invention of printing, when the so-called black letter, or set hand of the r5th century in Germany and other countries, furnished models for the types. But in Italy, with the revival of learning, a more refined taste set in in the production of MSS., and scribes went back to an earlier time in search of a better standard of writing.’ Hence; in the first quarter of the 15th century, MSS. written on the lines of the Italian hand of the early 12th century begin to appear, and become continually more numerous. This revived hand was brought to perfection soon after the middle of the century, just at the right moment to be adopted by the early Italian printers, and to be perpetuated by them in their types. English Cursive Charter-Hands.—It must also not be forgotten that by the side of the book-hand of the later middle ages there was the cursive hand of everyday use. This is represented in abundance in the large mass of charters and legal or domestic documents which remains. Some notice has already been taken of the development of the national cursive hands in the earliest times. From the 12th century downwards these hands settled into well defined and distinct styles peculiar to different countries, and passed through systematic changes which can be recognized as characteristic of particular periods. But, while the cursive hand thus followed out its own course, it was still subject to the same laws of change which governed the book-hand; and the letters of the two styles did not differ at any period in their organic formation. Confining our attention to the charter- hand, or court-hand, practised in England, a few specimens may be taken to show the principal changes which it developed. In the 12th century the official hand which had been introduced after the Norman Conquest is characterized by exaggeration in the strokes above and {below the line, a legacy of the old Roman cursive, as already noted. There is also a tendency to form the tops of tall vertical strokes, as in 6, h, J, with a notch or cleft. The letters are well made and vigorous, though often rugged. Wane Onan Delt fal Las eee VI mos/ Wu acon, Ca Peace” Mase’ Fic. 47.—Charter of Stephen, A.D. 1136-1139. (ef ministris et omnibws fidelibus suis Francés et Regine uxoris mee et Eustachii filii mei dedi e¢ concessi ecclesie Beate Marie) As the century advances, the long limbs are brought into better proportion; and early in the 13th century a very delicate fine-stroked hand comes into use, the cleaving of the tops being now a regular system, and the branches formed by the cleft falling in a curve on either side. This style remains the writing of the reigns of John and Henry III. © Gulls (Pherae Leeds mifletong cabin: Botiong that certain species actually pass from one formation ‘to another. In the meantime the foundations of palaeobotany were being laid, (1804) by .Ernst) Friedrich» von» Schlothéim: (1764-1832), (1822) by | Kaspar, Maria, Sternberg (x 7602 1838) and (1838) by: Théophile Brongniart, (1801-1876), Following Cuvier’s eens sur les ossemens: fossiles, the rich succession of Tertiary; mammalian life |) was! gradually revealed. to, France) through the, explorations.and descriptions of such authors,as Croizet, Jobert;:de Christol,,Eymar, Pomel and. Lartet, during,a period: of rather. dry; systematic work, which included, however, the broader: generalizations of Henri Marie on ae de Blainatlle (4778~2850);,and culminated ‘in the, comprehensive, treatises on Tertiary palacontology of Paul Gervais (1816-1879)... Extending the knowledge of the extinct mammals. of, Germany, the, principal: contributors; were Georg August. Goldfuss .(2782~1848),,- Georg: Friedrich; von, Jaegar (1785+1866),: Felix F;, Plieninger|(2807+1873). and: Johann. Jacob Kaup, (180371873)... As Cuvier founded: ‘the; palaeontology. of mammals, and, reptiles, so Louis.Agassiz’s epoch-making ‘works Recherches sur les poissons fossiles: (183371845) laid, the secure foundations,of palaeichthyology, and were,fcllowed by. Christian Heinrich Pander’s (279451865). classic. memoirs on the fossil fishes of, Russia;,,, In philosophy, Agassiz was distinctly a disciple of, Cuvier.and\supporter.of the doctrine;of special creation, and to a more limited)extent; of cataclysmic! extinctions.,..Animals of the next higher order; the amphibians.of the coal,measures andthe, Permian, were: first comprehensively, treated. in| the masterly, memoirs) ;of. Christian, Erich, Hermann von! Meyer (z801-1869),beginning in,2829,.,especially, in, his. Betirdge zur Petrefactenkunde (1829-1830) and, his Zur Fauna der Vorwelt (4, vols., 1845-1860). Successive discoveries gradually revealed the world of extinct Reptilia;in 1821 Charles Kénig (1784-1851), the first, keeper, of the; mineralogical collection: in: the, British Museum, described, Lchihyosaurus from; ‘the Jurassic; in the same, year, William .Daniel..Conybeare. (1787-1857). described Plesiosaurus; and,-a, year, later. (1822)..Mosasaurus; in.-1824 William.Buckland, described the great,carnivorous dinosaur Megalosaurus; while Gideon Algernon, Mantell (1790-1852), in 1848 announced the discovery of [guanodon.. .Some of the fossil Reptilia. of ,France’ were, made known through St Hilaire’s researches on the:Crocodilia (1831), and those of J. A.'Deslong- champs (1794-1867),and his'son,on. the teleosaurs,, or long- snouted..crocodiles.., Materials accumulated, far more rapidly, however, than the) power-of generalization, and, classification. Able as von. Meyer. was, his, classification of the Reptilia failed because, based .upon,the single; adaptive -characters..of foot structure.. The reptiles awaited. a great classifier, and such a one appeared. in England. in the person’ of Sir Richard Owen (180471892), the direct. successor of, Cuvier and a comparative anatomist of. the first rank,...Non-committal)as regards evolu- tion, he vastly, broadened, the field of vertebrate palaeontology by his descriptions of the extinct fauna.of England, of South America, (including especially ‘the great edentates revealed. by the voyage of the. “‘ Beagle’’), of Australia (the ancient and modern marsupials) and of, New Zealand, (the great struthious birds). His contributions. on, the, Mesozoic, reptiles. of Great Britain. culminated.in his,complete rearrangement and classifi- cation of this group, one of his greatest services to palaeontology. Meanwhile the researches of Hugh Falconer (1808-1865), and of Proby., Thomas .Cautley,. (1802-1871) in..the,-sub-Himalayas PALAEONTOLOGY brought, to light’ the marvellous *fauna® ofthe Siwalik hills of — India, published in Fauna antigua Sivalensis (London, 1845) _ and in the: volumes of) Falconer’s: individual reséarches.) ‘The ancient life:of the Atlantic: border of North America was also becoming known through the work of the pioneer vertébrate palaeontologists Thomas Jefferson (1743-1826), Richard Harlan (1796-1843), Jeffries) Wyman: (1814-1874) and Joseph Leidy (1823-1891). This was followed by the revelation of the vast ancient life of the western half of the American continent; which was destined to revolutionize: the science. » The: master ‘works of Joseph Leidy began with the first-fruits of western exploration in 1847 and extended through a series of grand memoirs, culmina- ting in 1874. Leidy adhered strictly to Cuvier’s exact descriptive methods, and while an evolutionist and recognizing clearly the genetic relationships of the horses and: other groups, he never indulged:in speculation. The history of invertebrate palaeontology during the second period is more closely connected with the rise of historic geology and: stratigraphy, especially with the settlement’ ofthe great andiiminor ‘time divisions of the earth’s shistory; ©The! path- breaking works of Lamarck were soon followed by the monu- mental treatise of (Gérard Paul Déshayes (1795-1875) entitled Descriptions des’ coquilles »fossiles des endirons:de | Paris \(1824= 1837), the first of a series-of great contributions by this and other authors.',These and other early ‘monographs: on’ the: Tertiary shells of the Paris basin, of thé environs of Bordeaux, and of the sub-Apennine formations | of) :Italy,|| brought .out«the- striking distinctness! of these’ faunas from each) othersand from other molluscan faunas. Recognition’ of- this. threefold: character led Deshayes to establish ‘a threefold division of ‘the ‘Tertiary based on» the percentage of molluscs: belonging! to ‘types now living found)in each. Te these divisions Lyell gave:in 1833 the names;Hocene, Miocene and Pliocene. ; James Hutton (1726-1797) hadiset forth'(1788) the principle that during all geological time there -has «been no | essential changein the character of events, and that uniformity. of law is perfectly’ consistent .with mutability. in’ the: results. Lyell marshalled all the observations he could collect:in; support of this | principle, teaching that-the present:is-the key tothe past; and, arraying all. obtainable evidence against. the»cataclysmic theories. of .Cuvier..; He: thus: exerted a. potent. influence on palaeontology through his persistent advocacy of ‘uniformi- tarianism, a doctrine with which Lamarck should also be credited. As among the vertebrates, materials were accumulating rapidly for the great generalizations. which. were to! follow! in: the. third period. De Blainville added, to. the knowledge of; the. shells of ‘the Paris basin; Giovanni Battista, Brocchi (1772-1826) in 1814, and Luigi: Bellardi (1818-1889)..and Giovanni Michelotti (born 1812),in 1840, described the Pliocene molluses of the sub- Apennine formation. of Italy; from» Germany and \Austria appeared the epoch-making works of. Heinrich Ernst Beyrick (2815-1896) and of Moritz Hoernes (1815-1868). We. shall pass over’here the labours of Adam ‘Sedgwick (1785-1873) and Sir Roderick Murchison (1792-1871)in: the Palaeozoic of England, which because:of their close relation to stratigraphy more properly concetn geology; but must mention the grand contributions ‘of Joachim. Barrande (1799-1883), published in his Systéme silurien du centre dela Bohéme, the first volume of which appeared in 1852. While establishing the historic divisions of the Silurian in Bohemia, Barrande also propounded: his famous theory of “colonies,” by, which, he attempted to. explain the aberrant. occurrence of ,strata, con- taining animals of a. more advanced stage among strata containing earlier and more primitive faunas; his assumption was that the second fauna: had migrated from) an unknown neighbouring region. It,is proved that the. specific instances on, which Barrande’s generalizations were founded were, due to his misinterpretation of the overturned and faulted strata, but his conception of the simultaneous existence of two faunas, one of more ancient.and one of more modern type, and of. their alternation in a given area, was based on sound philosophical principles and has been eonfirmed by more recent work. j '' PALAEONTOLOGY The greatest (generalization of this: second period, however, was that partly prepared for by d’Orbigny, as will be more fully explained later in this article, and clearly expressed by Agassiz —namely, the law of repetition of ancestral stages of life in the course of the successive stages of individual development. ‘This law. of -recapitulation, subsequently termed) the “‘ biogenetic law ” by Ernest’ Haeckel, was the greatest philosophic: contri- bution of this period; and proved to be not only: one of ‘the bulwarks,. ofthe. evolution theory but: one: of the’ most important principles in the method of palaeontology. On: the whole, as in the case of vertebrate palaeontology, the pre-Darwinian period of invertebrate palaeontology was one: of rather dry systematic description, in which, however, the applications of the science gradually extended. to many regions» of the world and to all divisions of the kingdom of invertebrates. TI]. —Tuirp Hisroric, PERIOD Beginning with the publication of «Darwin's great works, “Narrative of the Surveying Voyages of H.M.S: ‘Adventure’ and ‘ Beagle’ ’? (1839), and “‘ On the Origin of Species by Means of Natural. Selection.” (1859).—A review of the two first classic works! of Charles Robert Darwin (1809=1882)\ ‘and /of\ their influence proves that he was the founder of modern palaeon- tology: Principles of descent and other applications of uniformi- tarianism which had been struggling for expression in’ the writings of Lamarck, St Hilaire and de Blainville here found their true interpretation, because the geological succession, the rise; the migrations, the extinctions, were all connected: with the grand central idea of evolution from primordial forms. A close study of the exact modes of evolution and of the philosophy of evolution is the distinguishing feature of this period. It appears’ from comparison of the work ‘in the two great odivisions of vertebrate and invertebrate palaeontology made for the first time in this article that in accuracy of observa- tion and in close philosophical analysis of facts the students of invertebrate palaeontology led the way. This was due to the much greater completeness and abundance of material afforded among invertebrate fossils, and it was manifested in the demon- stration of two great principles or laws: first, the law of recapitu- lation, which is found in its most ideal expression in the shells of invertebrates; second, in the law of direct genetic succession through very gradual modification. It is singular that the second law is still ignored by many zoologists. Both laws were. of paramount importance, as direct evidence of Darwin’s theory of descent, which, it will be remembered, was at the time regarded merely as an hypcthesis. Nevertheless, the tracing of phylogeny, or direct lines of descent, suddenly began to attract far more interest than the naming and description of species. The Law of Recapitulation. Acceleration. » Retardation.—This law, that in| the stages of growth of individual development (ontogeny), an animal repeats the stages of its ancestral evolution (phylogeny) was, as we have stated, anticipated by d’Orbigny. He recognized the fact that the:shells of molluscs, which grow by successive additions, preserve unchanged the whole series of stages of their individual development, 30 that each shell of a Cretaceous ammonite, for example, represents five stages of progressive modification as follows: the first: is the période embryonnaire, during which the shell is smooth; the second and third represent periods of elaboration and ornamentation; the fourth is a period of initial degeneration; the fifth and last a period of degeneration when ornamentation becomes obsolete and the exterior smooth again, as in the young. »D’Orbigny, being: a special creationist, failed to recognize the bearing of these individual stages on evolution. Alpheus Hyatt (1838- 1902) was the first to discover (1866) that these changes in the form of the ammonite shell agreed closely with those which had been passed through in the ancestral history of the ammonites. In anepoch-making essay, On the Parallelism between the Different stages of Life in the individual and those in the entire group of the Molluscous Order Tetrabranchiata (1866), and in a number, of subsequent memoirs, among which Genesis of the Arietidaz (1889) 583 and) Phylogeny of ano cquired: Characteristic’ (1894) ‘should ‘be mentioned; ‘he laid the: foundations,» by methods of the most exact analysis, for'all future recapitulation work of invertebrate palaeontologists. -He'showed that«from each’ individual shell of an ammonite the entire ancestral series may be reconstructed, and that, :;whilesthe earlier: shell-whorls retain’ the characters of the adults of preceding members:of the series;‘a shell in its own adult: stage adds ‘a: new character, which in turn’ becomes the’ pre-adult: character of the: types which will succeed: it; finally, that this comparison between the revolutions of the’ life of an ‘individual and the life of the entire order of ammonites is wonder- fully harmonious.and precise. 'Moreover,)the last stages of ‘individual life are: propheticonot only -of) future: rising’ and progressing derivatives; but:in’ the case’ of senile individuals of future declining and degradational series. Thus the recapitulation law, which had been built up indepen- dently from:the observations and speculations on vertebrates by Lorenz Ofen (1779=1851), Johann Friedrich Meckel (1781-1833), St Hilaire, Karl ‘Ernst von Baer (1792~1876) and others, and had been applied (1842~1843) by Karl Vogt (1817-1895)'and Agassiz, in their respective: fields of observation, to comparison of indi- vidual’ stages with the adults of the same group in preceding geological periods, furnished the key to the determination of the ancestry of the invertebrates generally. Hyatt went further and: demonstrated that: ancestral characters are passed through by successive descendants at a more and more accelerated rate’ in each generation, thus giving ‘time for the appearance of new characters in’ the adult} His ‘law’ of acceleration’? together - with the complementary “law of retardation,” or the slowing up in'the development of certain characters (first’ propounded’ by E. D. Cope); was also a philo- Af la 2a 4 BRIb4 2b f 3b 4 CHie+ 2c Bc { 4c { DE ld+-24+f 34+ + 4d bd | Eble 2e4 se ——f fe | bef — be FE 1p 2t-+-— 33 + — 4¢ —+— 5 +-— 8t- + - 71 GE ep 2g 4 8g — 1g —J 5 be }-$_—— 2 —_ 4 HF 1b-+ 2h+—3h — 4h { Th aa! TE a +—3i —— 45 — 7 {_—gj—4 (From the American Naturalist.) Fic. 6. sophie contribution of the first importance (see fig. 6 and Plate III., fig. 7). In the same year, 1866, Franz Martin Hilgendorf (1839- ) studied the shells of Planorbis, from the Miocene lake. basin underlying the present’ village of Steinheim in Wiirttemberg, and introduced the method of examination of large numbers of individual. specimens, a» method which has become. of | prime importance in’ thé science. . He discovered the actual transmu- tations in direct genetic series: of species on the successive deposition levels of the old lake basin. This study of direct genetic series marked another great advance, and became possible in invertebrate palaeontology long before it' was introduced among the vertebrates: Hyatt; in’ a re-examination of the Steinheim deposits, proved that successive modifications occur at the same level as. well: as.in\ vertical, succession. Melchior Neumayr (1845-1890) and) C. M: Paul similarly: demonstrated genetic series of Paludina (Vivipara) in the Pliocene lakes of Slavonia (1875). ) . The Mutations of Waagen. . Orthogenesis:—In1869; Wilhelm Heinrich Waagén (1841-1900) ‘entered the field with the study of Ammonites subradiatus: He proposed the term “‘ mutations ”’ for the minute progressive changes of single characters: in definite directions as observed in successive stratigraphic levels. Even when seen. in minute features only he recognized them as constant progressive characters or ‘‘ chronologic: varieties ” in ''534 contrast’ with contemporaneous or ‘‘ geographic. varieties,” which he considered inconstant and of slight systematic value: More recent analysis has shown, however, that certain modifica- tions. observed within the same stratigraphic level are: really grades of mutations which show divergences: comparable to those found in successive levels. The collective term ‘“muta- tion,’ as now employed by palaeontologists, signifies a type modified to a slight degree in one or more of its characters along a progressive or definite line of phyletic development. The term “‘ mutation ”’ also applies to a single new character and for distinction!» may be known as “the mutation of Waagen.” This definitely directed evolution; or development in: a few determinable directions, has since been termed ‘“‘ orthogenetic evolution,” and is recognized: by all\ workers in invertebrate palaeontology and phylogeny as fundamental because the facts of invertebrate palaeontology admit of no other interpretation. Among the many who followed the method of attack first outlined. by Hyatt, or who independently. discovered: his method, only a few can be mentioned here—namely, Waagen (1869), Neumayr (1871), Wiirttemberger (1880), Branco (1880), Mojsisovics (1882), Buckman:(1887), Karpinsky (1889), Jackson (2890), Beecher (1890), Perrin-Smith (1897), Clarke © (1898) and Grabau (1904). Melchior Neumayr, the great Austrian palaeontologist, especially extended the philosophic foundations of modern invertebrate palaeontology, and: traced a number of continuous genetic series (formenrethe) in successive horizons. He also demonstrated that mutations have this special - or distinctive character, that they repeat in the same direction without oscillation or retrogression: | He expressed great reserve as to the causes of these mutations: He was the first to attempt a comprehensive treatment of all invertebrates from the genetic point of view; but unfortunately his great work, entitled Die Stimme des Thierreichs (Vienna and Prague, 1889), was uncompleted. The absolute agreement in the results independently obtained by these various investigators, the interpretation of individual development as the guide to phyletic~ development; ~the demonstration of continuous, genetic series, each mutation falling into its proper place and all showing a definite direction, constitute contributions to biological philosophy of the first importance, which have been little known or appreciated by zoologists because of their publication in monographs of very special character. Vertebrate..Palaeontology after Darwin.—The impulse which Darwin gave to vertebrate palaeontology was immediate and unbounded, finding expression especially in the writings of Thomas Henry Huxley (1825-1895) in England, of Jean Albert Gaudry (b. 1827) in’ France, in’ America’ of Edward Drinker Cope (1840-1897) and Othniel Charles Marsh (1831-1899). Fine examples'of the spirit ‘of the period as applied'to extinct Mammalia are: Gaudry’s Animaux fossiles et géologie de V Attique (1862) on the Upper Miocene fauna of Pikermi near Athens; and the remarkable memoirs of Vladimir: Onufrievich> Kowalevsky (1842=1883), published ini1873; These works swept aside the dry traditional fossil lore which had been accumulating in France and Germany. They breathed the new spirit of the recognition of adaptation and descent. . In 1867—1872 Milne Edwards published his memoirs cn the Miocene birds of central France. Huxley’s development of the method of palaeontology should be studied in his collected memoirs (Scientific Memoirs of Thomas Henry Huxley, 4 vols., 1898). Im Kowalevsky’s' Versuch einer natiir- lichen Classification der Fossilen Hufthiere (1873) we finda model union of detailed inductive study with theory and ‘working hypothesis. All these writers attacked the problem of descent, and published preliminary phylogenies of such animals as the horse, rhinoceros and elephant, which time has proved to be of only general value and not at all comparable to the exact phylogenetic series which were being established by invertebrate palaeontologists. Phyletic gaps began to be filled in this general way, however, by discovery, especially through remarkable 1 The Dutch botanist, De Vries, has employed the term in another sense, to mean a‘slight jump or saltation. PALAEONTOLOGY discoveries in North America by Leidy, Cope and Marsh, and the ensuing phylogenies gave enormous prestige to palaeontology. Cope’s philosophic contributions to palaeontology began in 1868 (see essays in The Origin of the Fittest; New York, 1887, and The Primary Factors of Organic Evolution, Chicago, 1896) with the independent discovery and: demonstration among verte- brates of the laws of acceleration and retardation. »To the law of “recapitulation” he unfortunately applied) Hyatt’s term “parallelism,” a term which is used now in another sense: He especially pointed out the laws of othe: ‘extinction: of the specialized” and ‘survival of the non-specialized’” forms of life, and’challenged Darwin’s principle of selection as an explana- tion of the origim of adaptations by saying that the ‘survival ‘of the fittest’? does not explain the “‘ origin of the fittest.”’ He personally sought to demonstrate) such origin, first, in the existence of a specific internal growth force, which he termed bathmic force, and second in the direct inheritance of acquired mechanical modifications of the teeth and feet. He thus re- vived Lamarck’s views. and. helped to found the ‘so-called: néo- Lamarckian school in America. To this’school A. Hyatt, W. H. Dall and:many other invertebrate palaeontologists subscribed. History of Discovery. Vertebrates.—In discovery the theatre of interest has shifted from continent to continent, often in a sensational manner. ‘After a long period of gradual revelation of the ancient life of Europe, extending eastward to'Greece, eastern Asia and to Australia, attentions became centred on» North America, especially on Rocky Mountain exploration. New and unheard-of orders of amphibians, reptiles and mammals came to the surface of knowledge, revolutionizing thought, demonstrating the evolution theory; and solving some of the most: important problems of descent. Especially noteworthy was the discovery of birds with teeth both in Europe (Archaeopteryx) and in North America (Hesperornis), of Eocene stages in the history of ithe horse, and of the giant dinosauria of the Jurassic and: Cretaceous in North America: Then the stage of novelty suddenly shifted to South America, where after the: pioneer labours of Darwin, Owen and Burmeister, the field of our knowledge was suddenly and vastly extended by explorations: by the brothers:Ameghino (Carlos.and Florentino): We were in the midst-of more thorough examination of the ancient world of Patagonia, of the Pampean region and of its submerged sister continent Antarctica, when the scene shifted to North Africa through the: discoveries of Hugh J. LeBeadnell and Charles W. Andrews: ‘These latter discoveries supply us with the ancestry of the elephants and: many other forms. © They round out our knowledge of Tertiary history, but leave the problems of the Cretaceous mammals and of their relations to, Tertiary mammals still unsolved: Similarly, the Mesozoic reptiles have been traced successively to various parts of the world from France, Germany, England, to North America and South America, ‘to Australia and New» Zealand. and to northern Russia, from Cretaceous times back into the Permian, and by latest reports into the Carboniferous. Discovery of Invertebrates~The most striking feature of exploration for invertebrates, next to the world-wide extent to which exploration has been carried:on and results: applied; is the early appearance of life. Until comparatively recent times the molluscs were considered as appearing on the limits of the Cambrian and Ordovician; but Charles D. Walcott has described a tiny lamellibranch (Modioloides) from the inferior Cambrian, and he reports the gastropod (?) genus Chuaria from the pre- Cambrian. Cephalopod molluscs have been traced) back to the straight-shelled nautiloids of the genus Volborthella, while true ammonites have been found in the inferior Permian of the Conti- nent and by American palaeontologists in the true coal measures. Similarly, early forms of the crustacean sub-class Merostomata have been traced to the pre-Cambrian of North America. Recent discoveries of vertebrates are of the same significance, the most primitive fishes being traced to the Ordovician or base of the Silurian,? which proves that we shall discover more 2 Professor Bashford Dean doubts the fish characters of these Ordovic Rocky Mountain forms. Frech admits their fish character but considers the rocks infaulted Devonic. '' | PALAEONTOLOGY ancient chordates in the Cambrian or even pre-Cambrian. ‘Thus all recent discovery tends to carry the centres of origin and of dispersal of all animal types farther and farther back in geological time. TV.—RELATIONS OF PALAEONTOLOGY TO OTHER PHYSICAL EARTH SCIENCES Geology and Palaeophysiography.—Fossils are not absolute timekeepers, because we have little idea of the rate of evolution; they are only relative timekeepers, which enable us to check off the period of deposition of one formation with that of another. Huxley questioned the time value of fossils, but recent research has tended to show that identity of species and of mutations is, 4 on the whole, a guide’to synchroneity, though the general range of vertebrate and invertebrate life as well as of plant life? is generally necessary for the establishment of approximate synchronism. Since fossils afford an immediate and generally a decisive clue to the mode of deposition of rocks, whether marine, lacustrine, fluviatile, flood plain or aeolian, they lead us naturally into palaeophysiography. Instances of marine and lacustrine analysis have been cited above. The analysis of continental faunas into those inhabiting rivers, lowlands, forests, plains or uplands, affords a key to physiographic con- ditions all through the Tertiary. For example, the famous bone-beds of the Oligocene of South Dakota have been analysed by W..D. Matthew, and are shown to contain fluviatile or channel beds with water and” river-living forms, and neighbouring flood-plain sediments containing remains of plains-living forms. Thus we may complete the former physiographic picture ofa vast flood plain east of the Rocky Mountains, traversed by slowly meandering streams. As already intimated, our knowledge of palaeometeorology, or of past climates, is derivable chiefly from fossils. Suggested two centuries ago by Robert Hooke, this use of fossils has in the hands of Barrande, Neumayr, the marquis de Saporta (1895), Oswald Heer (1809-1883), and.an army of followers developed into a sub-science of vast importance and interest. It is true that a great variety of evidence is afforded by the composition of the rocks, that glaciers have left their traces in glacial scratch- ings and transported boulders, also that proofs of arid or semi- arid conditions are found in the reddish colour of rocks in certain portions of the Palaeozoic, Trias and Eocene; but fossils afford the most precise and conclusive evidence as to the past history of climate, because of the fact that adaptations to temperature have temained constant for millions of years. All conclusions derived from the various forms of animal and plant life should be scrutinized closely and compared. The brilliant theories of the palaeobotanist, Oswald Heer, as to the extension of a sub-tropical climate to Europe and even to extreme northern latitudes in Tertiary time, which have appealed to the imagina- tion and found their way so widely into literature, are now challenged by J. W. Gregory (Climatic Variations, their Extent and Causes, International Geological Congress, Mexico, 1906), who holds that the extent of climatic changes in past times has been greatly exaggerated. It is to palaeogeography and-zoogeography in their reciprocal relations that palaeontology has rendered the most unique services. Geographers are practically helpless as historians, and problems of the former elevation and distribution of the land and sea masses depend for their solution chiefly upon the palaeontologist. With good reason geographers have given reluctant consent to some of the bold restorations of ancient continental outlines by palaeontologists; yet some of the greatest achievements of recent science have been in this field. The concurrence of botanical (Hooker, 1847), zoological, and finally of palaeontological evidence for the reconstruction of the continent of Antarctica, is one of the greatest triumphs of biological investigation. To the evidence advanced by a great number of authors comes the clinching testimony of the existence of a number of varieties of Australian marsupials in Patagonia, as originally discovered by Ameghino and more exactly described by members of the Princeton Patagonian expedition staff; while ee 585 the fossil shells of the Eocene of Patagonia’ as analysed by Ortmann’ give evidence of the existence of a continuous shore- line, or at least of shallow-water areas; between Australia, New Zealand and South America. This line of hypothesis and demonstration is typical of. the palaeogeographic méthods generally—namely, that vertebrate palaeontologists, impressed by the sudden appearance of extinct forms of continental life, demand land connexion or migration tracts from common centres of origin and dispersal, while the invertebrate palaeon- tologist alone is able to restore ancient coast-lines and determine the extent.and width of these tracts. Thus has been built up a distinct and most important branch. The great contributors to the palaeogeography of Europe are Neumayr and Eduard Suess (b. 1831), followed by Frech, Canu, de Lapparent and others. Neumayr was the first to attempt to restore the grander earth outlines of the earth as a whole in Jurassic times. Suess outlined the ancient relations of Africa and Asia through his ‘“‘ Gondwana Land,” a land mass practically identical with the “‘ Lemuria ”’ of zoologists. South American palaeogeography has been traced by von Ihring into a northern land mass, “ Archelenis,”’ and a southern mass, “‘ Archiplata,” the latter at times united with an antarctic continent. Following the pioneer studies of Dana, the American palaeontologists and strato- graphers Bailey Willis, John M. Clarke, Charles Schuchert and others have re-entered the study of the Palaeozoic geography of the North American continent with work of astonishing precision. Zoogeography.—Closely connected with palaeogeography is zoogeography, the animal distribution of past periods. The science of zoogeography, founded by Humboldt, Edward Forbes, Huxley, P. L. Sclater, Alfred Russel Wallace and others, largely upon the present distribution of animal life, is now encountering through palaeontology a new and fascinating series of problems. In brief, it must connect living distribution with distribution in past time, and develop a system which will be in harmony with the main facts of zoology and palaeontology. The theory of past migrations from continent to continent, suggested by Cuvier to explain the replacement of the animal life which had become extinct through sudden geologic changes, was prophetic of one of the chief features of modern method—namely, the tracing of migrations. With this has been connected the theory of “‘ centres of origin’ or of the geographic regions where. the chief characters of great groups have been established. Among invertebrates Barrande’s doctrine of centres of origin was applied by Hyatt to the genesis of the Arietidae (1889); after studying thousands of individuals from the principal deposits of Europe he decided that the cradles of the various branches of this family were the basins of the Cote d’Or and southern Germany. Ortmann has traced the centre of dispersal of the fresh-water Crawfish genera Cambarus, Potamobius and Cambaroides to eastern Asia, where their common ancestors lived in Cretaceous time. Similarly, among vertebrates the method of restoring past centres of origin, largely originating with Edward Forbes, has developed into a most distinct and important branch of historical work. This branch of the science has reached the highest development in its application to the history of the extinct ‘mammalia of the Tertiary through the original work of Cope and Henri Filhol, which has been brought to a much higher degree of exactness recently through the studies of H. F. Osborn, Charles Depéret, W. D. Matthew and H. G. Stehlin. V.—RELATIONS OF PALAEONTOLOGY TO OTHER ZOOLOGICAL METHODS Systematic Zoology.—It is obvious that the Linnaean binomial terminology and its subsequent trinomial refinement for species, sub-species, and varieties was adapted to express the differences between animals as they exist to-day, distributed contemporane- ously over the surface of the earth, and that it is wholly inadapted to express either the minute gradations of successive generic series or the branchings of a genetically connected chain of life. Such gradations, termed “ mutations”? by Waagen, are distinguished, as observed, in single characters; they are the ''586 PALAEONTOLOGY nuances, or grades of difference, which are the more gradual | which is essentially based:on a theory of interrupted ior: dis- the more finely we dissect’ the geologic column; while the terms species, sub-species and variety are generally based upon.a sum of changesin several characters. Thuspalaeontology has brought to light an entirely new nomenclatural problem, which’ can only be solved by resolutely adopting an entirely different principle. ontogeny—namely, continuous characters, is inapplicable. Embryology and Ontogeny.—In following the discovery of the law of recapitulation among palaeontologists we have clearly stated the chief contribution of palaeontology to the science of the correspondences and differences between Formations’ in Western United States and Characteristic Type of Horse in Each Fore Foot Hind Foot ‘Teeth Quaternary Recent , , : . . | or mie ( Age of Man] Pleistocen One Toe Qesiles oted ' Splints of Splints of, g Long- 2™and 4! digits 274 and 4'P digits Pliocene ' — Cement~ covered ‘Three Toes Three Toes Mi F Side toe toes Side toes je. ocene \ not touching the ground not touching the ground Tertiary | Three Toes 2 or Side toes Ww { Olid. ching the ground; _| 4 Age of -| Oligocene “pot tf stag Three Toes Side toes Mammals | i touching the ground \ z Short- ; » Crowned! a Four Toes M4 without ri | “Cement Eocene ; ; Four Toes. Thrée Toes 4 ZB Hyracotherium I Splint of 12 of 17 digit Splint of Stdigit. (Eohippus), , Whee 4 Cretaceous Hypothetical Ancestors with Five Toes on Each Foot e) 1 { ge Jurassic, and Teeth like those of Monkeys etc. Reptil feck Triassic. : i J ’ AIO! Reproduced by permission of the American Museum of Natural History BiGio, This revolution may be pepapnshed by adding the term “mutation ascending ” ‘mutation descending,’’. for the minute steps of go ope rsnS) To and theterm phylum, as employed in Germany, for the minor anid major, branches of genetic series. Bit by bit mutations are added to.each other in different single characters until a sum or degree of mutations is reached, which no zoologist would hesitate to place in a,separate species or ina separate genus, The minute gradations observed by Hyatt, Waagen and all invertebrate palaeontologists, in the hard parts, (shells) of molluses, &c., are analogous: to the equally minute gradations observed by vertebrate palaeontologists.in the hard parts of rep-_ tiles and mammals. The mutations of Waagen may possibly, in fact, prove to. be identical with. the,‘ definite variations ”’ or | “ rectigradations ’ ’ observed by Osborn i in the teeth of mammals. For example, in the grinders of Eocene horses (see Plate TL., fig. 8; also fig. 9).in a lower horizon a cusp.is adumbrated in adows form, in a slightly higher; horizon it is visible, in.a, still higher horizon it is full-grown; and we honour this final stage by assign- ing to the animal,which-bears it amew specific name:; When a number of such characters.accumulate, we further honour them by assigning a new generic name. This is exactly the nomen- clature system laid down by. Owen, Cope, Marsh and others, ‘although established without any understanding, of the law of mutation.. But besides the innumerable. characters, which are visible and, “measurable, there, are probably. thousands which we cannot measure or which have not. been discovered, . since every part of the organism enjoys its gradual and, independent _eyolution, In the face of the continuous, series of characters and types revealed by palaeontology, the Linnaean, terminology, the individual order of development and the ancestral order of evolution... The mutual relations of palaeontology and, embryo- logy, and comparative anatomy .as means of determining the ancestry, of animals are most. interesting. _ In tracing. the phylogeny, or ancestral history of organs, palaeontology affords the, only absolute criterion on the successive evolution of organs in time.as well as of (progressive) evolution zm. form. From comparative anatomy alone it is possible to arrange a series. of living. forms which, although. structurally a convincing array because; placed in a graded..series, may. be, nevertheless, in an order inverse,to. that. of, the actual, historical succession... The most marked.case of such inversion, in, comparative anatomy is that of Carl Gegenbaur (1826-1903), who. in arranging the fins of fishes.in: support of his theory that the fin of the Australian lung-fish (Ceratodus) was the most primitive (or Archipterygium), placed,.as the primordial type.a fin which palaeontology has proved to be one, of the latest: types.if not the last. It is equally true that palaeontological evidence has frequently failed where we most sorely needed it. _The student must therefore resort to what, maybe called a tripod of evidence, derived from the available facts of embryology, comparative, anatomy. and peleobisiegy. r oe THE PALAEONTOLOGIST AS HISTORIAN’ The modes of change among animals, and methods ,of anaiysing them.—As historian. the palaeontologist always, has, before him as. one, of, his, most, fascinating problems, phylogeny, . or the restoration .of the great tree of, animal, descent... Were the geologic record complete he, would be able to trace the ancestry of, man, and_of, all, other animals: back to, their, very. beginnings ''PALAEONTOLOGY in the ‘primordial protoplasm: Dealing with interrupted avidencé! however, it becomes necessary to exercise the closest analysis and synthesis as part of his general art asa restorer. The most fundamental distinction in analysis is that’ which must be made between homogeny, or true hereditary resemblance, and those multiple forms’ of adaptive’ resemblance’ which’ are variously known as ‘cases of “analogy,” parallelism,” ‘‘ con- vergence ” and “‘ homoplasy.” Of ‘these two kinds of genetic. and adaptive resemblance, homogeny ‘is the warp composed of the. vertical, ‘hereditary strands, which connect animals with | their ancestors, and their successors, while analogy.is the woof, composed of the horizontal strands which tie animals together | by their superficial xesemblances, , This wide distinction between similarity of descent. and similarity of adaptation applies to_ every organ,‘to all groups of organs, to animals as a whole, and | to all’ groups of animals. It is the old distinction between | homology and analogy on a grand scale. Analogy, in its power of transforming unlike and unrelated | animals or unlike and unrelated parts of animals into likeness, | has done such miracles that the inference of kinship is often | almost irresistible. During the past century it was and even now is the very “ will-o’-the-wisp ” of evolution, always tending to lead the phylogenist astray. It is the first characteristic of analogy that it is superficial. (After a drawing by Charles R. Knight, made under the direction of Professor Osborn.) Fic. 10.—Analogous or conyergent evolution in; Fish, Reptile and Mammal. The external similarity in the fore paddle and back fin of these three marine animals is absolute, although they are totally unrelated to each other, and have’ a’ totally different) internal ‘or skeletal structure. It is one of the most striking cases known of the law of analagous evolution. A, Shark (Lamna cornubica), with long lobe of tail upturned. B, Ichthyosaur (Ichthyosaurus quadricissus), with fin-like paddles, long lobe of tail down-turned. C, Dolphin (Sotalia fluviatilis), with horizontal tail, fin or fluke. and the dolphin (fig. 10) superficially resemble each other, but if the outer form be removed this resemblance proves to be a mere veneer of adaptation, because their internal skeletal parts are as radically different as are their genetic relations, founded on heredity. Analogy’ also produces equally remarkable internal or skeletal transformations. The ingentity of nature, however, in adapting animals is not infinite, because the same devices are repeatedly employed by her to accomplish the same adaptive ends whether in fishes, reptiles, birds or mammals; thus she has repeated herself at least twenty-four times in the evolution of long-snouted’ rapacious swimming types of animals. “The grandest application of analogy is that observed in the adapta- tions of groups of animals evolving on different’ continents,’ by which their various divisions tend to mimic’ those on other continents. Thus the collective fauna of ancient South America Thus the shark, the ichthyosaur, he £1 | step there have been established in palaeontology a number of O87 mimics the independently evolved: collective: fauna of North America, the collective fauna’of modern Australia mimics the collective ‘fauna of ithe Lower Eocene of) North: America. Exactly the same: principles ‘have: developed on even a vaster scale among the Invertebrata... Among) the: ammonites of the Jurassic: and Cretaceous, periods .types..cccur, which .in their external, appearance so closely resemble each other that they could be taken for members of a single series, and not infrequently have been taken for species of the same genus and even for the same species; but their early stages of development and, in fact, their entire individual history prove them to be distinct and not infrequently to belong to widely separated genetic series. Homogeny; .in'icontrast, the ‘ special homology ” of Owen, is the supreme test of kinship or of hereditary relationship, and thus the basis of all sound-reasoning in phylogeny. ‘The two joints of the thumb, for example, are homogenous throughout the whole series of the pentadactylate, or five-fingered animals, from the most primitive amphibian. tooman. 0” I-rrode The conclusion’ is that, the sum’ of homogenous parts, which may be similar or dissimilar in external form according to their similarity or diversity of function, and the recognition of former similarities of adaptation (see below) are the true bases for the critical determination of kinship and phylogeny. Adaptation andthe Independent Evolution. of Paris.—Step by laws relating to the evolution of the parts of animals which closely coincide with similar laws discovered by zoologists. All } are contained in the bread generalization that every part of an animal, however minute, has its separate and independent basis in the hereditary substance of the germ cells from which it is derived and may.enjoy consequently a.separate and independent history. The consequences of this principle when applied to the adaptations of animals bring us to the very antithesis of Cuvier’s supposed “‘law of correlation,” for we find that, while the end results of adaptation are such that all parts of an animal conspire to make the whole adaptive, there is no fixed correlation either in the form or rate of development of parts, and that itis there- fore impossible for the palaeontologist to predict the anatomy of an unknown animal from one of its parts only, unless the animal happens to belong to a type generally familiar. For example, among the land vertebrates the feet (associated with the structure of the limbs and trunk) may take one of many lines of adaptation to different media or habitat; either aquatic, terrestrial, arboreal or aerial; while the teeth (associated with the structure of the skull and jaws) also may take one of many lines of adaptation to different kinds of food, whether herbivorous, insectivorous or carnivorous. ‘Through this independent adaptation of different parts to their specific ends there have arisen among vertebrates an almost unlimited number of combinations of foot and tooth structure, the possibilities of which are illustrated in the accom- panying diagram (see fig. 11; also Plate IIT., fig. 8): As instances of such combinations, some of the (probably herbivorous) Eocene monkeys with arboreal limbs have teeth so difficult to distinguish from those of the herbivorous ground-living Eocene horses with cursorial limbs that at first in France and also in America they were both classed with the hoofed animals. Again, directly opposed to Cuvier’s principle, we have discovered carnivores with hoofs, such as° Mesonyx, and herbivores with’ sloth-like claws, such as Chalicotherium. This latter animal is closely related to one which Cuvier termed Pangolin gigantesque, and had he restored it according to his “ law of correlation ” he would have pictured a giant “scaly anteater,” a type as wide as the poles from the actual form of Chalicotherium, which in body, limbs and teeth is a modified ungulate herbivore, related remotely to the tapirs. In its claws alone does it resemble the giant sloths. This independence of adaptation applies to every detail of structure; the six cusps of a grinding tooth may all-evolve alike, or each may evolve independently and differently. Independent evolution of parts is well shown among invertebrates, where the shell of’ an ammonite, for example, may change markedly in form’ without a corresponding change in suture, or vice versa. ''588 Similarly, there is no correlation in the rate of evolution either of adjoining or of separated parts; the middle digit of the foot of the three-toed horse is accelerated in development, while the lateral digits on either side are retarded. Many examples might be cited among invertebrates also. ADAPTIVE TYPES OF LIMBS AND FEET VOLANT FOSSORIAL ARBOREAL ae Short-limbed, plantigrade, heii eal pentadactyl, unguicu- OR late Stem TERRESTRIAL NATATORIAL CURSORIAL Amphibious Digitigrade Aquatic Unguligrade ADAPTIVE TYPES OF TEETH OMNIVOROUS Grass Herb HERBIVOROUS4 Shrub Fish Fruit CARNIVOROUS Flesh Root Carrion MYRMECOPHAGOUS Dentition reduced Stem INSECTIVOROUS Law OF THE INDEPENDENT ADAPTIVE EVOLUTION OF Parts. Fic. -11.—Diagram, demonstrating that. there are an. indefinite number of combinations of various adaptive types of limbs. and feet with various adaptive types of teeth, and that there is no fixed law of correlation between the two series of adaptations. All..these. principles .are consistent with Francis Galton’s law of particulate inheritance in heredity, and with the modern doctrine of ‘‘ unity of characters ” held by students of Mendelian phenomena. Sudden versus Gradual Evolution of Parts—There is a broad and most interesting analogy between the evolution. of parts of animals and of groups of animals studied as a whole. Thus we observe persistent organs and. persistent. types. of animals, analogous organs and analogous types of animals, and this analogy applies still further to the rival and more or less contra- dictory hypotheses. of the sudden as: distinguished from the gradual appearance of new parts or organs of animals, and the sudden appearance.of new types of animals. The first exponent of the theory of sudden appearance of new; parts and new types, to our knowledge, was Geoffroy St Hilaire, who suggested saltatory evolution through the direct.action of the, environ- ment on development, as explaining the abrupt transitions in the Mesozoic: Crocodilia and the origin of the birds from the reptiles. Waagen’s law of mutation, or the appearance of new parts or organs so gradually that they can be perceived only) by following them through successive geologic time stages, appears to be directly contradictory to the saltation principle; it is cer- tainly one of the most firmly established principles of palae- ontology, and it constitutes the contribution par excellence of this branch of zoology to the law of evolution, since it is obvious that it could not possibly have been deduced from comparison of PALAEONTOLOGY living animals but only through the long perspective gained by comparison of animals succeeding each other in time... The essence of Waagen’s law is orthogenesis, or evolution in a definite direction, and, if there does exist an internal hereditary. principle controlling such orthogenetic evolution, there doesnot appear to be any essential contradiction between its gradual operation in the “mutations of Waagen.”’.and its,occasional hurried operation in the “ mutations of de Vries,” which are by their definition discontinuous or saltatory (Osborn, 1907). VII.—Mopes oF CHANGE IN ANIMALS AS A WHOLE OR IN GROUPS. OF ANIMALS, AND’ METHODS OF’ “ANALYSING THEM. 1. Origin from Primitive or Stem Forms.—As already observed, the same principles apply to groups of animals as to organs and groups of organs; an organ originates in a primitive and un- specialized stage, a group of animals originates in a primitive orstemform. It was early perceived by Huxley, Cope and many others that Cuvier’s broad belief in a universal progression was erroneous, and there developed the distinction between “ per- sistent primitive types” (Huxley) and “ progressive types.” The theoretical existence of primitive or stem forms was clearly perceived by Darwin, but the steps by which the stem form might be restored were first clearly enunciated by Huxley in 1880 (“On the Application of Evolution to the Arrangement of the Vertebrata and more particularly of the Mammalia,” Scient. Mem. iv..457) namely, by sharp separation of the primary or stem characters from the secondary or adaptive characters in all the known descendants or branches of a theoretical original form. The sum of the primitive characters approximately restores the primitive form; and the gaps in palaeontological evidence are supplied by analysis of the available zoological, embryological and anatomical evidence. Thus Huxley, with true prophetic instinct, found that the sum of primitive characters of all the higher placental mammals points to a stem form of a generalized insectivore type, a prophecy which has been fully confirmed by the latest research» On the other hand, Huxley’s summation of the primitive characters of all the mammals led him to an amphibian stem type, a prophecy which has proved faulty because based on erroneous analysis and comparison. More or less ‘independently, Huxley, Kowalevsky and Cope restored the stem ancestor of the hoofed animals;.or ungulates, a restoration which has been.nearly fulfilled by the discovery, in 1873, of the generalized type Phenacodus of northern Wyoming. Similar anticipations and verifications among the invertebrates have been made by Hyatt, Beecher, Jackson and others. In certain cases the character stem forms actually survive in unspecialized types. Thus the analysis of George Baur of the ancestral form of the lizards, mosasaurs, dinosaurs, crocodiles and phytosaurs led both to the generalized Palaeohatteria of the Permian and indirectly to the surviving Tuatera lizard of New Zealand. 2. Adaptations to Alternations of Habitat: Law of Irreversi- bility of Evolution.—In the long vicissitudes of time and proces- sion of continental changes, animals have been subjected to alternations of habitat either through their own migrations or through the “‘ migration of the environment itself,’ to employ Van den Broeck’s epigrammatic description.of the profound and sometimes sudden environmental changes. which may take place in a single locality..The traces of alternations of adaptations corresponding to these alternations.of habitat are recorded both in palaeontology and anatomy, although often after the obscure analogy of the earlier and later writings of a palimpsest.. Huxley in 1880 briefly suggested the arboreal origin, or primordial tree- habitat of all the marsupials, a suggestion abundantly confirmed by. the. detailed. studies of Dollo and of Bensley, according to which we may imagine the marsupials to have passed through (x) a former terrestrial phase, followed by (2) a primary arboreal phase—illustrated in the tree phalangers—followed by (3)..a secondary terrestrial phase—illustrated in the kangaroos and wallabies—followed by (4) a secondary arboreal phase—illus- trated in the tree kangaroos. Louis Dollo especially has '' PALAEONTOLOGY contributed most brilliant discussions of the theory of alter- nations of habitat as applied to the interpretation of the anatomy of the marsupials, of many kinds of fishes, of such reptiles as the herbivorous dinosaurs of the Upper Cretaceous. He has applied the theory with especial ingenuity to the interpretation of the circular bony plates in the carapace of the aberrant leather-back sea-turtles (Sphargidae) by prefacing an initial land phase, in which the typical armature of land tortoises was acquired, a first marine or pelagic phase, in which this armature was lost, a third littoral or seashore phase, in which a new poly- gonal armature was acquired, and a fourth resumed or secondary marine phase, in which this polygonal armature began to degenerate. Each of these alternate life phases may leave some profound modification, which is partially obscured but seldom wholly lost; thus the tracing of the evidences of former adaptations is of great importance in phylogenetic study. A very important evolutionary principle is that in such secondary returns to primary phases lost organs are never recovered, but new organs are acquired; hence the force of Dollo’s dictum that evolution is irreversible from the point of view of structure, while frequently reversible, or recurrent, in point of view of the conditions of environment and adaptation. 3. Adaptive Radiations of Groups, Continental and Local.— Starting with the stem forms the descendants of which have passed through either persistent or changed habitats, we reach the underlying idea of the branching law of Lamarck or the law of divergence of Darwin, and find it perhaps most clearly ex- pressed in the words ‘‘adaptive radiation’”’ (Osborn), which convey the idea of radii in many directions. Among extinct Tertiary mammals we can actually trace the giving off of these radii in all directions, for taking advantage of every possibility to secure food, to escape enemies and to reproduce kind; further, among such well-known quadrupeds as the horses, rhinoceroses and titanotheres, the modifications involved in these radiations can be clearly traced. Thus the history of continental life presents a picture of contemporaneous radiations in different parts of the world and of a succession of radiations in the same parts. We observe the contemporaneous and largely independent radiations of the hooféd animals in South America, in Africa and in the great ancient continent comprising Europe, Asia and North America; we observe the Cretaceous radiation of hoofed animals in the northern hemisphere, followed by a second radiation of hoofed animals in the same region, in some cases one surviving spur of an old radiation becoming the centre of a new one. As a rule, the larger the geographic theatre the grander the radia- tion. Successive discoveries have revealed certain grand centres, such as (1) the marsupial radiation of Australia, (2) the little- known Cretaceous radiation of placental mammals in the northern hemisphere, which was probably connected in part with the peopling of South America, (3) the Tertiary placental radiation in the northern hemisphere, partly connected with Africa, (4) the main Tertiary radiation in South America. Each of these radiations produced a greater or less number of analogous groups, and while originally independent the animals thus evolving as autochthonous types finally mingled together as migrant or invading types. We are thus working out gradually the separate contributions of the land masses of North America, South America, Europe, Asia, Africa, and of Australia to the mammalian fauna of the world, a result which can be obtained through palaeontology only. 4. Adaptive Local Radiation.—On a smaller scale are the local adaptive radiations which occur through segregation of habit and local isolation in the same general geographic region wherever physiographic and climatic differences are sufficient to produce local differences in food supply or other local factors of change. This local divergence may proceed as rapidly as through wide geographical segregation or isolation. This principle has been demonstrated recently among Tertiary rhinoceroses and titano- theres, in which remains of four or five genetic series in the same geologic deposits have been discovered. We have proof that in the Upper Miocene of Colorado there existed a forest-living horse, 589 or more persistent primitive type, which was contemporaneous with and is found in the same deposits with the plains-living horse (Neohipparion) of the most advanced or specialized desert type (see Plate IV., figs. 12, 13, 14, 15). In times of drought these animals undoubtedly resorted to the same water-courses for drink, and thus their fossilized remains are found associated. 5. The Law of Polyphyletic Evolution. The Sequence of Phyla or Genetic Series.—There results from continental and. local adaptive radiations the presence in the same geographical region of numerous distinct lines in a given group of animals. The polyphyletic law was early demonstrated among invertebrates by Neumayr (1889) when he showed that the ammonite genus Phylloceras follows not one but five distinct lines of evolu- tion of unequal duration. The brachiopods, generally classed collectively as Spirifer mucronatus, follow at least five distinct lines of evolution in the Middle Devonian of North America, while more than twenty divergent lines have been ebserved by Grabau among the species of the gastropod genus Fusus in Tertiary and recent times. Vertebrate palaeontologists were slow to grasp this principle; while the early speculative phylo- genies of the horse of Huxley and Marsh, for example, were mostly displayed monophyletically, or in single lines of descent, it is now recognized that the horses which were placed by Marsh in a single series are really to be ranged in a great number of contemporaneous but separate series, each but partially known, and that the direct phylum which leads to the modern horse has become a matter of far more difficult search. As early as 1862 Gaudry set forth this very polyphyletic principle in his tabular phylogenies, but failed to carry it to its logical application. It is now applied throughout the Vertebrata of both Mesozoic and Cenozoic times. Among marine Mesozoic reptiles, each of the groups broadly known as ichthyosaurs, plesiosaurs, mosasaurs and crocodiles were polyphyletic in a marked degree. Among land animals striking illustrations of this local polyphyletic law are found in the existence of seven or eight contemporary series of rhinoceroses, five or six contemporary series of horses, and an equally numerous contemporary series of American Miocene and Pliocene camels; in short, the polyphyletic condition ‘is the rule rather than the exception. It is displayed to-day among the antelopes and to a limited degree among the zebras and rhinoceroses of Africa, a continent which exhibits a survival of the Miocene and Pliocene. conditions of the northern hemisphere. 6. Development of Analogous Progressive and) Retrogressive Groups.—Because of the repetition of analogous physiographic and climatic conditions in regions widely separated both in time and in space, we discover that continental and local adaptive radiations result in’ the creation of analogous groups of radii among all the vertebrates and invertebrates. Illustrations of this law were set forth by Cope as early as 1861 (see ‘‘ Origin of Genera,” reprinted in the Origin of the Fittest, pp. 95-106) in pointing out the extraordinary parallelisms between unrelated groups of amphibians, reptiles and mammals. In the Jurassic period there were no less than’ six orders of, reptiles which independently abandoned terrestrial life and acquired more or less perfect adaptation to sea life. Nature, limited in) her resources for adaptation, fashioned so many of these animals in like form that we have learned. only recently to: distinguish similarities cf analogous habit from the similitudes of real kinship. From whatever order of Mammalia or Reptilia an animal may be derived, prolonged aquatic adaptation will model its outer, and finally its inner, structure according to certain advantageous designs. The requirements of an elongate body moving through the resistant medium of water are met by the evolution of similar entrant and exit curves, and the bodies of most swiftly moving aquatic animals evolve into forms resembling the hulls of modern sailing yachts (Bashford Dean). We owe especially to Willy Kiikenthal, Eberhard Fraas, S.W. Williston and R. C. Osburn a summary of those modifications of form to which aquatic life invariably leads. The law of analogy also operates in retrogression. A. Smith Woodward has observed that the decline of many groups of ''590 fishes is heralded by the tendency to assume elongate and finally eel-shaped forms, ‘as seen independently, for example, among the declining: Acanthodians or: palaeozoic sharks, among the modern crossopterygian Polypterus and Calamoichthys of the Nile, in the modern dipneustan Lepidosiren and Protopterus, in the Triassic chondrostean Belonorhynchus, as well as in the bow-fin (Amia) and the garpike (Lepidosteus). Among invertebrates similar analogous groups also develop. This is especially marked in: retrogressive, though ‘also -well- known in progressive series. The loss of the power to coil, observed in the terminals of many declining series of gastropods from the Cambrian tothe present time, and the similar loss of power among Natiloidea and ‘Ammonoidea of many: genetic series, as well as the ostraean form assumed by various declining series of pelecypods and by some: brachiopods, may: bé cited’ as examples. 7. Periods of Graduai Evolution of Groups:—It is certainly a very ‘striking fact that wherever we have been able to trace genetic series, either of invertebrates or vertebrates, in closely sequent geological horizons, or life zones, we find strong proof of evolution through extremely gradual mutation simultaneously. affecting many parts of each organism, as set forth above. This proof has been reached) quite independently by a very large number of observers studying a still greater variety of animais. Such diverse organisms as brachiopods, ammonites, horses and rhinoceroses absolutely conform to this law in all those rare localities where we have been able to observe closely sequent stages. The inference is almost irresistible that the law of gradual transformation through minute continuous change is by far the most universal; but many palaeontologists as well as zoologists and botanists hold a contrary opinion. 8. Periods of Rapid Evolution vy Groups.—The above law of gradual evolution is perfectly consistent with a second principle, namely, that at certain times evolution is much more rapid than at others, and that organisms are accelerated or retarded in development in a manner broadly analogous to the acceleration or retardation of separate organs. Thus H.S. Williams observes (Geological Biology, p. 268) that. the evolution of those funda- mental characters which mark differences between separate classes, orders, sub-orders, and even families of organisms, took place in relatively short periods of time. Among the brachiopods the chief expansion of each type is at a relatively early period in their life-history. Hyatt (1883) observed of the ammonites that each group originated suddenly and spread out with great rapidity. Depéret notes that the genus Newmayria, an ammonite of the Kimmeridgian, suddenly branches out into an “explosion” of forms. Depéret also observes the contrast between periods of quiescence and limited variability and periods of sudden efflorescence. A, Smith Woodward (“Relations of Palaeontology to Biology,” Annals and Mag. Natural Hist., 1906, p.317) notes that the fundamental advances in the growth of fish life have always been sudden, beginning with excessive vigour at the end of long periods of apparent stagnation; while each advance has been marked by the fixed and definite acquisition of some new anatomical character or “ expression point,” a term first used by Cope. One of the causes of these sudden advances is un- doubtedly to be found in the acquisition of a new and extremely useful character. Thus the perfect jaw and the perfect pair of lateral fins when first acquired among the fishes favoured a very rapid and for a time unchecked development. It by no means follows, however, from this incontrovertible evidence that the acquisition either of the jaw or of the lateral fins had not been in itself an extremely gradual process. Thus both invertebrate and vertebrate palaeontologists have reached independently the conclusion that the evolution of groups is not continuously ata uniform rate, but that there are, especially in the beginnings of new phyla or at the time of acquisition of new organs, sudden variations in the rate of evolu- tion which have been termed variously “ rhythmic,” “pulsating,” “efflorescent,” “intermittent ” and even “ explosive” (Depéret). This varying rate of evolution has (illogically, we believe) been compared with and advanced in support of the ‘mutation law confused with that for the law of rapid efflorescence PALAEONTOLOGY of De Vries,”’or the theory of saltatory evolution, which we may next consider. i 9. Hypothesis of the Sudden Appearance of News Parts or Organs.—The rarity of really continuous series has naturally led palaeontologists to support the hypothesis of brusque tran- sitions: of structure... As we have seen, this hypothesis was fathered by Geofiroy St Hilaire in 1830 from his studies of Meso- zoic Crocodilia, was sustained by Haldemann, and quite recently has been revived by: such eminent palaeontologists: as Louis Dollo and A. Smith Woodward. The evidence for it is not to be of groups just considered. It. should ‘be :remembered: that palaeontology ‘is the most unfavourable field of all for observation and demon- stration of sudden saltations or mutations of character, because of the limited materials available for comparison and the rarity of genetic series. It should be borne in mind, first, that wherever a new animal suddenly appears or a ‘new: character suddenly arises in a fossil horizon we must consider whether such appear- ance may be due to 'the non-discovery of transitional links with older forms, or to the sudden invasion of a new type or new organ which has gradually evolved elsewhere, The rapid variation of certain groups of animals or the acceleration of certain organs is also not evidence of the sudden appearance of new adaptive characters. Such sudden. appearances may be demonstrated possibly in zoology and embryology but never can be demon- strated by palacontology, because of the incompleteness of the geological record. 10. Decline or Senescence of Groups.—Periods of gradual evolution and.of efflorescence may be followed by stationary or senescent conditions. In his history of the Arietidae Hyatt points out that toward the close of the Cretaceous this entire group of ammonites appears to have been affected with some malady; the unrolled forms multiply, the septa are simplified, the ornamentation becomes heavy, thick, and finally disappears in the aduit; the entire group ends by dying ovt and leaving no descendants. This is not due to environmental conditions solely, because senescent branches of normal progressive groups are found in all geologic horizons, beginning, for gastropods, in the Lower Cambrian. Among the ammonites the loss of power to coil the shell is one feature of racial old age, and in others old age is accompanied by closer coiling and loss of surface orna- mentation, such as spines, ribs, spirals; while in other forms an arresting of variability precedes extinction. Thus Williams has observed that if we find a species breeding perfectly true we can conceive it) to have reached the end of its racial life period. Brocchi and Daniel Rosa (1899) have developed the hypothesis of the progressive reduction of variability. Such decline is by no means a universal law of life, however, because among many of the continental vertebrates at least we observe extinctions repeatedly occurring during the expression of maximum varia- bility. Whereas among many ammonites and gastropods smooth ness of the shell, following upon an ornamental youthful condition, is generally a symptom of decline, among many other invertebrates and vertebrates, as C. E. Beecher (1856=10905) has pointed out (1898), many animals possessing hard parts tend toward the close of their racial history to produce a superfluity of dead matter, which accumulates in the form of spines among invertebrates, and of horns among the land vertebrates, reaching a maximum when the animals are really on the down-grade of development. 11. The Extinction of Groups—We have seen that different lines vary in vitality and in longevity, that from the éarliest times senescent branches are given off, that different lines vary in the rate of evolution, that extinction is often heralded by symptoms of racial old age, which, however, vary widely in different groups. In general we find an analogy between the development of groups and of organs; we discover that each phyletic branch of certain. organisms traverses a geologic career comparable to the life of an individual, that we may often distinguish, especially amung invertebrates, a phase of youth, a phase of maturity, a phase of senility or degeneration fore- shadowing the extinction of a type. ''PALAEOSPONDYLUS Internal causes of extinction are to be found in exaggeration of body size, in the hypertrophy or over-specialization of certain organs, in the irreversibility of evolution, and possibly, although this has not been demonstrated, in a progressive reduction of variability. In a full analysis of this problem of internal and external causes in relation to the Tertiary Mammalia, H. F. Osborn (‘‘ Causes of Extinction of the Mammalia,” Amer. Natur- alist, 1906, pp. 769-795, 829-859) finds that foremost in the long series of causes which lead to extinction are the grander environ- mental changes, such as physiographic changes, diminished or contracted land areas, substitution of insular for continental conditions; changes of climate and secular lowering of temperature accompanied by deforestation and checking of the food supply; changes influencing the mating period as well as fertility; changes causing increased humidity, which in turn favours enemies among insect life. Similarly secular elevations of temperature, either accompanied by moisture or desiccation, by increasing droughts or by disturbance of the balance of nature, have been followed. by.great waves of extinction of the Mammalia... In the sphere of living environment, the varied evolution of plant life, the periods of forestation and deforestation, the introduction of deleterious plants simultaneously with harsh conditions of life and enforced. migration, as. well as of mechanically dangerous plants, are among the well-ascertained causes of diminution and extinction: .The evolution of insect life in driving animals from feeding ranges and in the spread of disease probably has been a prime cause of extinction. Food competition among mammals, especially intensified on islands, and theintroduction of Carnivora constitute another class of causes. Great waves of extinction have followed the long periods of the slow evolution of relatively inadaptive types of tooth and foot structure, as first demon- strated by Waldemar Kowalevsky; thus mammals are repeatedly observed in a cul-de-sac of structure from which there is no escape in an,adaptive direction.. Among still other causes are great bulk, which proves fatal under certain new conditions; rela- tively slow breeding; extreme specialization and development of dominant organs, such as horns and tusks, on which for a time selection centres to the detriment of more: useful characters. Little proof is. afforded among the mammals of extinction through arrested evolution or through the limiting of variation, although such laws undoubtedly .exist.. One of the chief deductions is that there are special dangers in numerical diminu- tion of herds, which.may arise from a. chief or original cause and be followed by.a conspiracy of other causes which are cumu- lative in effect. . This survey. of the phenomena of extinction. in one great class of animals certainly establishes the existence of an almost infinite variety of causes, some of which are internal, some external in origin, operating on animals of different kinds. VIII.— UNDERLYING BIOLOGICAL PRINCIPLES AS THEY APPEAR TO THE PALAEONTOLOGIST It follows from the above brief summary that palaeontology affords a distinct and highly suggestive field of purely biological research; that is, of the causes of evolution underlying the obsery- able modes which we have been describing. The net result of observation is not favourable to the essentially Darwinian view that the adaptive arises out of the fortuitous by selection; but is rather favourable to the hypothesis of the existence of some quite unknown intrinsic law of life which we are at present totally unable to comprehend or even conceive. We have shown that the direct observation of the origin of new characters in palaeontology brings them within that domain of natural law and order to which the evolution of the physical universe con- forms. The nature of this law, which, upon the whole, appears to be purposive or teleological in its operations, is altogether a mystery which may or may not be illumined by future research. In other words, the origin, or first appearance of new characters, which is the essence of evolution, is an orderly process so far as the vertebrate and invertebrate palaeontologist observes. it. The selection of organisms through the crucial test of fitness and the shaping. of the organic world is an orderly process when contemplated.on a grand scale, but. of, another, kind; here. the 5QF test of fitness is supreme... The only inkling of possible underlying principles in this orderly process is that there appears to be in respect to certain characters a potentiality or a predisposition through hereditary kinship to evolve in certain definite directions. Yet there is strong evidence against the existence of any law. in the nature of an internal perfecting tendency which would operate independently of external conditions... In other , words, a balance appears to be always sustained between the internal (hereditary and ontogenetic) and the external (environmental and selectional) factors of evolution. kiran BIBLIOGRAPHY.—Among the older works on the history of palaeontology are the treatises of Giovanni Battista Brocchi (1772- 1826), Conchiologia fossile Subappenina... Disc. sui progressi dello studio... 1843 (Milan); of Etienne Jules d’Archiac, Histoire du progres de la géologie de 1834 a 1862 (Paris, Soc. Géol. de France, 1847-1860); of Charles Lyell in his Principles of Geology. A clear narrative of the work of many of the earlier contributors is found in Founders of Geology, by Sir Archibald Geikie (London, 1897— 1905). The most comprehensive and up-to-date reference work on the history of geology and palaeontology is Geschichte der Geologic und Paldontologie, by Karl Alfred von Zittel (Munich and Leipzig, 1899), the final life-work of this great authority, translated ,into English in part by Maria M. Ogilvie-Gordon, entitled ‘‘ History of Geology and Palaeontology to the end of the 19th Century.” The succession of life from the earliest times as it was known at the close of the last century was treated by the same author in his Handbuch der Paléontologie (5 vols., Munich and Leipzig, 1876-1893). Abbre- viated editions of this work have appeared from the author, Grund- ziige der Paldontologie (Palaeozoologie) (Munich and Leipzig, 1895, 2nd ed., 1903), and in English form in Charles’ R. Eastman’s Texi- Book of Palaeontology (1900-1902). A classic. but unfinished work describing the methods of invertebrate palaeontolegy is Die Stdmme des Thierreichs: (Vienna, 1889), by Melchior Neumayr. In France admirable recent works are Elémenis de Paléontologie, by Felix Bernard (Paris, 1895), and the still more recent philosophical treatise by Charles Depéret, Les Transformations du monde animal (Paris, 1907). Huxley’s researches, and especially his share in the development of the philosophy of palaeontology, will be found in his essays, The Scientific Memoirs of Thomas Henry Huxley (4 vols., London, 1898-1902). The whole subject is treated systematically in Nicholson and Lydekker’s A Manual of Palaeontology (2 vols., Edinburgh and London, 1889), and A. Smith Woodward’s Outlines of Vertebrate Palaeontology (Cambridge, 1898). Among American contributions to vertebrate palaeontology, the development of Cope’s theories is to be found in the volumes of his collected essays, The Origin of the Fittest (New York, 1887), and The Primary Factors of Organic Evolution (Chicago, 1896). A brief summary of the rise of vertebrate palaeontology is found in the address of O. Marsh, entitled ‘‘ History and Methods of Palaeonto- logical Discovery ’’ (American Association for the Advancement of Science, 1879). The chief presentations of the methods of the American: school of invertebrate palaeontologists are to be found in A. Hyatt’s great memoir ‘‘ Genesis of the Arietidae ’’ (Smithsonian Contr. to Knowledge, 673, 1889), in Hyatt’s “ Phylogeny of an Acquired Characteristic’’ (Philosophical Soc. Proc., vol. xxxii. 1894), and in Geological Biology, by H.S. Williams (New York, 1895). In preparing the present article the author has drawn freely on his own addresses: see H. F.. Osborn, “‘ The Rise of the Mammalia in North America’ (Proc. Amer. Assn. Adv. Science, vol. xlii., 1893), ‘‘ Ten Years’ Progress in the Mammalian Palaeontology of North America’ (Comptes rendus du 6° Congres intern. de zoologie, session de Bern, 1904), ‘‘ The Present Problems of Palaeontology ”’ (Address before Section of Zool. International Congress of Arts and Science, St Louis, Sept. 1904), ‘‘ The Causes of Extinction of Mammalia ”’ (Amer. Naturalist, x\. 769-795, 829-859, AE 0) PALAEOSPONDYLUS, a small fish-like organism, of which the skeleton is found fossil in the Middle Old Red Sandstone ki . Kau 5 p Beeler teary ee t J SESS Sao. , ie — From British Museum Guide to Fossil Reptiles and Fishes, by ‘ ; permission of the Trustees. Palaeospondylus gunni, restored by Dr R. H. ‘Traquair. (Nearly twice nat. size.) - of Achanarras, near Thurso, Caithness. It was thus named (Gr, ancient vertebra) by Dr R. H. Traquair in 1890, in allusion to its. well-developed. vertebral rings; and, its structure was ''594 studied in detail in 1903 by Professor and Miss Sollas, who succeeded in making enlarged models of the fossil in wax. The skeleton as preserved is carbonized, and indicates an ¢el- shaped animal from 3 to 5 cm., in length. The skull, which must have consisted of hardened cartilage, exhibits pairs of nasal and auditory capsules, with a gill-apparatus below its hinder part, but no indications of ordinary jaws. The anterior opening of the brain-case is surrounded by a ring of hard cirri. A pair of “ post-branchial plates ” projects backwards from the head. The vertebral axis shows a series of broad rings, with distinct neural arches, but no ribs. Towards the end of the body both neural and haemal arches are continued into forked radial cartilages, which support a median fin. ‘There are no traces either of paired fins or of dermal armour: The affinities of Palaeospondylus are doubtful, but it is probably related to the. contemporaneous armoured Ostracoderms. REFERENCES.—R. H. Traquair, paper in Proc. Roy. Phys. Soc. Edin., xii. 312, (1894); W. J. Sollas and I. B. J. Sollas, paper in Phil. Trans. Roy. Soc. (1903 B.). (A. S. Wo.) PALAEOTHERIUM (i.e. ancient animal), a name applied by Cuvier to the remains of ungulate mammals recalling tapirs in general appearance, from the Lower Oligocene gypsum quarries of Paris. These were the first indications of the (From the Paris gypsum.) Restoration of Palaeotherium magnum. (About } nat. size.) occurrence in the fossil state of perissodactyle ungulates allied to the horse, although it was long before the relationship was recognized. The palaeotheres, which range in size from that of a pig to that of a small rhinoceros, are now regarded as repre- senting a family, Palaeothertidae, nearly related to the horse- tribe, and having, in fact, probably originated from the same ancestral stock, namely, Hyracotherium of the Lower Eocene (see EquipaE). The connecting link with H-yracotherium was formed by Pachynolophus (Propalacotherium), and the line apparently terminated in Paloplotherium, which is also Oligocene. Representatives of the family occur in many parts of Europe, but the typical genus is unknown in North America, where, however, other forms occur. Although palaeotheres resemble tapirs in general appearance, they differ in having only three toes on the fore as well as on the hind foot. The dentition normally comprises the typical series of 44 teeth, although in some instances the first premolar is wanting. The cheek-teeth are short-crowned, generally with no cement, the upper molars having a W-shaped outer wall, from which proceed two oblique transverse crests, while the lower ones carry two crescents. Unlike the early horses, the later premolars are as complex as the molars; and although there is a well-marked gap between the canine and the premolars, there is only a very short one between the former and the incisors. The orbit is completely open behind. In other respects the palaeo- theres resemble the ancestral horses. They were, however, essentially marsh-dwelling animals, and exhibit no tendency to the cursorial type of limb so characteristic of the horse-line. They were, in fact, essentially inadaptive creatures, and hence rapidly died out. CRele) PALAEOZOIC ERA, in geology, the oldest of the great time divisions in which organic remains have left any clear record. The three broad divisions—Palaeozoic, Mesozoic, Cainozoic— PALAEOTHERIUM—PALAEPHATUS which are employed by geologists to mark three stages in the development of life on the earth, are based primarily upon the fossil contents of the strata which, at one point’ or another, have been continuously forming since the very earliest times. The precise line in the “‘record of the rocks” where the chronicle of the Palaeozoic era closes and that of the Mesozoic era opens— as in more recent historical documents—is a matter for editorial caprice. The early geologists took the most’ natural dividing lines that came within their knowledge, namely, the line of change in general petrological characters, ¢.g. the ““ Transition Series ” (Ubergangsgebirge), the name given to rocks approximately of Palaeozoic age by A. G. Werner because they exhibited a transi- tional stage between the older crystalline rocks and the younger non-crystalline; later in Germany these same rocks were said to have been formed in the “ Kohlenperiode ” by H: G: Bronn and others, while in England H. T. de la Beche classed them as a Carbonaceous and Greywacke group. Finally, the divisional time separating the Palaeozoic record from that of the Mesozoic was made to coincide with a great natural break or unconformity of the strata. This was the most obvious course, for where such a break occurred there would be the most marked differences between the fossils found below and those found above the physical discordance. The divisions in the fossil record having been thus established, they must for convenience remain, but their artificiality cannot be too strongly emphasized, for the broad stratigraphical gaps and lithological groups which made the divisions sharp and clear to the earlier geologists are proved to be absent in other regions, and fossils which were formerly deemed characteristic of the Palaeozoic era are found in some places to commingle with forms of strongly marked Mesozoic type. In short, the record is more nearly complete than was originally supposed. The.Palaeozoic or Primary era is divided into the following periods or epochs: Cambrian, Ordovician, Silurian, Devonian, Carboniferous and Permian. The fact that fossils found in the rocks of the three earlier epochs—Cambrian, Ordovician, Silurian —have features in common, as distinguished from those in the three later epochs has led certain authors to divide this era into an earlier, Protozoic (Proterozoic) and a later Deuterozoic time. The rocks of Palaeozoic age are mainly sandy and muddy sediments with a considerable development of limestone in places. These sediments have been altered to shales, slates, quartzites, &c., and frequently they are found in a highly meta- morphosed condition; in eastern North America, however, and in north-east Europe they still maintain their horizontality and primitive texture over large areas. The fossils of the earlier Palaeozoic rocks are characterized by the abundance of trilobites, graptolites, brachiopods, and the absence of all vertebrates except in the upper strata; the later rocks of the era are distinguished by the absence of graptolites, the gradual failing of the trilobites, the continued predominance of brachiopods and tabulate corals, the abundance of crinoids and the rapid development of placoderm and heterocercal ganoid fishes and amphibians. The land plants were all cryptogams, Lepidodendron, Sigillaria, followed by Conifers and Cycads. It is obvious from the advanced stage of development of the organisms found in the earliest of these Palaeozoic rocks that the beginnings of life must go much farther back, and indeed organic remains have been found in rocks older than the Cambrian; for convenience, therefore, the base of the Cambrian is usually placed at the zone of the trilobite Olenellus. (J.A.H.) PALAEPHATUS, the author of a small extant treatise, entitled Tlepi ’"Ariorwy (On “ Incredible Things”). It consists of a series of rationalizing explanations of Greek legends, without any attempt at arrangement or plan, and is probably an epitome, composed in the Byzantine age, of some larger work, perhaps the Avoes tév pvOiKds elpnuevwv, mentioned by Suidas as the work of a grammarian of Egypt or Athens. Suidas himself ascribes a Tlepi ’Arriorwv, in five books, to Palaephatus of Paros or Priene. The author was perhaps a contemporary of Euhemerus (3rd century B.c.). Suidas mentions two other writers of the name: (1) an epic poet of Athens, who lived before the time of ''PALAEONTOLOGY Prare I. Frc. 1.—An ichthyosaur (I. quadriscissus) containing in the body cavity the partially preserved skeletons of seven young, proving that the young of the animal developed within the maternal body and were brought forth alive; z.e. thet the ichthyosaur was a viviparous animal. (Specimen presented to the American Museum of Natural History by the Royal Museum of Stuttgart through Professor Eberhard Fraas.) Fic. 2.—A_ hypothetical pictorial restoration of the mother ichthyosaur accompanied by five of its newly born young, from the information furnished by actual fossils. (From a drawing by Charles R. Knight made under the direction of Professor Osborn.) Fe Fic. 3.—One of the most perfect of the many specimens discovered and prepared by Herr Bernard Hauff, and showing the extra- ordinary preservation of the epidermis of the ichthyosaur, which gives the complete contour of the body in silhouette, the out- lines of the paddles, of the remarkably fish-like tail, into the lower lobe of which the vertebral column extends, and the great integumentary dorsal fin. Materials for the Restoration of Ichthyosaurs.—This plate illustrates the exceptional opportunity afforded the palaeontologist through the remarkably preserved remains of Ichthyosaurs in the quarries of Holzmaden near Stuttgart, Wiirttemberg, excavated for many years by Herr Bernard Hauff. (Illustrations reproduced by permission from specimens in the American Museum of Natural History, New York.) XX. 580. ''Prats i. PALA EON TOLOGY Fic. 4SKELETON OF ALLOSAUR us oS Te 2 : vo é sas sii: Baad ! aa ‘eg mae iho, a vous hs cca Fic. 5—RESTORATION OF ALLOSAURUS. Materials for the Restoration of Dinosaurs.—Carniverous dinosaur (Allosaurus) of the Upper Jurassic period of North America, an ani- mal closely related to the Megalosaurus type of England. The skeleton (fig. 4) was found neatly complete in the beds of the Morrison formation, Upper Jurassic of central Wyoming, U.S.A. Near it was discovered the posterior portion of the skeleton of a giant herbivorous dinosaur (Brontosaurus Marsh). It was observed that ten of the caudal vertebre of the latter skeleton bore tooth marks and grooves corresponding exactly with the sharp pointed teeth in the jaw of the carnivorous dinosaur. This proved that the great herbivorous dinosaur had been preyed upon by its smaller carnivorous contemporary. Teeth of the carnivorous dinosaur scattered among the bores of the herbivorous dinosaur completed the line of circumstantial evidence. Upon this testimony the restoration (fig. 5) of the Megalosaur has been drawn by Charles R. Knight under the direction of Professor Osborn. (Originals reproduced by permission of the American Museum of Natural History.) al ''PALAEONTOLOGY Pirate IIL. This series of feet represents the evolutionary succession from the Eocene Hypohippus (1) to the modern Equus (6) seen in front and in side view. The top bone is the os calcts, or hock bone, to which the tendon Achilles is attached. The bottom bone is the terminal phalanx which is inserted in the heart of the hoof. Equus Modern caballus. horse. | Merychippus | 4 Sp. , Merychippus insignis ' (milkmolar).| yfiocene. Parahippus pawniensis. Pa The stages are as follows: . Hypohippus, Lower Eocene. 4. Protohippus, Upper Miocene. I 2, Mesohippus, Lower Oligocene. 5. Neohipparion, Upper Miocene. J 3 Parahippus, Lower Miocene. 6. Equus, Pleistocene and recent PPPS : : Mesohippus Oligocene : : ; ‘ intermedius. (White The evolution consists first in progressive in- river for- crease in size; second, in the acceleration of the mation). median digit and retardation of the lateral digits, the latter becoming more and more elevated from the ground until finally in Equus (6) they are the iateral splints, which in the embryonic condition Mesohippus have vestigial cartilages attached bairdi? representing the last traces of the lateral phalanges. Oligocene (White ~ river for- mation). Mesohippus batrdt. Middle Eocene Orohippus (Bridger Sp. for- mation). Eohippus Lovee SP (Wind river for- 2 mation). Eohippus (Wasatch Sp. for- | mation). | Fic. 8.—TEN STAGES IN THE EVOLU- 4 6 TION OF pe SECOND vee Fic. 7—LAW OF ACCELERATION AN 4 J f MOLAR TOOTH Tae 3 7 ee - A _ RETARDATION ILLUSTRATED IN SIDE, ARRANGED ACCORDING TO * EVOLUTION OF THE HIND FEET OF THE HORSE. CHOLOGIGAL, ERVBE. (From photos lent by the American Museum of Natural History.) (Nos. 1-9 from “American Equidae.’’) XX. 584. ''"AI SIVId Ze) > ee > cs © = eS © ‘ © @ K bohippus, a forcst- .—WNeohipparion, a plains-living horse with very slender limbs and lateral digits small and well raised from the ground, g ar view, showing adapted to a dry, hard soil. large lateral digits on the fore ‘and hind feet, adapted ‘animal from sinking into the soft soil. Laws of Local Adaptive aws 2 Radiation and _ Polyphy- letic Evolution, illustrated by two Upper Miocene Horses of the Plains Region of North Amer These horses are of the same geologic age (Upper Mio- cene) and were found in the same geographic region (South Dakota, U.S‘A:). One is supposed to have lived in the forests along the stream borders, and the other in the open plains. m (lilustrations reproduced by as permission of the Amert- A lass Hypoh Beans (From a drawing by c aaa R. Knight, can Museum of Natural F1G.15.—Restoration of : Nudebeiar on. made under the direction of Professor Osborn.) History, New York.) made under the direction of Profe ''PALAESTRA—PALAMCOTTAH Homer; (2) an historian of Abydus, an intimate friend of Aristotle. ; See edition by N. Festa, in Mythographi graect (1902), in the Teubner series, with valuable prolegomena supplementary to Intorno all’ opuscolo di. Palefato de incredibilibus (1890), by the same writer. PALAESTRA (Gr. 7aXatorpa), the name apparently applied by the Greeks to two kinds of places used for gymnastic and athletic exercises. In the one case it seems confined to the places where boys and youths received a general gymnastic training, in the other to a part of a gymnasium where the athletae, the competitors in the public games, were trained in wrestling (radatew, to wrestle) and boxing. The boys’ palaestrae were private institutions and generally bore the name of the manager or of the founder; thus at Athens there was a palaesira of Taureas (Plato, Charmides). The Romans used the terms gymnasium and palaestra indiscriminately for any place where gymnastic exercises were carried on. PALAFOX DE MENDOZA, JUAN DE (1600-1659), Spanish bishop, was born in Aragon. He. was appointed in 1839 bishop of Angelopolis (Puebla de los Angeles) in Mexico, and there honourably distinguished himself by his efforts to protect the natives from Spanish cruelty; forbidding any methods of con- version other than persuasion. In this he met with the uncom- promising hostility of the Jesuits, whom in 1647 he laid under an interdict. He twice, in 1647 and 1649, laid a formal complaint against them at Rome. The pope, however, refused to approve his censures, and all he could obtain was a brief from Innocent X. (May 14, 1648), commanding the Jesuits to respect the. episcopal jurisdiction. In 1653 the Jesuits succeeded in securing his trans- lation to the little see of Osma in Old Castile. In 1694 Charles II. of Spain petitioned for his canonization; but though this passed through the preliminary stages, securing for Palafox the title of “ Venerable,”’ it was ultimately defeated, under Pius VI., by the intervention of the Jesuits. See Antonio Gonzalez de Resende, Vie de Palafox (French trans., Paris, 1690). PALAFOX Y MELZI, JOSE DE (1780-1847), duke of Sara- gossa, was the youngest son of an old Aragonese family. Brought up at the Spanish court, he entered the guards at an early age, and in 1808 as asub-lieutenant accompanied Ferdinand to Bayonne; but after vainly attempting, in company with others, to secure Ferdinand’s escape, he fled to Spain, and after a short period of retirement placed himself at the head of the patriot movement in Aragon. He was proclaimed by the populace governor of Saragossa and captain-general of Aragon (May 25, 1808). Despite the want of money and of regular troops, he lost no time in declaring war against the French, who had already overrun the neighbouring provinces of Catalonia and Navarre, and soon afterwards the attack he had provoked began. Saragossa as a fortress was both antiquated in design and scantily provided with munitions and supplies, and the defences resisted but-a short time. But it was at that point that the real resistance began. A week’s street fighting made the assailants masters of half the town, but Palafox’s brother succeeded in forcing a passage into the city with 3000 troops. Stimulated by the appeals of Palafox and of the fierce and resolute demagogues who ruled the mob, the inhabitants resolved to contest possession of the remaining quarters of Saragossa inch by inch, and if necessary to retire to the suburb across the Ebro, destroying the bridge. The struggle, which was prolonged for nine days longer, resulted in the withdrawal of the French (Aug. 14), after a siege which had lasted 61 days in all. Palafox then attempted a short campaign in the open country, but when Napoleon’s own army entered Spain, and destroyed one hostile army after another in a few weeks, Palafox was forced back into Saragossa, where he sustained a still more ‘memorable second siege. This ended, after three months, in the fall of the town, or rather the cessation of resistance, for the town was in ruins and a pestilence had swept away many thousands of the defenders. Palafox himself, suffering from the epidemic, fell into the hands of the French and was kept 593 prisoner at Vincennes until December 1813. In June 1814 he was confirmed in the office of captain-general of Aragon, but soon afterwards withdrew from it, and ceased to take part in public ‘affairs. ‘From 1820 to 1823 he commanded the royal guard of King Ferdinand, but, taking the side of the Constitution in the civil troubles which followed, he was stripped of all his honours and offices by the king, whose restoration by French bayonets was the triumph of reaction and absolutism. Palafox remained in retirement for many years. He received the title of duke of Saragossa from Queen Maria Christine. From 1836 he took part in military and political affairs as captain-general of Aragon and a senator. He died at Madrid on the rsth of February 1847. A biographical notice of Palafox appeared in the Spanish trans- lation of Thiers’s Hist. des consulates de l’empire,. by P. de Madrago. For the two sieges of Saragossa, see C. W. C. Oman, Peninsular War, vol. i.; this account is both more accurate and more just than Napier’s. . PALAMAS, GREGORIUS (c. 1296-1359), Greek mystic and chief apologist of the Hesychasts (g.v.), belonged to a dis- tinguished Anatolian family, and his father held an important position at Constantinople. Palamas at an early age retired to Mt Athos, where he became acquainted with the mystical theories of the Hesychasts. In 1326 he went to Skété near Beroea, where he spent some years in isolation in a cell specially built for him. His health having broken down, he returned to Mt Athos, but, finding little relief, removed to Thessalonica. About this time Barlaam, the Calabrian monk, began his attacks upon the monks of Athos, and Palamas came forward as their champion. In 1341 and 1351 he took part in the two synods at Constantinople, which definitively secured the victory of the Palamites. During the civil war between John Cantacuzene and the Palaeologi, Palamas was imprisoned. After Cantacuzene’s victory in 1347, Palamas was released and appointed arch- bishop of Thessalonica; being refused admittance by the inhabitants, he retired to the island of Lemnos, but subsequently obtained his see. Palamas endeavoured to justify the mysticism of the Hesychasts on dogmatic grounds. The chief objects of his attack were Barlaam, Gregorius Acindynus and Nicephorus Gregoras. Palamas was a prolific writer, but only.a few of his works have been published, most of which will be found in J. P. Migne, Patro- logia graeca (cl., cli.). They consist of polemics against the Latins and their doctrine of the Procession of the Holy Ghost; Hesychastic writings; homilies; a life of St Peter (a monk of Athos); a rhetorical essay Prosopopeia (ed. A. Jahn, 1884), containing the accusations brought against the body by the soul, the defence made by the body, and the final pronouncement of the judges in favour of the body, on the ground that its sins are the result of inadequate teaching. See the historical works of John Cantacuzene and Nicephorus Gregoras, the Vita Palamae by Philotheus, and the encomium by Nilus (both patriarchs of Constantinople); also C. Krumbacher, Geschichte der byzantinischen Litteratur (1897). PALAMAU, a district of British India, in the Chota-Nagpur division of Bengal. It was formed out of Lohardaga, in 1894, and takes its name from a former state or chiefship. The administrative headquarters are at Daltonganj: pop. (1901), 5837. It consists of the lower spurs of the Chota-Nagpur plateau, sloping north to the valley. of the Son. . Area 4914 sq. m.; pop. (1901), 619,600, showing am increase of 38% in the decade; average density, 126 persons per sq. m., being the lowest in all Bengal. Palamau suffered severely from drought in 1897. A branch of the East Indian railway from the Son valley to the valuable coalfield near Daltonganj was opened in 1902. The only articles of export are jungle produce, such as lac and tussur silk. The forests are unprofitable. See Palamau District Gazetteer (Calcutta, 1907). PALAMCOTTAH, a town of British India, in the Tinnevelly district of Madras, on the opposite bank of the Tambraparni river to Tinnevelly town, with which it shares a station on the South Indian railway, 444 m. south of Madras. Pop. (r90r), 39,545. It is the administrative headquarters of the district, and also the chief centre of Christian missions in south India. Among many educational institutions may be mentioned the Sarah Tucker College for Women, founded in 1895. ''594 PALAMEDES, in Greek legend, son of Nauplius king of Euboea, one of the heroes of the Trojan War, belonging to the post-Homeric cycle of legends. During the siege of Troy, Aga- memnon, Diomedes and Odysseus (who had. been detected by Palamedes in an attempt to escape going to Troy by shamming madness) caused a letter containing money and purporting to come from. Priam to be concealed in his tent. They then accused Palamedes. of treasonable correspondence with the enemy, and he was ordered to be stoned to death. His father exacted a fearful vengeance from the Greeks on their way home, by placing false lights on the promontory of Caphareus. The story of Palamedes was first handled in. the Cypria of Stasinus, and formed the subject of lost plays by Aeschylus (Palamedes), Sophocles (Nauplius), Euripides (Palamedes), of which some fragments remain. Sophists and rhetoricians, such as Gorgias and Alcidamas,; amused themselves by writing declamations in favour of or against him. Palamedes was regarded as’ the inventor of the alphabet, lighthouses, weights and measures, dice, backgammon and the discus. See Euripides, Orestes, .432 and schol.; Ovid, Metam. xiii. 56; Servius on Virgil, Aeneid, ii. 82, and Nettleship’ s note in Conington’s edition ; Philostratus, Heroica, 11; Euripides, Frag. 581; for different versions of his death see Dictys Cretensis ii. 15; Pausanias ii. 20, 3;X. 31, 2; Dares. Phrygius, 28; monograph by O. Jahn (Hamburg, 1836). PALANPUR, a native state of India, in the Gujarat ey of Bombay, on the southern border of Rajputana. Area, 1766 sq. M.; pop. (1901), 222,627, showing a decrease of 19 % in the decade. The country is mountainous, with much forest towards the north, but. undulating and cpen in the south and east. The principal rivers are the Saraswati and Banas. The estimated gross revenue is £50,000; tribute to the gaekwar of Baroda, £2564. The chief, whose title is diwan, is an Afghan by descent. Thestate is traversed by the main line of the Rajputana—Malwa railway, and contains the British cantonment of Deesa. Wheat, rice and sugar-cane are the chief products. The state has suffered severely of recent years from plague. The town of PALANPUR is a railway junction for Deesa, 18 m. distant. Pop. (1901), 17,799. Palanpur also gives its name to a political agency, or collection of native states; total area, 6393 sq. m.; pop. (i901), 467,271, showing a decrease of 28 % in the decade, due to the effects ofifamnine. 32:1: PALANQUIN (pronounced palankeen, & en in which it ‘is sometimes spelled), 2 covered litter used in India and other Eastern countries. It is usually some eight feet long by four feet in width and depth, fitted with movable blinds or shutters, and slung on poles carried by four bearers. Indian and Chinese women of rank always travelled in palanquim, and they were largely used by European residents in India before the railways. The norimono of Japan and the kiaotsu of China differ from the Indian palanquin only in the method of attaching the poles to the body of the conveyance. The word came into European use through Port. palanguim, which represents an East Indian word seen in several forms, e.g. Maiay and Javanese palangki, Hindostani palki, Pali pallanko, &c., all in the sense of litter, couch, bed. The Sansk. paryanka, couch, bed, the source of all these words, is derived from pari, round, about, and anka, hook. The New. English. Dictionary points out the curious resemblance of these words with the Latin use of phalanga (Gr. dadayé) for a bearing or carrying pole, whence the Span. palanca and palanquino, a bearer. PALATE (Lat. palatum, possibly from the root of pascere, to feed), the roof of the mouth in man ‘and! vertebrate animals. The palate is divided, into two parts, the anterior bony “ hard palate’? (see Mout), and the posterior fleshy “ solt’ palate ” (see ,PHARYNX)., For’ the malformation. consisting in a longi- tudinal fissure in the roof of the mouth, see CLerr PALATE. _ PALATINATE (Ger. Pfalz),.a name. given’ generally to any district ruled by a count palatine, but particularly to a district of Germany,,a province of, the kingdom of, Bavaria, lying west of the Rhine. It is bounded on the N. by the Prussian, Rhine province and .the Hessian province of Rhein-Hessen; on the E. PALAMEDES—PALATINATE by Baden, from which it is separated by the, Rhine; on the - S. by the imperial province of Alsace-Lorraine, from which itis | divided by the Lauter; and on the W. by the administrative districts of Trier and Coblens belonging to the Prussian Rhine province. It has an area of 2288 sq. m., and a population (1905) of 885,280, showing a density of 386-9 to the square mile. As regards religion, the inhabitants are fairly equally distributed into Roman Catholics and Protestants. The rivers in this fertile tract of country are the Rhine, Lauter, Queich, Speirbach, Glan and Blies. ‘The Vosges, and their continuation the Hardt, run through the land from south to north and divide it into ‘the fertile and mild plain of the Rhine, together with the slope of the Hardt range, on the east, and the rather inclement district on the west, which, running between the Saarbrtick carboniferous mountains and the northern spurs of the Hardt: range, ends in a porphyrous cluster of hills, the highest point of which is the Donnersberg (2254 ft.). The country on the east side and on the slopes of the Hardt yield a number of the most varied products, such as wine, fruit, corn, vegetables, flax and tobacco. Cattle are reared in great quantity and are of excellent quality: The mines yield iron, coal, quicksilver and salt... The industries are very active, especially in iron, machinery, paper, chemicals, shoes, woollen goods, beer, leather:and tobacco.. The province is well served by railway communication and, fer purposes of administration, is divided’) into the» fellowing 16 districts: Bergzabern, Diirkheim, Frankenthal, Germersheim, Homburg, Kaisers- lautern, /Kirchheimbolanden, Kusel, Landau, Ludwigshafen, Neustadt, Pirmasens,’ Rockenhausen, St Ingbert, Spires and Zweibriicken. Spires (Speyer) is the seat of government, and the chief industrial centres are Ludwigshafen on the Rhine, which is the principal river port, Landau, and Neustadt, the seat of /the wine trade. See A. Becker, Die Pfalz und die Pfalzer (Leipzig, 1857); Mehlis, Fahrien durch die Pfalz (Augsburg, 1877); Kranz, Handbuch der Pfalz (Spires, 1902); Hensen, Pfalzfiihrer (Neustadt, 1905); and Naher, Die Burgen der rheinischen Pfalz (Strassburg, 1887). History.—The count palatine of the Rhine was a royal official who is first mentioned in the 1oth century. The first count was Hermann I., who ruled from 945 to 996, and although the office was not hereditary it appears to have been held mainly by his descendants until the death of Count Hermann III. in rrss. These counts had gradually extended their powers, had obtained the right of advocacy over the archbishop of Trier and the bishopric of Juliers, and ruled various isolated districts along the Rhine. «In 1155 the German king, Frederick I., appointed his step-brother Conrad as) count palatine. Conrad took up his residence at the castle of Juttenbuhel, near Heidelberg, which became the capital of the Palatinate. In 1195 Conrad was succeeded by his son-in-law Henry, son of Henry the Lion, duke of Saxony, who was a loyal supporter of the emperor Henry VI. After the latter’s death in 1197 he assisted his own brother Otto, afterwards the emperor Otto IV., in his attempts to gain the German throne. ‘Otto refused to reward Henry for this support, so in 1204 he assisted his rival, the German king Philip, but returned to Otto’s side after Philip’s murder in 1208. In 1211 Henry abdicated in favour of his son Henry, who died in 1214, when the Palatinate: was given by the German king Frederick IL. to Otto, the infant son of Louis L., duke of Bavaria, a member of the ‘Wittelsbach family, who was betrothed to Agnes, sister of the late count, Henry. The break-up of the duchy of Franconia had increased the influence of the count palatine of the Rhine, and the importance of his position among the princes of the empire is shown by Roger of Hoveden, who, writing of the election to the German throne in 1108, singles out four princes as chief electors, among whom is the count palatine of the Rhine. In the Sachsenspiegel, a collection of German laws which was written before 1235, the count is given as the butler (dapifer) of the emperor, the first place among the lay selectors. The Palatinate was ruled by Louis of Bavaria on behalf of his son until 1228, when it passed. to Otto who ruled until his death in 1253. Otto’s possessions were soon afterwards divided, ''PALATINE and his elder son Louis II: received the Palatinate and Upper Bavaria. Louis died in 1294 when these’ districts passed to his son Rudolph I. (dv 1319), and subsequently to his grandson Louis, afterwards the emperor Louis IV. By the Treaty of Pavia’ in 1329) Louis granted the Palatinate to his nephews Rudolph II. and Rupert I., who received from him at the same time’a portion of the duchy of Upper Bavaria, which was called the upper Palatinate to’ distinguish it from the Rhenish, or lower Palatinate: Rudolph died in 1353, after which Rupert ruled alone until his death in’ 1390. In 1355 he had sold a portion of ‘the upper Palatinate to’ the emperor Charles IV., but by various purchases he increased the area of the Rhenish Palatinate.’ His’ successor was his nephew’ Rupert II., who bought’ from) the German king’ Wenceslaus a’ portion of the territory ‘that his uncle‘had» sold to ‘Charles IV. He: died in 1398 and was'succeeded by his 'son'Rupert IIL.) In 1400 Rupert was elected German king, and when he ‘died in ‘1410 his posses- sions were divided among his four sons: the eldest, Louis IIL, received the Rhenish Palatinate proper; the second son, John, obtained the upper Palatinate; while the outlying districts of Zweibritcken and Simmern passed to Stephen, and that of Mosbach to Otto. When the possessions of the’ house ‘of Wittelsbach were divided in ‘1255 and the branches of Bavaria and the Palatinate were founded; a dispute arose over the exercise of the electoral vote, and the question was not settled until in 1356 the Golden Bull bestowed the privilege upon the count palatine of the Rhine, who exercised it until 1623.. The part played by Count Frederick V., titular king of Bohemia, during the Thirty Years’ War. induced the emperor Ferdinand II. to deprive him of his vote ard to transfer it to the duke of Bavaria, Maximilian I. By the Peace of Westphalia in 1648 an eighth electorate was created for the count palatine, to which was added the office of treasurer. In 1777, however, the count resumed the ancient position of his family in the electoral college, and regained the office of steward which he retained until the formal dissolution of the empire in 1806. To return to the history of the Palatinate as divided into four parts among the sons of the German king Rupert in 1410. John, the second of these brothers, died in 1443, and his son Christopher, having become king of Denmark in 1440, did not inherit the upper Palatinate, which was again united with the Rhenish Palatinate. Otto, the son of Otto (d. 1461), Rupert’s fourth son, who had obtained Mosbach, died without sons in 1499, and this line became extinct, leaving only the two remaining lines with interests in the Rhenish Palatinate. After Rupert’s death this was governed by his eldest son, thetelector Louis III. (d. 1436), and then by the latter’s sons, Louis IV. (d. 1449) and Frederick I. The elector Frederick, called the Victorious, was. one of the foremost princes of his time. His nephew and successor, the elector Philip, carried on a war for the possession of the duchy of Bavaria~-Landshut, which had been bequeathed to his son Rupert (d. 1504), but, when in 1507 an end was put to this struggle, Rupert’s son, Otto Henry, only received Neuburg and Sulzbach. Louis’ V. and then Frederick HI. succeeded Philip, but both died without sons and Otto Henry became elector. He too died without sons in 1559, when the senior branch became extinct, leaving only the branch descended from Rupert’s third son, Stephen. » Already on Stephen’s death in 1459 ‘this’ family had been divided into two branches, those of Simmern and of Zwei- briicken, and in 1514 the latter branch had been divided into the lines of Zweibriicken proper and of Veldentz. It. was Frederick, count palatine of Simmern, who’ succeeded to the Palatinate on Otto Henry’s death, becoming the elector Frederick III. The new elector, a keen but not a very bigoted Calvinist, was one of the most active of the Protestant princes. His son and successor, Louis VI. (d. 2583), was a Lutheran, but another son, John Casimir, who ruled the electorate on behalf of his young nephéw, Frederick IV., from 1583 to 1502, gave every encouragement to the Calvinists. A similar line of action was followed by Frederick IV. himself after 1592. §95 He was the founder and head) of the Evangelical: Union estab- lished to combat the: aggressive tendencies of the Roman Catholics. His son, the elector Frederick V., accepted the throne of Bohemia and thus brought on: the Thirty Years’ War. He was quickly driven from that country, and his own electorate was devastated by the Bavarians and Spaniards. At the peace of Westphalia in'1648 the Palatinate was restored to Frederick’s son, Charles Louis, but it was shorn of the upper’ Palatinate, which Bavaria retained as the prize of war. e Scarcely had the: Palatinate begun to recover when it was attacked by Louis XIV.) For six’ years (1673-79) the electo- rate was devastated by the French troops, and even after the Treaty of Nijmwegen it suffered from the aggressive policy of Louis. In: August) 1680°the elector Charles Louis died, and when his son and ‘successor;‘Charles, followed him to the grave five years later the ruling family became: extinct in the senior line. Mention has:already: been: made of a division of this family into two lines after 1459, and of a further division of the Zweibriicken line in 1514, when again two lines were founded. The junior of: these, that. of Veldentz, became’ extinct in 1694, but the senior, that of Zweibriicken proper, was still very flourishing. Under Count Wolfgang (d. 1569) it» had: pur- chased Sulzbach and Neuburg in 1557, and in'the person of his grandson, Wolfgang William (d. 1653) it had secured the coveted duchies of Juliers‘and Berg: It was Philip William of Neuburg, the son of Wolfgang: William, who became ‘elector palatine in succession to Charies in 1685. The French king’s brother, Philip, duke of Orleans, had married Charlotte Elizabeth, a sister of the late elector Charles, and consequently the French king claimed a part of Charles’s lands in 1680. His troops took Heidelberg and devastated the Palatinate, while Philip’ William took refuge in Vienna; where he died in. 1690... Then in’ 1697, ‘by the Treaty of ‘Ryswick, Louis abandoned his claim in return for a sum of money. | Just before this date the Palatinate began to be disturbed by troubles about religion. The great’ majority of the inhabitants were Protestants, but the family which succeeded in 1685 belonged to the Roman Catholic: Church: Philip William, however, gave equal rights to all his subjects, but under his son and successor, the elector John William, the Protestants were deprived of various civil rights until the intervention of Prussia and of Brunswick in 1705 gave them some redress. The next elector, a brother of the last one, was Charles Philip, who removed his ‘capital from Heidelberg to Mannheim in» 1720. He died without male issue in December 1742. His successor was his kinsman, Charles Theodore, count palatine of Sulzbach, a cadet of the Zweibriicken-Neuburg line, and now with the exception of one or two small pieces the whole of the Palatinate was united under one ruler. Charles: Theodore was'a prince of refined and educated tastes and during his long reign his country enjoyed prosperity. In 1777 on the extinction of the other branch of the house of Wittelsbach, he became elector of Bavaria, and the Palatinate was henceforward united’ with Bavaria, the elector’s capital being Munich. Charles Theodore died without legitimate sons’ in 1799, and his successor was Maximilian Joseph, a member of the Birkenfeld branch of the Zweibriicken family, who later became king of Bavaria as Maximilian T. In 1802 the elector was obliged to cede the portion of the Palatinate lying on the left bank of the Rhine to France, ‘and other portions to Baden and to Hesse-Darmstadt. Much of this, however, was regained in 1815, and since that date the Palatinate has formed part of the kingdom of Bavaria. See Widder; Versuch einer vollstindigen geographisch-historischen Beschreibung der Kurfiirstlichen Pfalz (Frankfort, 1786-1788) ; L: Hausser, Geschichte’ der Rheinischen Pfalz, (Heidelberg, 1845); Nebenius, Geschichte der Pfalz (Heidelberg, 1874) ; Giimbel, Geschichte der protestantischen Kirche der Pfalz (Kaisezslautern, 1885); the Regesten der Pfalzgrafen am Rhein, 1214-1508, edited by Koch and Wille (Innsbruck, 1894); ‘and Wild, Bilderatlas zur badisch- pfalzischen Geschichte (Heidelberg, 1904). PALATINE (from Lat. palatium, a palace,) pertaining to the palace and therefore to the emperor, king or other sovereign ''596 ruler. In the later Roman Empire certain officials attending on the emperor, or discharging other duties at his court, were called palatini; from the time of Constantine the Great the term was also applied to the soldiers stationed in or around the capital to distinguish them from those stationed on the frontier of the empire. In the East Roman Empire the word was used to designate officials concerned with the administration of the finances and the imperial lands. This use of the word palatine was adopted by the Frankish kings of the Merovingian dynasty. They employed a high official, the comes palatinus, who at first assisted the king in his judicial duties and at a later date discharged many of these himself. Other counts palatine were employed on military and administrative work, and the system was maintained by the Carolingian sovereigns. The word paladin, used to describe the followers of Charlemagne, is a variant of palatine. A Frankish capitulary of 882 and Hincmar, archbishop of Reims, writing about the same time, testify to the extent to which the judicial work of the Frankish Empire had passed into their hands, and one grant of power was followed by another. Instead of remaining near the person of the king, some of the counts palatine were sent to various parts of his empire to act as judges and governors, the districts ruled by them being called palatinates. Being in a special sense the representatives of the sovereign they were entrusted with more extended power than the ordinary counts. Thus comes the later and more general use of the word palatine, its application as an adjective to persons entrusted with special powers and also to the districts over which these powers were exercised. By Henry the Fowler and especially by Otto the Great, they were sent into all parts of the country to support the royal authority by checking the independent tendencies of the great tribal dukes. We hear of a count palatine in Saxony, and of others in Lorraine, in Bavaria and in Swabia, their duties being to administer the royal estates in these duchies. The count palatine in Bavaria, an office held by the family of Wittelsbach, became duke of this land, the lower title being then merged in the higher one; and with one other exception the German counts palatine soon became insignificant, although, the office having become hereditary, Pfalzgrafen were in existence until the dissolution of the Holy Roman Empire in 1806. The exception was the count palatine of the Rhine, who became one of the four lay electors and the most important lay official of the empire. In the empire the word count palatine was also used to designate the officials who assisted the emperor to exercise the rights which were reserved for his personal, consideration. They were called comites palatini caesarii, or comites sacri palatii; in German, Hof pfalzgrafen. From Germany the term palatine passed into England and Scotland, into Hungary and Poland. It appears in England about the end of the 11th century, being applied by Ordericus Vitalis, to Odo, bishop of Bayeux and earl of Kent. The word palatine came in England to be applied to the earls, or rulers, of certain counties, men who enjoyed exceptional powers. Their exceptional position is thus described by Stubbs (Const. Hist. vol. i): ‘They were ‘earldoms in which the earls were endowed with the superiority of whole counties, so that all the landholders held feudally of them, in which they received the whole profits of the courts and exercised all the regalia or royal rights, nominated the sheriffs, held their own councils and acted as independent princes except in the owing of homage and fealty to the king.”” The most important of the counties palatine were Durham and Chester, the bishop of the one and the earl of the other receiving special privileges from William I. Chester had its own parliament, consisting of barons of the county, and was not represented in the national assembly until 1541, while it retained some of its special privileges until 1830. The bishop of Durham retained temporal jurisdiction over the county until 1836. Lancashire was made a county, or duchy, palatine in 1351, and kept some of its special judicial privileges until 1873. Thus for several centuries the king’s writs did not run in these three palatine counties, and at the PALATKA—PALAZZOLO ACREIDE present day Lancashire and Durham have their own courts of chancery. Owing to the ambiguous application of the word palatine to Odo of Bayeux, it is doubtful whether Kent was ever a palatine county; if so, it was one only for a few years during the 11th century. Other palatine counties, which only retained their exceptional position for a short time, were Shropshire, the Isle of Ely, Hexhamshire in Northumbria, and Pembroke- shire in Wales. In Ireland there were palatine districts, aad the seven original earldoms of Scotland occupied positions some- what analogous to that of the English palatine counties. In Hungary the important office of palatine (Magyar Nddor) owes its inception to St Stephen. At first the head of the judicial system, the palatine undertook other duties, and became after the king the most important person in the realm. At one time he was chosen by the king from among four candidates named by the Diet. Under the later Habsburg rulers of Hungary the office was several times held by a member of this family, one of the palatines being the archduke Joseph. The office was abolished after the revolution of 1848. In Poland the governors of the provinces of the kingdom were called palatines, and the provinces were sometimes called palatinates. In America certain districts colonized by English settlers were treated as palatine provinces. In 1632 Cecilius Calvert, and Lord/Baltimore, received a charter from Charles I. giving him palatine rights in Maryland. In 1639 Sir Ferdinando Gorges, the lord of Maine, obtained one granting him as large and = prerogatives as were enjoyed by the bishop of Durham. Carolina was another instance of a palatine province. In addition to the authorities mentioned, see R. Schréder, Lehrbuch der deutschen Rechtsgeschichte (Leipzig, 1902); C. Pfaff, Geschichte des Pfajzgrafenamtes (Halle, 1847); G. T. Lapsley, The County Palatine of Durham (New York, 1900), and D. J. Medley, English Constitytional History (1907). GW) PALATKA, a city and the county-seat of Putnam county, Florida, U.S.A., in the N.E. part of the state, on the W. bank of the St John’s river, about 100 m. from its mouth, and at the head of deep-water navigation. Pop. (1905) 3950; (1910) 3779. Palatka is served by the Georgia Southern & Florida (of which it is the southern terminal), the Atiantic Coast Line, and the Florida East Coast railways, and also has connexion by water with Baltimore, New York and Boston. | Palatka is situated in a rich agricultural, orange-growing and timber region, for which it is the distributing centre. Large quantities of cypress lumber are shipped from Palatka. Palatka was incorpo- rated as a town in 1853, and in 1872 was chartered as a city. PALAVER (an adaptation of Port. palavra, a word or speech; Ital. parola; Fr. parole, from the Low Lat. parabola, a parable, story, talk; Gr. rapaBon7, literally ‘‘comparison”’; the Low Lat. parabolare, “to talk,” gives Fr. parler, “to speak,” whence “ parley,” ‘‘ parliament,” &c.), the name used by the Portuguese traders on the African coast for their conversations and. bargain- ing with the natives. It was introduced into English in the 18th century through English sailors frequenting the Guinea coast. It has now passed into general use among the negroes of West and West Central Africa for any conference, either among themselves or with foreigners. From the amount of unnecessary talk characteristic. of such meetings with natives, the word is used of any idle or cajoling talk. PALAWARAM, a town of British India, in Chingleput district, Madras, 11 m. S. of Madras.city, with a station on the South Indian railway; pop. (1901), 6416. Formerly called the presi- dency cantonment, as containing the native garrison for Madras city, it is now a dépét for native infantry and the residence of European pensioners. There are several tanneries. PALAZZOLO ACREIDE, a town of Sicily, in the province of Syracuse, 28 m. by road W. of it, 2285 ft. above sea-level. Pop. (1901), 14,840. The town occupies the site of the ancient Acrae, founded by Syracuse about 664 B.c. It followed in the main the fortunes of the mother city. In the treaty between the Romans and Hiero II. in 263 B.c. it was assigned to the latter. The ancient city lay on the hill above the modern town, the ''PALE—PALENCIA approach to it being defended by quarries, in which tombs of all periods have been discovered. The auditorium of the small theatre is well preserved, though nothing of the stage remains. Close to it are ruins of other buildings, which bear, without justification, the names Naumachia, Odeum (perhaps a bath establishment) and Palace of Hiero.. The water supply was obtained by ‘subterranean aqueducts. In the cliffs of the. Monte Pineta to the south are other tomb chambers, and to the south again are the curious bas-reliefs called Santoni or Santicelli, mutilated in the roth century by a peasant proprietor, which appear to be sepulchral also. Near here too is the necropolis of the Acrocoro della Torre, where many sarcophagi have been found. Five miles north lies Buscemi, near which a sacred grotto has been discovered; and also a church cut in the rock and surrounded by a cemetery.’ See G. Judica, Antichita di Acre (Messina, 1819). (Baron Judica’s collection of antiquities was dispersed after his death.) J. Schu- bring, Jahrbuch fur Philologie, Suppl. 1V., 662-672. PALE (through Fr. pal, from Lat. palus, a stake, for paglus, from the stem pag- of pangere, to fix; “pole ”’ is from the same original source), a stake, particularly one of a closely set series driven into the ground to form the defensive work known as a “palisade ’’; also one of the lighter laths or strips of wood set vertically and fastened to a horizontal rail to form a “ paling.” Used as am historical term, a pale is a district marked off from the surrounding country by a different system of government and law or by definite boundaries. The best known of these districts was the “‘ English Pale” in Ireland, dating from the reign of Henry II., although the word “ pale.” was not used in this connexion until the latter part of the 14th century. The Pale varied considerably, according to the strength or weakness of the English authorities, and in the time of Henry VIII. was bounded by a line drawn from Dundalk to Kells, thence to Naas, and from Naas E. to Dalkey, embracing, that is, part of the modern counties of Dublin, Louth, Meath, and Kildare. The Pale existed until the complete subjugation of Ireland under Elizabeth; the use of the word is frequent in Tudor, times. There was an “ English Pale ” or “‘ Calais Pale” also in France until 1558, extending from Gravelines to Wissant, and for a short. time under the Tudors an English Pale in Scotland. In heraldry, a ‘‘ pale”? is a band placed vertically in the centre of a shield, hence ‘‘ in pale”’ or “ to impale”’ is used of the marshalling of two coats side by side on a shield divided vertically. “Pale,” in the sense of colourless, whitish, of a shade of colour lighter than the normal, is derived through O. Fr. palle, mod. pale, from Lat. pallidus, pallor, pallere; and in that of a baker’s shovel, or “ peel ”’ as it is sometimes called, from Lat. pala, spade, probably connected with the root of pandere, to spread out. PALEARIO, AONIO (c. 1500-1570), Italian humanist and reformer, was born about 1500 at Veroli, in the Roman Campagna. Other forms of his name are Antonio Della Paglia, A. Degli Pagliaricci. In 1520 he went to Rome, where “he entered the brilliant literary circle of Leo X. When Charles of Bourbon stormed Rome in 1527 Paleario went first to Perugia and then to Siena, where he settled as a teacher. In 1536 his didactic poem in Latin hexameters, De immortalitate animarum, was published at Lyons. It is divided into three books, the first containing his proofs of the divine existence, and. the remaining two the theological and philosophical arguments for immortality based on that postulate.. The whole concludes with a rhetorical description of the occurrences of the Second Advent. In 1542 a tract, written by him and entitled Della Pienezza, sufficienza, et satisfazione della passione di Christo, or Libellus de morte Christi, was made by the Inquisition the basis of a charge of heresy, from which, however, he successfully defended himself. In Siena he wrote his Actio in pontifices romanos et corum asseclas, a Vigorous indictment, in twenty ‘‘testimonia,” against what he now believed to be the fundamental error of the Roman Church in subordinating Scripture to tradition, as well as against various particular doctrines, such as that of 1P. Orsi in Notizie degli Scavi (1899), 452-471; Rémische Quartal- schrift (1898), 624-631. 507 purgatory; it was not, however, printed until after his death (Leipzig, 1606). In 1546 he accepted a professorial chair at Lucca, which he exchanged in 1555 for that of Greek and Latin literature at Milan. Here about 1566 his enemies renewed their activity, and in 1567 he was formally accused by Fra Angelo the inquisitor of Milan. He was tried at Rome, condemned to death in October 1569, and executed in July 1570. An edition of his works (Ant. Paleariit Verulant Opera), including four books of Epistolae and twelve Orationes besides the De im- mortalitate, was published at Lyons in 1552; this was followed by two others, at Basel, and several after his death, the fullest being that of Amsterdam, 1696. A work, entitled Benefizio di Cristo (“‘ The Benefit of Christ’s Death ’’), has been attributed to Paleario on insufficient grounds. Lives by Gurlitt (Hamburg, 1805); Young (2 vols., London, 1860); Bonnet (Paris, 1862). PALENCIA, an inland province of Spain, one of the eight into which Old Castile was divided in 1833; bounded on the N. by Santander, E. by Burgos, S. by Valladolid, and W. by Valladolid and feat. POD. (l0GG)) 1oz4y2, died. 2750 sc. IM. Iie surface of the province slopes gradually S. to the Duero (Douro) valley. The principal rivers are the Pisuerga and the Carrion, which unite at Duefias and flow into the Duero at Valladolid. The chief tributaries of the Pisuerga within the province are the Arlanzon, the Burejo, the Cioza, and the united streams of the Buedo and Abanades; the Carrion is joined on the right by the Cueza. The north is traversed by the Cantabrian Mountains, the highest summit being the culminating point of the Sierra del Brezo (6355 ft.). There are extensive forests in this region and the valleys afford good pasturage. Theremainder of Palencia, the “ Tierra de Campos,” belongs to the great Castilian table-land. In the south is a marsh or lake, known as La Laguna de la Nava. The mountainous district abounds in minerals, but only coal and small quantities of copper are worked. The province is crossed in the south-east by the trunk railway connecting Madrid with France via Irun, while the line to Santander traverses it throughout from north to south; there are also railways from the city of Palencia to Leon, and across the north from Mataporquera in Santander to La Robla in Leon. A branch of the Santander line gives access to the Orbo coal-fields. The main highways are good; the other roads often bad. The Canal de Castilla, begun in 1753, and completed in 1832, connects Alar del Rey with Valladolid. Wheat and other cereals, vegetables, hemp and flax are extensively grown, except in the mountainous districts. Flour and wine are made in large quantities, and there are manufactures of linen and woollen stuffs, oil, porcelain, leather, paper and rugs. Palencia rugs are in great demand through- out Spain. The only town with more than 5000 inhabitants is Palencia (q.v.). For the history, inhabitants, &c., see CASTILE. PALENCIA, an episcopal city, and the capital of the Spanish province of Palencia; on the left bank of the river Carrion, on the Canal de Castilla, at the junction of railways, from Leon and Santander, and 7 m. N. by W. of Venta de Bafios on the Madrid-Irun line. Pop. (1900), 15,940.,. Palencia is built in the midst of the level plains called the Tierra de Campos, 2690 ft. above sea-level. Three bridges across the Carrion afford access to the modern suburbs on the right bank. The older and by far the more important part of the city is protected on the west by the river; on the other sides the old machicolated, walls, 36 ft. high by 9 ft. in thickness, are in fairly good. preservation, and beautified by alamedas or promenades, which were laid out in 1778.. The cathedral was begun in 1321, finished in 1504, and dedicated, to St Antolin; it is a large building in the later and florid Gothic style of Spain., The site was previously occupied by a church erected by Sancho III. of Navarre and Castile (1026-1035) over the cave of St Antolin, which, is still shown. The cathedral contains some, valuable paintings, old Flemish tapestry, and beautiful carved woodwork and stonework. The church of San Miguel is a good and fairly well-preserved example of 13th-century work; that of San Francisco, of the same date, is inferior and has suffered more from modernization. The ''598 hospital of San Lazaro is said to,date'in part from the time of the. Cid, (g.v.), who here married Ximena in: 1074. J ‘Much has been done for education. Palencia has also hospitals, a foundling refuge, barracks and a bull-ring.. Local industries include iron-founding, and the making of rugs, alcohol, leather, soap, porcelain, linen, cotton, wool, machinery and, matches. Palencia, the, Pallantia of Strabo and Ptolemy, was the chief town, of the Vaccaei., Its history, during the Gothic and Moorish periods is obscure; but it was a Castilian town of some importance in the 12th: and 13th centuries... The university founded here in 1208 by Alphonso TX. was removed in ‘1239 to Salamanca, PALENQUE, the modern name of. a deserted .city in Mexico, | in the narrow valley of the Otolum, in the north part of'the state of Chiapas, 80 m. S. of the Gulf port of Carmen. About;30,m. away, on the left bank of the Usumacinta river, stand the ruins of Men-ché or Lorillard city.. The original:name of Palenque has been lost, and its present name is taken from the neighbour- ing village, Santo Domingo del Palenque. Unlike the dead cities of the Yucatan plains, Palenque is surrounded by wooded hills and overgrown by tropical vegetation. ._., There is less stone carving on the exterior walls, door jambs and pillars of the buildings than on those of the Yucatan Penin- sula; this is due to the harder and more uneven, character of the limestone. Probably. owing to the same; cause, there is less cut stone in the walls, the Palenque builders using plaster to obtain smooth surfaces. There is, however, considerable. carving on the interior walls, the best. specimens: being on the tablets, affixed to the walls with plaster., Modelling in stucco was exten- sively used. A few terra-cotta images have been found. Paint and coloured washes. were liberally used to cover. plastered surfaces and for ornamentation, and paints seem to have been used to bind plastered surfaces. The Palenque builders apparently used nothing but stone tools in their work. The so-called Great Palace consists of a group of detached buildings, apparently ten in number, standing on two platforms of different elevations. Some of the interior structures. and the detached one on the lower. southern terrace are in a fair state, of preservation. The plan of construction shows three parallel walls enclosing two corridors covered with the peculiar pointed arches. or vaults. characteristic, of Palenque. The buildings appear to. haye been erected at different periods. A square tower rises from. a. central part of the platform to a height of about 40 ft., divided into a solid masonry base and three storeys connected by interior stairways. The Temple. of Inscriptions, one of the largest and. best preserved, is. distin- guished chiefly for its tablets, which contain only hieroglyphics. Sculptured slabs form balustrades to the steps leading up: to the temple, and its exterior is ornamented. with figures in stucco, the outer faces of the four pillars in front having life-size figures of women with children in their armsi°’ The small Temple of Beau Relief stands on a narrow ledge of rock against the steep slope of the mountain. ‘Its’most important feature is a large stucco bas-relief, occupying a central position on the back wall of the sanctuary. It consists of a single figure, seated on a throne, beautifully modelled ‘both in form, drapery and ornaments, with the face turned to one side and the arms out- stretched, and is reproduced by H. H. Bancroft. “The temples on the east side of the Otolum: are distinguished by tall narrow vaults, perforated by numerous square openings giving the appearance of coarse lattice work. The Temple of the Sun stands upon a comparatively low pyramidal foundation. The interior consists of the usual’ pair of vaulted corridors. The sacred tablet on the back wall of the sanctuary is carved in low relief in limestone, and consists of two figures, apparently a priest and his assistant making offerings. There are rows of hiero- glyphics on the sides and over the central design. The Temple of the Cross is a larger structure of similar design and construc- tion. The tablet belonging ‘to’ this temple has excited contro- versy, because the design contains a representation of a Latin cross. ‘The Temple of the Cerro, called that of the Cross No. 2, because its tablet is very similar to that just mentioned, which was revived under various forms in later times. PALENQUE+{-PALERMO stands back) against.the slope of the mountain, and is in great part. a ruin., (For: history and, further details see .CENTRAL America; § Archaeology.) ; PALERMO (Greek, [dvopyos; Latin, Panhormus, Panormus), a city of, Sicily, capital ,of.a-province, of the same name, in the kingdom. of Italy, and) the, see ofan archbishop. Pop. (1906),, town. 264,036, \commune 323,747. The: city. stands in the N.W. of the island,.on.a small bay.looking.E., the coast forming the chord of a.semicircle.of mountains which hem in the campagna.of Palermo, called the Conca, d’Oro.. The most striking point is the mountain.of Hiercte, now called Pellegrino (from the grotto of Santa Rosalia, a favourite place of pilgrimage) at the N. of this semicircle;:at the $:E.-is: the: promontory. of Zaffarano, on which: stood Soluntum (q.v:). A neolithic settlement and,-necropolis were. idiscovered -in 1897, at the foot of Monte Pellegrino, on the N.E. side (E. Salinas in Notizie degli Scavi; 1907, 307). | Palermo’ has ‘been commonly thought to be an original Phoenician ‘settlement of unknown date (though its true*Phoenician'name is unknown), but: Holm (Archivio ‘storio siciliano, 1880} iv. 421) has suggested ‘that ‘the settlement’ was originally Greek: There:is no ‘record! of ‘any Greek colonies/in that part’ of ‘Sicily, and’ Panormus ‘certainly was’ Phoenician ‘as far back as history can carry us: According to ‘Thucydides (vi. 2), as'the Greeks colonized’ ‘the E. of the island, the Phoenicians withdrew to the N.W:, and concentrated themselves at Panormus, Motye, and Soluntum. Like the other Phoenician colonies in the west, Panormus came under the power of Carthage, and became the head of the Carthaginian dominion in Sicily. As such it became the centre of that strife between’ Europe and’ Africa, between Aryan and Semitic man, in its later stages between Christendom and Islam, which forms the great interest of Sicilian history. As the’ Semitic head of Sicily, it stands opposed to Syracuse, the Greek head. Under the Carthaginian it was the head of the Semitic part of Sicily; when, under the Saracen all Sicily.came under Semitic rule, it was the chief seat of that rule’ It was thrice won for Europe, by’ Greek, Roman and Norman conquerors—in 276 B.c. by the Epirot’ king Pyrrhus, in 254 Bc. by the Roman ‘consuls Aulus Atilius and Gnaeus’ Cornelius Scipio, and in A.D. 1071 by Robert Guiscard and his brother Roger, the first count of Sicily. “After the conquest’ by Pyrrhus the city was ‘soon recovered by Carthage, but this first Greek’ occupation was the beginning of a connexion with western Greece’and its islands After the Roman conquest an attempt to recover the city for Carthage was made. in 250 B.c.,; which led only to.a great Roman victory (see Punic) Wars). .Later,-in the First Punic War, Hamilcar Barca was encamped for three years on Hiercte or Pellegrino, but the Roman possession of the city was not disturbed. Panor- mus received the privileges of autonomy and immunity from taxation. It seems probable that at the end of the republic the coinage for the west of Sicily was struck here (Mommsen, Rom. Miinzwesen, 665). A colony was sent: here by Augustus, and ‘the place remained of considerable importance, though inferior to Catana: A fortunate chance has preserved to us alarge number of the inscriptions set up in the Forum (Mommsen, Corpus inscr. lat. x:'752). The town was taken by the Vandal Genseric in A.D.-440.° It afterwards became a part of the East- Gothic dominion, and was recovered for the empire by Belisarius in 535. It again remained a Roman possession for exactly three hundred years, till it was taken | by theSaracens in 835. Panormus now became the Moslem capital. In\1062:the Pisan fleet broke through the chain’ of the: harbour and carried off much»spoil, which was spent on the building of the great church of Pisa. After the Norman conquest the city remained: for a short time in the hands of the dukes of ‘Apulia. But in 1093 half the city was ceded to Count Roger, and in 1122 the rest was ceded ‘to ‘the second Roger. When ‘he took the kingly: title in 1130 it became “‘ Prima sedes; corona regis, et’ regni caput.” 1 The coins ‘bearing the name of mind are no longer assigned to Panormus; but certain coins with the name js (Ziz; about 410 B.C.) belong to it. ''PALERMO During the Norman reigns Palermo was the main centre of Sicilian history, especially during the disturbances in the reign of William the Bad (1154-1166). The emperor Henry VI. entered Palermo in 1194, and it was the chief scene of his cruelties.. In 1198 his son Frederick, .afterwards, emperor, was crowned there. After his death: Palermo, was for a. moment a, commonwealth. It passed under the dominion of Charles. of Anjow in) 1266. In the next ‘year; when the greater part of Sicily revolted ‘on behalf of Conradin, Palermo was one of the few towns which was held for Charles; but the famous. Vespers of 1282 put an - - ———— TST. PALERMO. fegtreetei at Jarconte -aMonte Pellegrino : | Scale, 1537,000. MEN \ ° % % Mile [ 3 to eet le 1. Maseum %. Palazzo Chiaramonte i 2. Cathedral 8. S.Maria della Catena it 3. PalazzoReale 9. Quattro Cantoni ly i) ms 4.LaMartorana 10. Politeama Garibaldi K 11; Teatro Massimo Hau 6. San Giovanni__12. University 1 i. D Acquasante 5. San Cataldo degli Eremiti 13. Piazza Vittoria: \ \\ Wot") SSSA ie + oN Ot, Ba , Lot SSS Swear a Castellammare VOCE S NY. DVS oe Z SR .\Porta Fetice Emery Walker sc. end to the Angevin dominion. From’ that’time Palermo shared in the many changes of the Sicilian kingdom. In 1535 Charles V. landed there on ‘his ‘return from Tunis. The last kings crowned at Palermo were Victor Amadeus of Savoy, in 1713, and Charles IfI. of Bourbon, in 1735. The loss of Naples by the Bourbons in 1798, and again in 1806, made Palermo once more the seat ‘of a separate Sicilian kingdom. The’ city rose against Bourbon rule in 1820’and in 1848. In 1860 came the final deliverance, at the hands’ of Garibaldi; but ‘with it came also the yet’ fuller loss of the position of Palermo as the capital of a kingdom of Sicily. 7, Site —The original city was built on a tongue of land between two inlets of the sea. ‘There is no doubt that the’ present main street, the Cassaro (Roman castrum, Arabic Kasr), Via Marmorea or Via Toledo (ViasVittorio Emmanuele); represents the line of the ancient town, with water on each sidevof:it.: ~Another city itself. ao" peninsula ‘with one side to the open sea, meeting | as it were the main city at right angles, formed in Polybius’s time the Neapolis, or new town, in Saracen times Khalesa, a name which still survives in that of Calsa. But the two ancient) harbours have been dried up; the two peninsulas have met; the long street has been extended to the present coast-line; ‘ai small inlet, called the Cala; alone represents ‘the old haven. The'city kept its ancient shape till after the time of the Norman kings. The old state of things fully explains the name Ilavopyos. ‘ There: are not many early remains in) Palermo.: ‘The Phoeni- cian and Greek antiquities in the museum.do‘not belong to the The earliest existing buildings date from the time of the Norman kings, whose’ palaces and churches were built in the Saracenic’ and Byzantine styles prevalent in the island. Of Saracen’ works actually belonging»to the time of Saracen occupation there are no whole: buildings remaining, but many inscriptions and a good many columns, often inscribed) with passages from: the Koran, which have been used up again in later buildings, specially in the porch of the metropolitan church. This last: was built -by Archbishop Walter (fl..1170)— an English- man sent by Henry II: of England as tutor to William: II. of Sicilyand corsecrated in 1185, on the site of an ancient basilica, which on the Saracen conquest’ became“a mosque, and on the Norman conquest became ‘a church again, first of the Greek and then of the Latin rite. What remains of Walter’s building is a rich example of the Christian-Saracen style,’ disfigured, un- fortunately, by. the addition ofa totally unsuitable dome by Ferinando Fuga in 1781-1801.) This:church contains the tombs of the emperor Frederick Il..and his parentsmassive sarcophagi of red porphyry with canopies above them—and also the royal throne, higher than that:of the:archbishop: for the king of Sicily, as hereditary legate of the see of Rome, was’ the higher ecclesi- astical officer of the two.' But far'the best example of the style is the. chapel of the king’s palace (cappella palatina), at the west end of the city. » This is earlier than Walter’s church, being’ the work: of King Roger in’ 1143. The’ wonderful mosaics, the wooden roof, elaborately fretted and painted, and'the marble incrustation of the lower part of: the: walls and the floor are very fine. Of the palace itself the greater part»was rebuilt and added in Spanish times, but there are some other parts of Roger’s work left, specially the hall called Sala Normanna. Alongside of the churches of this: Christian-Saracen type, there is another class which follows the Byzantine type. Of these ithe most perfect is’ the very small church of San Cataldo. But the best, much altered, but now largely restored’ to its former state, is the adjoining church of La Martorana, the work of George of Antioch, King Roger’s admiral: This is rich with mosaics; among them: the portraits of the king and the founder. Both these and the royal chapel have several small ‘cupolas, and there is a still greater display in that way in the church of San Giovanni degli Eremiti, which it is hard to believe never was a mosque. It is the only church in Palermo: with ‘a’ bell-tower, itself crowned with a cupola. } Most of these buildings are witnesses in different ways to the peculiar position of Palermo in the rath century as the “city of the ‘threefold tongue,” » Greek, Arabic, and Latin. King Roger’s sun-dial in the palace’ is: commemorated in all three, and it is to be noticed that the three inscriptions do not translate one another: In: private ‘inscriptions a fourth tongue, the Hebrew, is also often found. ‘For in Palermo under the Norman kings ‘Christians: of both rites, Mahommedans and Jews were all allowed’ to flourish after their several fashions. In Saracen times there was a Slavonic quarter on the southern side of the city, and there is still a colony of United Greeks, or more strictly Albanians. I S18 ¢ ( The series’ of \Christian-Saracen ‘buildings is continued in the country houses of! the kings which surround the city, La Favara and» Mimnerno, the works’ of:Roger, and’ the better known Ziza-and°Cuba, the works severally’ of William the Bad and William the Good! The Saracenic architecture'and Arabic inscriptions of these buildings have often caused’ them to be taken foroworks ‘ofthe ancient:ameers; but’ the inscriptions of ''600 themselves prove their date. All these buildings are the genuine work of Sicilian art, the art: which had: grown up in the island through the presence of the two most civilized races of the age, the Greek and the Saracen. Later in the r2th century the Cistercians brought in a type of church which, without any great change of mere style, has a very different effect, a high choir taking in some sort the place of the cupola. The greatest example of this is the neighbouring metropolitan church of Monreale (q¢.v.); more closely connected with Palermo is the church of San Spirito, outside the city on the south side, the scene of the Vespers. Domestic and civil buildings from the 12th century to the 15th abound in Palermo, and they present several types of genuine national art, quite unlike anything in Italy. Of palaces the finest is perhaps the massive Palazzo Chiaramonte, now used. as the courts of justice, erected subsequently to 1307. One of the halls has interesting paintings of 1377-1380 on. its wooden ceiling; and in the upper storey of the court isa splendid three-light Gothic window. The later houses employ a very flat arch, the use of which goes on in some of the houses and smaller churches of the Renaissance. S. Maria della Catena may be taken as an especially good example. But the general aspect of the streets is later still, dating from mere Spanish times. Still many of the houses are stately in their way, with remarkable heavy balconies. The most striking point in the city is the central space at the crossing of the main streets, called the Quattro Cantoni. Two of the four are formed by the ancient Via Marmorea, but the Via Macqueda, which supplies the other two, was cut through a mass of small streets in Spanish times. The city walls are now to a great extent removed. Of the gates only two remain, the Porta Nuova and the Porta Felice; both are fine examples of the baroque style, the former was erected in 1584 to commemorate the return of Charles V. fifty years earlier, the latter in 1582. Outside the walls new quarters have sprung up of recent years, and the Teatro Massimo and the Politeama Garibaldi; the former (begun by G. B. Basile and completed by his son in 1897) has room for 3200 spectators and is the largest in Italy. The museum of Palermo, the richest in the island, has been transferred from the university to the former monastery of the Filippini.. Among the most important are the objects from prehistoric tombs.and the architectural fragments from Selinus, including several metopes with reliefs, which are of great impor- tance as illustrating the development of Greek sculpture. None of the numerous Greek vases and terra-cottas is quite of the first class, though the collection is important. The bronzes are few, but include the famous ram from Syracuse. There is also the Casuccini collections of Etruscan sarcophagi, sepulchral urns and pottery. Almost the only classical antiquities from Palermo itself, are Latin inscriptions of the imperial period, and two large coloured mosaics with figures found in the Piazza Vittoria in front of the royal palace in 1869: in 1906 excavations in the same square led to the discovery of a large private hcuse, apparently of the and or 3rd century A.D., to which these mosaics no doubt belonged. Of greater local interest are the medieval and Renaissance sculptures from Palermo itself, a large picture gallery, and an extensive collection of Sicilian majolica, &c. The university, founded in 1779, rose to importance in recent years (from 300 students in 1872 to 1495 in 1897), but: has slightly lost in numbers since. The city wears a prosperous and busy appearance. | The Marina, or esplanade at the south of the town, affords a fine sea front with a view of the bay; near it are beautiful public gardens.. In the immediate neigh- bourhood of the city are the oldest church in or near Palermo, the Lepers’ church, founded by the first conqueror or deliverer, Count Roger, and the bridge over the forsaken stream of the Oreto, built in King Roger’s day by the admiral George. There are also some later medieval houses and towers of some impor- tance. These all lie on to the south of the city, towards the hill called Monte Griffone (Griffon-Greek), and the Giant’s Cave, which has furnished rich stores for the palaeontologist. On PALES—PALESTINE the other side; towards Pellegrino, is the new harbour of Palermo, round which a new quarter has sprung up, including a yard capable of building ships up to 475 ft. in length, and a dry dock for vessels up to 563 ft. The steamship traffic at Palermo in 1906 amounted to 2035 vessels, with a total tonnage of 2,403,851 tons. Palermo is one of the two headquarters (the other being Genoa) of the Navigazione Generale Italiana, the chief Italian steamship. company... The principal imports were 36,567 tons of timber (a large increase on the normal figures), 21,4011 tons of wheat and 151,360 tons of coal; while the chief exports were 116,400 gallons of wine, 37,835 tons of sumach and 122,023 tons of oranges and lemons. F. inding most of its valuable rates hypothecated to the meeting of old debts, the municipality of Palermo has embarked upon municipal owner- ship and trading in various directions. The plain of Palermo is very fertile, and well watered by springs and streams, of the latter of which the Oreto°is the chief: “It is planted with orange and lemon groves, the products of which are largely exported, and with many palm-trees, the fruit. of. which, however, does not attain maturity. It also contains many villas of the wealthy inhabitants of Palermo, among the most beautiful of which is La Favorita, at the foot of Monte Pellegrino on the west, belonging to the Crown. AuTHORITIES—Besides works dealing with Sicily generally, the established local work on Palermo is Descrizione di Palermo antico, by Salvatore Morso (2nd ed., Palermo, 1827). Modern research and criticism have been applied in Die mitteldlterliche Kunst in Palermo, by Anton Springer (Bonn, 1869); Historische Topographie vo Panormus, by Julius Schubring (Liibeck, 1870); Studi di storia palermitana, by Adolf Holm (Palermo, 1880). See also ‘‘ The Normans in Palermo,” in the third series of Historical Essays, by E. A. Freeman (London, 1879). The description of Palermo in the second volume of Gselfel’s guide-book, Unter-Italien und Sicilien (Leipzig), leaves nothing to wish for. Various articles in the Archivio storico siciliano and the series of Documenti per servire alla storia della Sicilia, both published by the Societa siciliana per la storia patria, may also be consulted. GSA &.; TXAs.) PALES, an old Italian goddess of flocks and shepherds. The festival called Parilia (less correctly Palilia) was celebrated in her honour at Rome and in the country on the 21st of April. In this festival Pales was invoked to grant protection and increase to flocks and herds; the shepherds entreated forgiveness for any unintentional profanation of holy places of which their flocks might have been guilty, and leaped three times across bonfires of hay and straw (Ovid, Fasti, iv.-731-805). The Parilia was not only a herdsmen’s festival, but was regarded as the birthday celebration of Rome, which. was supposed to have been founded on the same day. Pales plays a very sub- ordinate part in the religion of Rome, even the sex of the divinity being uncertain, A’ male Pales was sometimes spoken of, corresponding in some. respects to Pan; the female Pales was associated with Vesta-and-Anna Perenna. PALESTINE, a geographical name of rather loose application. Etymologica! strictness would require it to denote exclusively the narrow strip of coast-land once occupied by the Philistines, from whose name it.is derived. It is, however, conventionally used as a name for the territory which, in the Old Testament, is claimed as the inheritance of the pre-exilic Hebrews; thus it may be said generally to denote the southern third of the province of Syria. Except in the west, where the country is bordered by the Mediterranean Sea, the limit of this territory cannot be laid down on the map as a definite line. .The modern subdivisions under the jurisdiction of the Ottoman Empire are in no sense conterminous, with those of antiquity, and hence do not afford a boundary by which Palestine can be separated exactly from the rest of Syria in the north, or from the Sinaitic and Arabian. deserts in the south and east; nor are the records of ancient boundaries sufficiently fuil and definite to make possible the complete demarcation of the country. Even the convention above referred to is inexact: it includes the Philistine territory, claimed but never settled by the. Hebrews, and excludes the outlying parts of the large area claimed in Num. xxxiv. as the Hebrew possession (from the ‘‘ River of Egypt.’ to. Hamath). However, the Hebrews themselves have preserved, in the 1 The figures for 1905 (40,005 tons, almost entirely from Russia) were abnormally high, while those for 1906 are correspondingly below the average. ''PHYSICAL FEATURES] proverbial expression ‘from Dan to Beersheba ” (Judg. XX. 1, &c.), an indication of the normal north-and-south limits of their land; and in defining the area of the country under discussion it is this indication which is generally followed. Taking as a guide the natural features most nearly correspond- ing to these outlying points, we may describe Palestine as the strip of land extending along the eastern shore of the Mediter- ranean Sea from the mouth of the Litany or Kasimiya River (33° 20’ N.) southward to the mouth of the Wadi Ghuzza; the latter joins the sea in 31° 28’ N., a short distance south of Gaza, and runs thence in a south-easterly direction so as to include on its northern side the site of Beersheba. Eastward there is no such definite border. The River Jordan, it is true, marks a line of delimitation between Western and Eastern Palestine; but it is practically impossible to say where the latter ends and the Arabian desert begins. Perhaps the line of the pilgrim road from Damascus to Mecca is the most convenient possible boundary. The total length of the region is about 140 m.; its breadth west of the Jordan ranges from about 23 m. in the north to about 80 m.-in the south. According to the English engineers who surveyed the country on behalf of the Pales- tine Exploration Fund, the area of this part of the country is about 6040 sq. m. East of the Jordan, owing to the want of a proper survey, no figures so definite as these are available. The limits adopted are from the south border of Hermon to the mouth of the Mojib (Arnon), a distance of about 140 m.: the whole area has been calculated to be about 3800 sq. m. The territory of Palestine, Eastern.and Western, is thus equal to rather more than one-sixth the size of England. There is no ancient geographical term that covers all this area. Till the period of the Roman occupation it was subdivided into independent provinces or kingdoms, different at different times (such as Philistia, Canaan, Judah, Israel, Bashan, &c.), but never united under one collective designation. The exten- sion of the name of Palestine beyond the limits of Philistia proper is not older than the Byzantine Period. Physical Features.—Notwithstanding its small size, Palestine presents a variety of geographical detail so unusual as to be in itself sufficient to mark it out as a country of especial interest. The bordering regions, moreover, are as varied in character as is the country itself—sea to the west, a mountainous and sandy desert to the south; a lofty steppe plateau to the east, and the great masses of Lebanon ‘to’ the north. In describing the general physical features of the country, the most significant point to notice is that (though it falls westward to the sea and rises eastward to an elevated plain) the rise from west to east is not continuous, but is sharply interrupted by the deep fissure of the Ghor or Jordan valley; which, running from north to south—for the greater part of its length depressed below sea-level—forms a division in the country of both physical and political importance. In this respect the function of the river Jordan in Palestine offers a strange contrast, often remarked upon, to that of the Nile in Egypt. The former is of no use for irrigation, except in the immediate neigh- bourhood of its banks, and is a barrier to cross which involves the labour of a considerable ascent at any point except its most northern section. The latter is at once the great fertilizer and the great highway of the country which it serves. Western Palestine is a region intersected by groups of mountain peaks and ranges, forming a southern extension of the Lebanon system and running southward till they finally lose themselves in the desert. The watershed of this system is so placed that from two-thirds to three-fourths of the country is on its western side. This fact, taken in connexion with the great depth of the depres- sion of the Ghor below the Mediterranean—already 682 ft. at the Sea of Galilee—has a peculiar effect on the configuration of the country. On the west side the slope is gradual, especially in the broad plain that skirts the coast for the greater part cf its length; on the east side it is steep—precipitous indeed, towards the southern end—and intersected by valleys worn to a tremendous depth by the force of the torrents that once ran down them. This territory of Western Palestine divides naturally into two longitudinal strips—the maritime plain and the mountain region. These it will be convenient to consider separately. I. The Maritime Plain, which, with a few interruptions, extends along the Mediterranean coast from Lebanon to Egypt, is a strip of land of remarkable fertility. It is formed of raised. beaches and sea-beds, ranging from the Pliocene period downwards, and resting on Upper Eocene sandstone. It varies greatly in width. At the mouth of the Kasimiya it is some 4 m. across, and this breadth it maintains to a short distance south of Tyre, where it suddenly narrows; until, at Ras el-Abiad, it has been necessary to PALESTINE 601 cut a passage in the precipitous face of the cliff to allow the coast- road to be carried past it. This ancient work is the well-known. “Ladder of Tyre.’’ South of this promontory the plain begins to widen again; on the latitude of Acre (Akka), from which this art of the plain takes its name, it is from 4 to 5 m. across; while arther south, at Haifa, it is of still greater width, and opens into the extensive Merj Ibn ‘Amir (Plain of Esdraelon) by which almost the whole of Western Palestine is intersected.. South) of Haifa the promontory of Carmel once more effaces the plain; here the passage along the coast is barely 200 yds. in width. At ‘Athlit, 9 m. to the south, it is about 2 m.; from this point it expands uni- formly to about 20 m., which is the breadth at the latitude of Ascalon. South of this it is shut in and broken up by groups of. low. hills. From the Kasimiya southwards the maritime plain is crossed by numerous river-beds, with a few exceptions winter torrents only. Among the perennial streams may be mentioned the Na‘aman, south of Acre; the Mukatta‘ “Kishon, at Haifa; the Nahr ez-Zerka, sometimes called the Crocodile River—so named from the crocodiles still occasionally to be seen in it; the Nahr el-Falik; the ‘Aujeh a few miles north of Jaffa and the Nahr Rubin. The surface of the plain rises gradually from the coast inland to an altitude of about 200 ft. It is here and there diversified by small hills. Il. The Mountain Region, the great plain of Esdraelon, which forms what from the earliest times has been recognized to be the easiest entrance to the interior of the country, cuts abruptly through the mountain system, and so divides it into two groups. Each of these may be subdivided into two regions presenting their own special peculiarities. a. The Galilean Mountains, north of the plain of Esdraelon, fall into two regions, divided by a line joining Acre with the north end of the Sea of Galilee. The northern region (Upper Galilee) ‘is virtually an outlier of the Lebanon Mountains. At the north end is an elevated plateau, draining into the Kasimiya. The mountains are intersected by a complex system of valleys, of which some thirty run down to the Mediterranean. The face toward the Jordan valley is lofty and steep. The highest poirt is Jebel Jermak, 3934 ft. above the sea; about it, on the eastern and northern sides, are lofty plateaus. The region is fruitful, and in places weil wooded; it is beyond question the most picturesque part of Palestine. The southern region (Lower Galilee) shows somewhat different characteristics. It consists of chains of comparatively low hills, for the greater part running east and west, enclosing a number of elevated plains. The principal of these plains is El-Buttauf, a tract 400 to 500 ft. above sea-level, enclosed within hills 1700 ft. high and measuring 9 m. east to west and 2 m. north to south. It is marshy at its eastern end and very fertile. This is the plain of Zebulun or Asochis, of antiquity. The plain of Tur‘an, south- east of El-Buttauf, is smaller, but equally fertile. Among the principal mountains of this district may be named Jebel Tur‘an, 1774 ft. and Jebel et-Tur (Tabor) 1843 ft.; the latter is an isolated mass of regular shape which commands the plain of Esdraelon. Eastward the country falls to the level of the Ghor by a succession of steps, among which the lava-covered Sahel el-Ahma may be mentioned, which lies west of the cliffs overhanging the Sea of Galilee. The chief valleys of this region are the Nahr Na‘aman and its branches, which runs into the sea south of Acre, and the Wadi Mukatta‘, or Kishon, which joins the sea at Haifa. On the east may be mentioned the Wadi er-Rubadiya, Wadi el-Hamam and Wadi Fajjas, flowing into the Sea of Galilee or else into the Jordan. b. The great plain of Esdraelon is one of the most important and striking of the natural features of Western Palestine. . It is a large triangle, having its corners at Jenin, Jebel et-Tur, and the outlet of the Wadi Mukatta‘t, by which last it communicates with the sea-coast. On the south-west it is bounded by the range of hills that terminates in the spur of Carmei. The modern name, as | above-mentioned, is Merj Ibn ‘Amir (‘‘ the meadow-land of the son of ‘Amir’’); in ancient times it was known as the Valley of Jezreel, of which name Esdraelon is a Greek corruption; and by another name (Har-Magedon) derived from that of the impeor- tant town of Megiddo—it is referred to symbolically in Rev. xvi. 16. It is the great highway, and also the great battlefield, of Palestine. At the village of Afuleh its altitude is 260 ft. above the sea-level. In winter it is swampy, and in places almost impassable. The fertility of this region is proverbial. There are several small subsidiary plains that extend from it both north and south into the surrounding mountain region; of these we need only mention a broad valley running north-eastwards between Jebel Duhi, a range 15 m. long and 1690 ft. high, on the one side, and Mt Tabor and the hills of Nazareth on the other side. East of the watershed are a number of valleys running to the Ghor; the most remarkable of these are the Wadi el-Bireh and the Wadi Jalud, the latter containing the river that flows from the fine spring called ‘Ain Jalud. c. The second of the divisions into which we have grouped the mountain system lies south of the plain of Esdraelon. This is divisible into the districts of Samaria and Judaea. In the first of these the mountain ranges are complex, appearing to radiate from a centre at which lies Merj el-Ghuruk, a small plain about 4 m. east to west and 2 m. north to south, This plain has no cutlet and is marshy in the rainy season. Connected with it are other small plains unnecessary to enumerate. For the greater part the ''602 principal mountains‘are' near the watershed; they include Jebel Fuku‘a (Gilboa), a range ‘that forms the watershed at the eastern extremity of the plain of Esdraelon. The range of Carmel: (highest PALESTINE [PHYSICAL FEATURES Judaean range, On, the eastern jside,of the. watershed the most important feature is perhaps the great valley system that connects the Mukhnah (the plain south of Nablus) with the Ghor—be- ginning with the impressive Wadi Bilan and proceeding ‘through the! important and: abundantly watered 35° NE point 1810 ft.) must also be included in this district; it runs from the A 34°30" B. oO 35230! D St George's Bay Ee PALESTI saan e Wen Na Scale; '1:1;600.000 English Miles 10 1, 20 Go Railways... Principal Roads. Canals & Aqueducts. Biblical & Classical Names... ‘Ain = Spring N.el-Zaherani | Sarafand 7 3 | Bahr = Sea 30’ | Beit, Beth = House v,, Jebel = Mountain USAREPTA, ZAREPHATHES Kefr = Village Kh., Khan =Inn Kh., Khurbet = Ruin N., Nahr = River (perennial) fas = Cape. Head Te/l = Mound ; W.=Wadi, Watercourse’ ne Ras e/-Abiad A TYRIORUMS er of TYTE A SCAL (Lada W.! ACHziB, E eis ¢, del é Rath @. fulei We B tito Wy BD I Fy Ez-ZibhW ‘CDIF PAST He va KOFBETES A = Oy <¢, Buyir Tubariya, SEA OF GALILE! pod se uw 9 Bera pari ° GENNE SEP ik Suge IER i « i Oo}. 33 mw i Akka, acRE.PTOLEM *, Bay of 4), Tell cs-Semak 8yCAMINUM 1 EvTird = Surafend/b Tells Tanturah, DOR, OORAB iit i Zimmari vi Sewish Colo ys Nahr ez-Zerka{-y “CROCODILE A. Kaisariya CAESAREA PALESTINA, N.el-Mefyir IGADARA JEq, i = x 2 jo N. Iskanderuna SALT AL Mukhalid/o Nahr el-Falik & & wy AN ~ ) te Arsu APOLLON) Rahr el-"Auja Jaffa sx? JOPPA N YA RRS x Arona jerinan , Colony) 5 ceri ve ea NYO 7 K & eG ES only. tha Suaf! ‘Askalan ASCALU 5 OE | pis Toi! Faras, 3 a : ye Gi Periatah rg S UinmK dis SO" BIL A arly % MOA P zg j E1nAl, ELEAL AN, NNHE Umm er-Ra ellA; far: {§ ASHTAAOTH ARETH J sh; Shg ELIOBOLS Wadi Far‘a.. Tell; sAsur.stands a) short distance north of Beitein (Bethel). South of it is the long zigzag range known as Jebel el-Kuds, “named from’ Jerusalem (el-Kuds) the chief town built upon it. The highest, point is Neby Samwil (Mizpah), 2935 ft. above the sea, north of Jerusalem. This city itself stands at an altitude of 2500 ft. To the south of it begins the ‘ subdivision of the Judaean mountains now known’ as: Jebel, el-Khalil, from, Hebron (el-Khalil),, which stands in an elevated basin some 500 ft. above the altitude of Jerusalem; ‘it ‘is here that the Judaean Mountains attain: their greatest height. 2 South, of Hebron) the ridge gradually, be- sea . comes. lower, and finally breaks. up and ci re loses itself in the southern desert. ‘ a On the west side of the watershed the yey mountainous’ district. extends about half way to the sea, broken. by deep valleys and passes. Among these the most im- portant are the Wadi Selman (Valley of Aijalon) which seems to have been the principal route to Jerusalem “in ancient times; the, Wadi, Isma‘in. south of this, along which.runs the modern carriage road from Jaffa to Jerusalem; and the Wadi es-Surar, a higher section of the bed of the Nahr Rubin, along which now runs: the railway line; farther to. the, south we may mention the, Wadi es-Sunt, which opens up the country from Tell es-Safi (Gath?) eastward. Between the mountainous country of Judaea and the maritime plain is an un- dulating region anciently known as the Shephelah. It is composed of horizontal! strata of limestone, forming groups of hills intersected by’ a network of small and fertile valleys.,: In. this region, which is of great historical importance, are the re- mains of many ancient cities. The ad- jacent part of the maritime plain is com- posed ‘of a rich, light. brown loamy. soil. Although. cultivated. with most primitive appliances, and with little or no attempt at irrigation or artificial fertilization, the average yield is eight- to, twelve-fold annually: This part of the plain is (in European nomenclature) divided into two at about the latitude of Jaffa, that to the north being the plain of Sarona (Sharon), the southern half being the plain of the Philistines. On the east side of the watershed the ground slopes rapidly from, its height of 2500 ft. above sea-level to: a. maximum depth of 1300 ft. below sea-level, within.a distance of about 20 m.. It is a waste, destitute of water and with but scanty vegetation... It has never been brought into cultivation; but in the first Christian OS#- VEEL is wai) Mania gloat | #Sho‘ar Noara Te Melty 4 PBs-Sunamei J Ia aos ra Tell e!-Ha. Kuneiydad HA Ss N Sshei¢n Sheikh Miski LGhair lita ALEMA a) y ie LF tiresh Shahe df Arar. > ‘Core d- Dera cty Gharz b “Soy Riel tour 0 ug EE abompa.QUuwAro BY. LN seh neurastel ange Dibdan,0iBON o PEs Lou centuties the caves in its valleys were the chosen refuge of Christian monasticism. It descends to the level, of the Ghor by terraces, deeply cut through by profound ravines such as the Wadi es-Suweinit, Wadi Kelt, Wadied-+Dabr, Wadien-Nar (Kedron) and Wadi el ‘Areijeh. e The southern district; which includes the white marl region of Beersheba, was in ancient times called the Negeb. It isa wide steppe region) which (though it contains vy J.er-Ram 7, 6 A3430° many remains of ancient towns and settle- ments, and was evidently at one timea terri- EB central ‘point’ above mentionéd—though interrupted by many passes—to the end of the promontory which’ makes the harbour of Haifa, at its foot, the best on the Palestine coast. The highest mountains in the Samaria district are, however, in the neighbour- hood of Nablus (Shechem).' They include the rugged bare mass of Gerizim (2849 ft.), the smoother cactus-clad cone of Ebal (3077), and farther south’ Tell’ ‘Asur (3318) at’ which point ‘begins the 4 Finery Wolk ae on tory of great importance) is now almost en- tirely inhabited by nomads. It should, however, be mentioned that the Turkish government has developed a town at Beersheba, under the jurisdiction of a’ Kaimmakam (lieutenant-governor), since the beginning of the 20th century. i g9214 The Ghor or Jordan ley, is treated in a separate article | (see JorpDAN).. There has been no systematic survey of Eastern Palestine such as'was carried out in Western Palestine between 1875 and. 1880 ''PHYSICAL FEATURES] by the officers of the) Palestine, Exploration Fund, A good, deal of work has been done by individual travellers, but the material for a full description. of its physical character is as yet lacking. Two great rivers, the’ Yarmuk (Hieromax) and the Zerka (Jabbok), divide Eastern ) Palestine. into three séections,) namely Hauran (BASHAN, g.v.), with the Jaulan west, of it; spel Ajlun: (GILEAD, g.v.); and the Belk’a (the southern portion of Gilead and the ancient territory of the tribe of Reuben). The latter extends southward to the’ Mojib, which, as we have already seen, is the southern boundary of Eastern Palestine. Lt It is a matter of dispute whether Hauran. should, be’ included within Palestine proper, accepting its definition as the ‘‘ ancient Hebrew territory.” It is a large volcanic region, entirely covered with lava and’ other igneous rocks: Two remarkable rows of these run in lines from north to south, through the region of the Jaulan parallel to the Ghor, and from a. long. distance are conspicuous features in the landscape. The soil is fertile, and there are many remains of ancient wealth and civilization scattered over its surface. South of the’ Yarmuk the formation is Cretaceous, Hauran basalt being found only» in the eastern portion.: That region)is much more mountainous than Hauran.; South. of the Zerka, the country culminates.in Jebel ‘Osha, a.peak of Jebel Jil‘ad. (‘the mountain of Gilead’), 3596 ft. high. From this point southward the country assumes the appearance ‘which is familiar to’ those who’ have visited Jerusalem—an elevatéd plateau, bounded on the west by the pre- cipitous cliffs known as the mountains of. Moab,,with but.a few peaks, such as Jebel Shihan (2781 ft.) and Jebel Neba (Nebo, 2643 {t.), con- spicuous above the level of the ridge by reason of superior height. Geology.—The oldest rocks consist of gneiss' and schist, penetrated by dikes and bosses» of: granite, syenite,) porphyry. ‘and’ other in; trusive rocks. All of, these,are pre-Carboniferous jin,age and most, of them probably belong to the Archean period. They are, gener- ally concealed by later’ deposits, but are exposed to view along the eastern margin ‘of’ the Wadi Araba, at ‘the foot of the plateau of Edom. «Similar rocks: occur) also at one or two places im ithe desert. of et-Tih, while.towards the.south they, attain a); greater extension, forming nearly the whole of Sinai and of the hills on the east side of the Gulf of Akaba. These ancient rocks, which form, the foundation of the country, are overlaid unconformably by a series of conglomerates and» sandstones,» generally unfossiliferous and often red or purpie,im colour, very,similar.in, character to. the Nubian sandstone of Upper Egypt.. In the midst. of. this. series: there is an inconstant band of fossiliferous limestone, which has been found im the Wadi’ Nasb and at other places on’ the southern border of et-Tih, and: also along :the western escarpment of the Edom plateau. ..The, fossils. include. Syringopora, Zaphrentis, Productus, Spirifer,, &c., and belong to the Carboniferous. The sandstone which lies below the limestone is also, no doubt, of Carboniferous ages but the’ sandstone above is conformably over- laid | by Uppers Cretaceous beds; and iis generally ‘referred to! 'the Lower, Cretaceous;, .No,..unconformity,; however, has.jyet -been detected anywhere in the sandstone series, and in the absence of fossils the’ upper “sandstone may represent any’ period from the Carboniferous to the Cretaceous: The Upper Cretaceous is repre- sented by limestones with bands of chert, and:contains Ammonites, Baculites, Hippurites..and other, fossils. .:It. covers; by. far. the greater part of Palestine, capping the table-lands of Moab. and Edom, and forming most of the high land between the Jordan and the Mediterranean.) It is overlaid towards’ the’ wést by similar limestones, which contain nummulites);and belong’ to the Eocene period; and these are followed, near the.coast by, the) calcareous’ sandstone of Philistia, which is referred by Hull to the Upper Eocene. Lava flows of basic character, belonging to the Tertiary period, cover extensive areas’in Jaulan’and Hauran; and smaller patches occur in the land of Moab. and ialso’ west, of: the. Jordan, especially near the Sea of Gennesareth, Of, Recent deposits the most interesting are the raised beaches near the coast and the terraces of the Jordan-Araba depression. The latter indicate that at one period nearly the whole of this depression ‘was filled with ‘water’ up to a level somewhat above that of the Mediterranean. The geological structure of the country is very.simple,in its. broad features, but of exceptional interest. In general. the stratified deposits lie nearly flat and in regular conformable succession, the lowest resting upon the floor of ancient crystalline rocks. ‘There is, however, a slight dip.towards) the west, soothat the newest deposits lie near the coast.. Moreover, along the eastern side of the Jordan- Araba valley there is a great fault, and on the eastern side of this fault the whole series of rocks stands at a much higher level than on the west. Consequently, west of the Jordan almost the whole country is formed ofthe newer beds (Upper Cretaceous and later), while east of the Jordan the older rocks, sometimes down.to, the Archean floor, are exposed at the foot of the plateau. The western margin of the valley is possibly defined by another fault which has not yet been detected; but in’any ‘case itis clear that the great depression owes its extraordinary depth to faulting. A. line of depressions of similar character has been traced,by,.E..Suess.as far south as Lake Nyasa.t See Lortet, La Mer Morte (Paris, 1877); E. Hull, Mount Seir, Sina and Western Palestine (London, 1885); and Memoir. on the PALESTINE 603 Climate.—Palestine belongs’ to the’-sub-tropical zone: at the summer solstice the sun is ten degreessouth of the zenith. Thelength of the day ranges) from ten to fourteen hours.’ The great variety of altitude and of surface characteristics gives rise to a considerable numberof local climatic peculiarities. On the maritime-plain the mean, annual temperature is 70° F., the normal extremes being about: 50° to) about 90°. The harvest ripens about a fortnight earlier than’ among the mountains. Citrons and oranges flourish, as do melons and palms: the latter do not fruit abundantly, but this is less the fault of climate than of carelessness in fertilization. The rainfall is rather lower than) among the mountains. In’ the mountainous regions the mean annual temperature is about 62°, but there is a great range of variation. In winter there are often several degrees of frost, though snow very rarely lies) for: more than a day or two. In summer the thermometer. occasionally registers as much as 100° in)» the: shade, or evem a degree or two more:/ this however ‘is exceptional, and. 80°-90° is a more normal maximum for the year... The rainfall is about 28 in., sometimes less; and in exceptional years as much as 10\in. in excess. of this figure has been registered. . The, vine, fig, and: olive grow. well in this region... The climate of the Ghor, again, is different. Here the thermometer, may, rise as,high as,130°., The rainfall is scanty, but as no civilized. person inhabits the southern end of the Jordan valley throughout the ‘year, and) it has. hitherto proved impossible ‘to establish self-registering instruments, no systematic meteorological observations have been taken. In Eastern Palestine there is even a greater range cf temperature; the loftier heights are covered in winter with snow. ( The’thermometer may range within twenty-four hours from freezing-point to;80°. qs The rainy season, begins about the end of .November, - usually with a heavy thunderstorm: the rain, at this part of the year is the’ “ former rain”’ of the Old Testament. The earth, baked hard by the summer heat, is thus softened, and ploughing begins at once.»,The wettest month, as indicated by meteorological obser- vation, is January; February, is: second to.it, and. December third; March is also a very wet month. In April the rains come to an end (the‘‘“latter rains”) and the winter crops receive their final fertilization. The winter crops (barley and’ wheat) are harvested from. April to. June. The; summer, crops. -(millet;! sesame, figs, melons, grapes,., olives, &c.); are fertilized by. the heayy \‘‘ dews.”’ which are one of the most remarkable. climatic features of the country ‘and’'to a large extent atone for the ‘total’ lack’ of rain forone half the -year:. These crops'are harvested ‘from’ August to October. is Water Supply—Notwithstanding the long drought, it must..not be supposed that Palestine is a waterless country, except in certain districts. There ate very few spots from which a spring of some sort is not accessible. | Perennial’'streams are, and in the recent geological ages always have been, rare in the country. The whole face of the land is pitted with ancient cisterns; indeed, many hillsides and fields are on that account most dangerous to walk over by night, except for those who are’thoroughly familiar with the landmarks. These: cisterns’ are: bell-shaped or ‘bottle-shaped ‘excavations,’ with a narrow circular shaft in the top, hollowed; in the rock:and lined with cement. Besides these, more ambitious works are. to be found, all now more or less ruined, in various parts of the country (see AQueDucts: Axcient). Such are the aqueducts, of which remains exist at Jericho, Caesarea and other places east and’ west of the Jordan; but especially must be, mentioned the enormous reservoirs known as Solomon’s Pools, in a valley between Jerusalem , and Hebron, by which the former city was supplied with water through an elaborate system of conduits. Many of these aqueducts, as well as countless numbers of now leaky ‘cisterns, could’ with’ but. little trouble be brought into, use, again, and would greatly enhance the fertility of the country... The most abundant springs in Palestine ‘are the sources of the Jordan at Banias and at Tell el-Kadi. A considerable number of springs in the country are brackish, being impregnated with chemicals of various kinds or (when near a town) with sewage... The latter is the case, of the Virgin’s Fountain (Ain Umm ed-Daraj), which is the only natural source of water in the neighbourhood of Jerusalem. 4 Hot springs are found in various parts of the country, especially at El-Hamma, about 1 m. south of ‘Tiberias, where! the water has: a temperature of 140° F, . This, is still, used for curative purposes, as it was in the days of Herod, but it is neglected and dirty... The spring of the Zerka Ma‘in (Calirrhoe) has'a temperature of 142° F. There are also ‘hot sulphur springs on the west side of the Dead Sea. Those of El-Hamma, ‘below Gadara, are from 104° to 120° F. in temperature, Fauna.—lIt has been calculated that about 595 different species of vertebrate animals are recorded or still to be found in Palestine— about 113 being mammals (including 'a few now extinct), 348 birds (including: 30 species peculiar: to the country), 91 reptiles and '43 fishes. Of the. invertebrata' the number. is) unknown, but, it must be enormous. The most important domestic animals are the sheep and the goat; the breed of oxen is small/and poor. The camel, the horse and the donkey are the draught animals; the flesh of the first Geology and Geography of Arabia Petraea, Palestine and adjoining Districts (London, 1886). i ''604 is eaten by the poorer classes, as is also occasionally that of the second. The dogs, which prowl in large numbers round the streets of towns and villages, are scarcely domesticated; much the same is true of the cats. Wild cats, cheetahs and leopards are found, but they are now rare, especially the latter. The lion, which inhabited the country in the time of the Hebrews, is now extinct. The most important wild animals are the hyena, wolf (now compara- tively rare), fox and jackal. Bats, various species of rodents, and gazelles are very common, as is the ibex in the valleys of the Dead Sea... Among the most characteristic birds may be mentioned eagles, vultures, owls, partridges, bee-eaters and hoopoes; singing birds are on the whole uncommon. Snakes—many of them venomous—are numerous, and there are many varieties of lizards. The crocodile is seen (but now very rarely) in the Nahr ez-Zerka. Scorpions and large spiders are a universal pest. Flora.—The flora of Palestine has a considerable range and variety, owing to the variation in local climatic conditions. In the Jordan valley the vegetation has a semi-tropical character, consonant with the great heat, which here is normal. The coast-plain has another type, i.e. the ordinary vegetation of the Mediterranean littoral. In the mountains the flora is, naturally, scantier than in these two more favoured regions, but even here there is a rich variety. In all parts of the country the contrast between thelandscape in early spring and later, when the cessation of rains and the increase of heat has burnt up the vegetation, is very remarkable. Population—The inhabitants of Palestine are composed of a large number of elements, differing widely in ethnological affinities, language and religion. It may be interesting to men- tion, as an illustration of their heterogeneousness, that early in the 20th century a list of no less than fifty languages, spoken in Jerusalem as vernaculars, was there drawn up by a party of men whose various official positions enabled them to possess accurate information on the subject.! It is therefore no easy task to write concisely and at the same time with sufficient fullness on the ethnology of Palestine. There are two classes into which the population of Palestine can be divided—the nomadic and the sedentary. The former is especially characteristic of Eastern Palestine, though Western Palestine also contains its full share. The pure Arab origin of the Bedouins is recognized in common conversation in the country, the word ‘“ Arab” being almost restricted to denote these wanderers, and seldom applied to the dweilers in towns and villages. It should be mentioned that there is another, entirely independent, nomad race, the despised Nowar, who correspond to the gipsies or tinkers of European countries. These people live under the poorest conditions, by doing smith’s work; they speak among themselves a Romani dialect, much contaminated with Arabic in its vocabulary. The sedentary population of the country villages—the fellahin, or agriculturists—is, on the whole, comparatively unmixed; but traces of various intrusive strains assert themselves. It is by no means unreasonable to suppose that there is a funda- mental Canaanite element in this population: the “ hewers of wood and drawers of water’ often remain undisturbed through successive occupations of a land; and there is a remark- able correspondence of type between many of the modern fellahin and skeletons of ancient inhabitants which have been recovered in the course of excavation. New elements no doubt came in under the Assyrian, Persian and Roman dominations, and in more recent times there has been much contamination. The spread of Islam introduced a very considerable Neo-Arabian infusion, Those from southern Arabia were known as the Yaman tribe, those from northern Arabia the Kais (Qais). These two divisions absorbed the previous peasant population, and still nominally exist; down to the middle of the roth century they were a fruitful source of quarrels and of bloodshed. The two great clans were further subdivided into families, but these minor divisions are also being gradually broken down. In the roth century the short-lived Egyptian government introduced into the population an element from that country which still persists in the villages. These newcomers have not been completely assimilated with the villagers among whom they 1 This list was intentionally made as exhaustive as possible, and included some languages (such as Welsh) spoken by one or two individual residents only. But even if, by omitting these accidental items, the list be reduced to thirty, a sufficient number will be left to indicate the cosmopolitan character of the city. PALESTINE [POPULATION have found a home; the latter despise them, and discourage intermarriage. _ Some of the larger villages—notably Bethlehem—which have always been leavened by Christianity, and with the develop- ment of industry have become comparatively prosperous, show tangible results of these happier circumstances in a_ higher standard of physique among the men and of personal appearance among the women. It is not uncommon in popular writings to attribute this superiority to a crusader strain—a theory which no one can possibly countenance who knows what miserable degenerates the half-breed descendants of the crusaders rapidly became, as a result of their immoral life and their ignorance of the sanitary precautions necessary in a trying climate. The population of the larger towns is of a much more complex nature. In each there is primarily a large Arab element, consisting for the greater part of members of important and wealthy families. Thus, in Jerusalem, much of ‘the local influence is in the hands of ‘the families of El-Khalidi, EI- Husseini and one or two others, who derive their descent from the heroes of the early days of Islam. The Turkish element is small, consisting exclusively of officials sent individually from Constantinople. There are very large contingents from the Mediterranean countries, especially Armenia, Greece and Italy, principally engaged in trade. The extraordinary development of Jewish colonization has since 1870 effected a revolution in the balance of population in some parts of the country, notably in Jerusalem. There are few residents in the country from the more eastern parts of Asia—if we except the Turkoman settle- ments in the Jaulan, a number of Persians, and a fairly large Afghan colony that since 1905 has established itself in Jaffa. The Mutawileh (Motawila), who form the majority of the inhabitants of the villages north-west of Galilee, are probably long-settled immigrants from Persia. Some tribes of Kurds live in tents and huts near Lake Huleh. If the inmates of the count- less monastic establishments be excluded, comparatively few from northern or western Europe will remain: the German “Templar ”’ colonies being perhaps the most important. ‘There must also be mentioned a Bosnian colony established at Caesarea Palestina, and the Circassian settlements placed in certain centres of Eastern Palestine by the Turkish government in order to keep a restraint on the Bedouin: the ‘latter are also found in Galilee. There was formerly a large Sudanese and Algerian element in the population of some of the large towns, but these have been much reduced in numbers since the beginning of the 20th century: the Algerians however still maintain themselves in parts of Galilee. The most interesting of all the non-Arab communities in the country, however, is without doubt the Samaritan sect in Nablus (Shechem); a gradually disappearing body, which has maintained an independent existence from the time when they were first settled by the Assyrians to occupy the land left waste by the captivity of the kingdom of Israel. The total population of the country is roughly estimated at 650,000, but no authentic official census exists from which satisfactory information, on this point is obtainable. Some two-thirds of this number are Moslems, the rest Christians of various sects, and Jews. The largest town in Palestine is Jerusalem, estimated to contain a population of about 60,000. The other towns of above 10,000 inhabitants are Jaffa (45,000), Gaza (35,000), Safed (30,000), Nablus (25,000), Kerak (20,000), Hebron (18,500), Es-Salt (15,000); Acre (11,000), Nazareth (11,0c0). The above remarks apply to the permanent population. They would be incomplete without a passing word on the non-permanent elements which at certain seasons of the year are in the principal centres the most conspicuous. Especially in winter and early spring crowds of European and American tourists, Russian pilgrims and Bokharan devotees jostle one another in the streets in picturesque incongruity. Political. Divisions —Under the Ottoman jurisdiction Palestine has no independent existence. West of the Jordan, and to about half-way between Nablus and Jerusalem, is the southern portion of ''OLD TESTAMENT HISTORY] the vilayet or province of Beirut. South of this point is the sanjak? of Jerusalem, to which Nazareth with its immediate neighbourhood is added, so as to bring all the principal ‘‘ Holy Places ’’ under one jurisdiction. East of the Jordan the country forms part of the large vilayet of Syria, whose centre is at Damascus. Communications.—Until 1892 communication through the country was entirely by caravan, and this primitive method is still followed over the greater part of its area. On the 26th of September of that year a railway between Jaffa and Jerusalem, with five intermediate stations, was opened, and has much facilitated transit between the coast and the mountains of Judaea. A railway from Haifa to Damascus was opened in 1905; it runs across the Plain of Esdraelon, enters the Ghor at Beisan, then, turning northwards, impinges on the Sea of Galilee at Samakh, and runs up the valley of the Yarmuk to join, at ed-Der‘a, the line of the third railway. This was undertaken in 1901 to connect Damascus with Mecca; in 1906 it was finished as far as Ma‘an, and in 1908 the section to Medina was completed. Carriage-roads also began to be constructed during the last decade of the 19th century. They are on the whole carelessly made and maintained, and are liable to go badly and more or less permanently out of repair in heavy rain. Of completed roads the most important are from Jaffa to Haifa, Jaffa to Nablus, Jaffa to Jerusalem, Jaffa to Gaza; Jerusalem to Jericho, Jerusalem to Bethlehem with a branch to Hebron, Jerusalem to Khan Labban —ultimately to be extended to Nablus; and Gaza to Beersheba. Other roads have been begun in Galilee (e.g. Haifa to Tiberias and to Jenin); but in this respect the northern province is far behind the southern. For the rest there is a network of tracks, all practically impassable by wheeled vehicles, extending over the country and connecting the towns and villages one with another. Industries —There are no mines and few manufactures of impor- tance in Palestine: the country is entirely agricultural. Although the processes are primitive and improvements are discouraged, both by the policy of the government and by an indolence and suspiciousness of innovation natural to the people themselves, fine crops of cereals are yielded, especially in the large wheat-lands of Hauran. Besides wheat, the following crops are to a greater or less extent cultivated—barley, millet, sesame, maize, beans, peas, lentils, kursenni (a species of vetch used as camel-food) and, in some parts of the country, tobacco. The agriculturist has many enemies to contend with, the tax-gatherer being perhaps the most deadly; and drought, earthquakes, rats and locusts have at all periods been responsible for barren years. The fruit trade is very considerable. The value of the oranges exported from Jaffa in 1906 was £162,000; this amount increases annually, and of course in addition a considerable quantity is retained for home consumption. Besides these are grown melons, mulberries, bananas, apricots, quinces, walnuts, lemons and citron. The culture of the vine—formerly an important staple, as is proved by the countless ancient wine-presses scattered over the rocky hillsides of the whole country—fell to some extent into desuetude, no doubt owing to the Moslem prohibition of wine-drinking. It is, however, rapidly returning to favour, principally under Jewish auspices, and numerous vineyards now exist at different centres. All over the country are olive-trees, the fruit and oil of which are a staple product of the country; the trade is however hampered by an excessive tax on trees, which not only discourages plantation, but has the unfortunate effect of encouraging destruction. Other fruit trees are abundant, though less so than those we have men- tioned: such are pomegranates, pears, almonds, peaches, and, in the warmer part of the country, palms. Apples are few and poor in quality. The kharrub (carob) is common and yields a fruit eaten by the poorer classes.?, Of ordinary table vegetables a considerable quantity and variety are grown: such are the cabbage, cauliflower, solanum (egg-plant), cucumber, hibiscus (bamieh), lettuce, carrot, artichoke, &c. The potato is also grown in considerable quantities. Beside the agricultural there is a considerable pastoral industry, though it is principally confined to production for home consump- tion. Sheep and goats are bred throughout the country; but the breeding of the beasts of burden (donkeys, horses, camels) is chiefly in the hands of the Bedouin. Of the manufactures the following call for mention: pottery (at Gaza, Ramleh and Jerusalem); soap (from olive oil, principally at Nablus); we may perhaps also extend the term to include the collecting of salt (from the Dead Sea). This last is a government monopoly, but illicit manufacture and smuggling are highly organized. Some of the minor industries, such as bee-keeping, are practised with success by a few individuals. Other industries of less importance are basket-making, weaving, and silk and cotton 1 A sanjak is usually a subordinate division of a vilayet, but that of Jerusalem has been independent ever since the Crimean War. This change was made on account of the trouble involved in referring all complications (arising from questions relating to the political standing of the holy places) to the superior officials of Beirut or Damascus, as had formerly been necessary. 7 Sometimes imagined to be the “locusts” eaten by John the Baptist, on which account the tree is often called the locust-tree. But it was the insect which John used to eat; it is still eaten by the fellahin. PALESTINE 605 manufacture. Stone-quarrying has been fostered since 1900 by the great development of building at Jerusalem and other places. _ Wine is manufactured by several of the German and Jewish colonies, and by some of the monastic establishments. Regular industrial work is however handicapped by competition with the tourist trade in its several branches—acting as guides and camp servants, manu- facture and sale of ‘‘ souvenirs ” (carved toys and trinkets in mother- of-pearl and olive-wood, forged antiquities and the like), and the analogous trade in objets de piété (rosaries, crosses, crude religious pictures, &c.) for pilgrims. Travellers in the country squander their money recklessly, and these trades, at once easy and lucrative, are thus fatally attractive to the indolent Syrian and_ prejudicial to the best interests of the country. CR AS Me) HIsTOoRY I.—Old Testament History. Palestine is essentially a land of small divisions, and its configuration does not fit it to form a separate entity; it ‘‘ has never belonged to one nation and probably never will.’”* Its position gives the key to its history. Along the west coast ran the great road for traders and for the campaigns which have made the land famous. The seaports (more especially in Syria, including Phoenicia), were well known to the pirates, traders and sea-powers of the Levant. The southernmost, Gaza, was joined by a road to the mixed peoples of the Egyptian Delta, and was also the port of the Arabian caravans. Arabia, in its turn, opens out into both Babylonia and Palestine, and a familiar route skirted the desert east of the Jordan into Syria to Damascus and Hamath. Damascus’ is closely connected with Galilee and Gilead, and has always been in contact with Mescpotamia, Assyria, Asia Minor and Armenia. Thus Pales- tine lay at the gate of Arabia and Egypt, and at the tail end of a number of small states stretching up into Asia Minor; it was encircled by the famous ancient civilizations of Babylonia, Assyria, South Arabia and Egypt, of the Hittites of Asia Minor, and of the Aegean peoples. Consequently its history cannot be isolated from that of the surrounding lands. Recent research in bringing to light considerable portions of long-forgotten ages is revolutionizing those impressions which were based upon the Old Testament—the sacred writings of a small fraction of this great area; and a broad survey of the vicissitudes of this area furnishes a truer perspective of the few centuries which concern the biblical student. The history of the Israelites is only one aspect of the history of Palestine, and this is part of the history of a very closely interrelated portion of a world sharing many similar forms of thought and custom. It will be necessary here to approach the subject from a point of view which is less familiar to the biblical student, and to treat Palestine not merely as the land of the Bible, but as a land which has played a part in history for certainly more than 4000 years. The close of Old Testament history (the book of Nehemiah) in the Persian age forms a convenient division between ancient Palestine and the career of the land under non-oriental influence during the Greek and Roman ages. It also marks the culmination of a lengthy historical and religious development in the establish- ment of Judaism and its inveterate rival Samaritanism. The most important data bearing upon the first great period are given elsewhere in this work, and it is proposed to offer here a more general survey.® To the prehistoric ages belong the palaeolithic and neolithic flints, from the distribution of which an attempt might be made to give a synthetic sketch of early Palestinian man.® A burial cave at Gezer has revealed the. existence of a race of slight build and stature, muscular, with elongated crania, and thick and heavy skull-bones. The 3G, A. Smith, Hist. Geog. of the Holy Land, p. 58. This and the author’s art. ‘‘ Trade and Commerce,”’ Ency. Bib. vol. iv., and his Jerusalem (London, 1907), are invaluable for the relation between Palestinian geography and history. For the wider geographical relations, see especially D. G. Hogarth, Nearer East (London, 1902). ‘See especially the writings of H. Winckler, in the 3rd ed. of Schrader’s Keilinschriften und das Alte Test. (Berlin, 1903); his Religionsgeschichtlicher u. geschichtlicher Orient (1906), &c. 5 See the articles on the surrounding countries and peoples, and, for the biblical traditions, art. Jews. 6See H. Vincent, Canaan d’apres l’exploration récente (Paris, 1907), pp. 374 sqq., also pp. 392-426. Beginning of history. ''606 people lived in caves or rude huts, and had domesticated animals (sheep, cow, pig, goat), the bones of which they fashioned into various implements. Physically they are quite distinct from the normal type, also found at Gezer, which was. taller, of stronger build, with well-developed skulls; and is akin both to the Sinaitic and Palestinian type illustrated upon Egyptian monuments from ¢. 3000 B.c., and to the modern native! The study of Oriental ethnology in the light. of history is still very incomplete, but the regular trend of events points to a mixture of races from the south (the home cf the Semites) and the north. At what period Palestine first became the “ Semitic” land, which it has always remained, is uncertain; nor can one decide whether the characteristic megalithic monuments, especially to the east of the Jordan, are due to the first wave which introduced the Semitic. (Canaanite) dialect and the place-names. At all events during the last centuries of the third millennium B.c., remarkable for the high state of civilization in Babylonia, Egypt and Crete, Palestine shares in the active life and intercourse of. the age; and while its fertile fields are visited by Egypt, Babylonia (under Gimil-Sin, Gudea and Sargon) claims some supremacy over the west as far as the Mediterranean. A more definite stage is reached in the period of the Hyksos (c. 1700), the invaders of Egypt, whose Asiatic origin is sug- gested inter alia by the proper-names which include “ Jacob,” and <‘Anath *? as, deities.? ., After. their expulsion it. is very significant to find that. Egypt forthwith enters upon a series of campaigns in Palestine and Syria as far as the Euphrates, and its successes over a, district whose political fate was bound up with Assyria and Asia Minor laid the foundation of a policy which became traditional. Apart from rather disconnected details which belong properly tothe history of Babylonia and Egypt, it-is not until about the 16th century B.c. that Palestine appears in the clear light of history, and henceforth, its course can be traced with some sort of con- tinuity., Of fundamental importance are the Amarna cuneiform tablets discovered in 1887, containing some of the political correspondence between. Western Asia and. Egypt for a few years of the reigns of Amenophis III. and IV. (c. 1414-1360).* The first Babylonian dynasty, now well known for its Kham- murabi, belonged to the past, but the cuneiform script and language are still used among the Hittites of Asia Minor (centring at Boghaz-keui) and the kings of Syria and Palestine. Egypt itself was now passing from its greatness, and the Hittites (g.v.)—the term is open to some criticism—were its rivals for the possession of the. intervening lands. , Peoples (apparently Iranian) of Hittite connexion from the powerful state of Mitanni (Northern Syria and Mesopotamia) had already left. their mark as far south as Jerusalem, as may be inferred from the personal names,? and to the intercourse with (apparently) Aegean culture. revealed by excavation, the letters add references to. mercenaries and bands from Meluhha_ (viz. Arabia), Mesopotamia and the Levant. The diminutive cities of this cosmopolitan Palestine were ruled by kings, not necessarily of the native stock; some were, appointed—and even anointed—by the Egyptian king, and the small extent of these city-states is obvious from the references to the kings of such near-lying sites as Jerusalem, Gezer, Ashkelon and Lachish. Torn by mutual jealousy and intrigue, and forming little confederations among Seyptian suzerainty, 1 For fuller treatment of the data see R. A. S. Macalister’s complete memoir of the Gezer éxcavations. cae 2 Reference may! be made to Ed. Meyer’s admirable survey of Oriental history down to this age, Gesch. d. Altertums. (Berlin, 1909), also. to J. H. Breasted, Hist. of Egypt (London, 1906), bks. iiv.; and L. W. King, Hist. of Bab. and, Ass. vol. i. (London, 1910). Some knowledge of the culture, religion, history and interrelations over the area of which Palestine formed part is indispensable for any careful study of the ages upon which we now enter. ® See the admirable edition by J. A. Knudtzon, with full notes by O. Weber (Leipzig, 1907-1910). For their bearing on Palestine, see especially P. Dhorme, Rev. biblique (1908), pp. 500-519; (1909), PP. 50-73, 368-385. . *Dhorme, op. cit. (1909), pp. 60 sqq.;H. R. Hall, Proc. Soc. Bibl. Arch, (1909), xxxi. 233 seq.; Weber, op. cit., p. 1088 seq.; ef. A. H. Sayce, Arch. of Cuneiform Inscr. (1907), pp. 193 sqq. PALESTINE — anatase [OLD TESTAMENT HISTORY | themselves, they were united by their common recognition of | the Egyptian ‘suzerain, their court of appeal, or in some short- - lived attempt to withstand him. Apart from Jerusalem and | a few towns on the. coast, the real weight lay to the north, and | especially in the state of Amor.® It is an age of internal dis- i organization and cf heavy pressure by land and by sea from | Northern Syria and Asia Minor. The land seethes with excite- | ment, and Palestine, wavering between allegiance to Egypt and | intrigues with the great movements at its north, is unable to_ take any independent line of action: The letters vividly describe _ the approach of the enemy, and, in appealing to Egypt, abound | in protestations of loyalty, complaints of the disloyalty of other kings and excuses for the writers’ suspicious conduct. Of exceptional interest are the letters from Jerusalem describing | the hostility of the maritime coast and the disturbances of the Habiru, (“ allies ””), a name which, though often equated with that of the Hebrews, may have no ethnological or: historical significance. But Egypt was unable ‘to’ help the loyalists, even ancient Mitanni lost its political independence, and the supremacy of the Hittites was assured... The history of the age illustrated by the’ Amarna letters is continued in the tablets found at Boghaz-keui, the capital of the old Hittite Empire.” Subsequent Egyptian evidence records that Seti I. (c. 1320) of the XIXth Dynasty. led. an, expedition. into .Palestine, but struggles with the Hittites continued until Rameses II.-(c. 1300) concluded with them an elaborate treaty ‘which ‘left him ‘little more than Palestine. Even this province was with ‘difficulty maintained: the disturbances in the Levant. and in Asia Minor (which beiong to Aegean and Hittite history). and the revival of Assyria were reshaping the political history of Western: Asia. Under Rameses IIT. (c. 1200-1169) we may recognize another age of disorganization in Palestine, in the movements with which the, Philistines (g.v.) were concerned. . Nevertheless, Egypt seems to have enjoved a fresh spell of extended supremacy, and Rameses apparently succeeded in recovering Palestine and some part of Syria.. But it was the close of.a lengthy period during which Egypt-had endeavoured to keep Palestine detached from Asia, and Palestine had realized the significance of a powerful empire at its south-western border. ‘Somewhat later Tiglath-Pileser (c. 1100) pushed the limits of Assyrian suzerainty westwards over the lands formerly held by the great Hittite Empire. It is at this age, when the external evidence becomes extremely fragmentary, that new political’ movements were inaugurated and new confederations of states sprang into existence. . Palestine had ,been politically part of Egypt. or. of the Hittite Empire; we now reach the stage where it becomes more closely identified with Israelite history. Palestine had not as yet been absorbed by any of the great powers with whose history and culture it had been. so closely bound up for so many centuries. In the ““Amarna’”’ age the little kings had a certain measure of inde- pendence, provided they guarded the royal caravan routes, paid tribute, refrained from. conspiracy, and generally supported their suzerain and his agents... However profound the influence of Babylonia’ may have been, excavation has discovered comparatively few specific traces of it. Although cuneiform was used, the Palestinian letters show that the native language, as in the case of earlier proper-names, was most nearly akin to the later “‘ Canaanite”? (Hebrew, Moabite and Phoenician). In view of the relations subsisting among Pales- tine, Mitanni and the Hittites, it is evident that Babylonian 5 Amor (Ass. Amurru, Bibl. Ameriie), lay north of Lebanon and behind Phoenicia; but the term fiuctuates (Weber, of. cit., 1132 sqq.). See art. AMorITEs, and A. T. Clay, Amurru (Phila- deiphia, 1909). : 6 See H. Winckler, Altor. Forschung. (1902), iii. 22; W. M. Miller in J. Benzinger, Heb. Archéol..(1907), p. 445; B. Eerdmans, Alitest. Stud. (1908), ii. 61 sqq.; Dhorme, op. ¢zt: (1909), pp. 677 sqq. The movement of the Habiru cannot be isolated from that represented in other letters (where the enemy are not described by this term), and their steps do not agree with, those of the invading Israelites in the book of Joshua (g.z.). ‘ My HL Ra ay Mitieil d. deutschen Orieni-Gesell. z. Berlin (1907) No. 35; cf. J. Garstang, Land of Hittites (London, 1910), 326 sqq. TheAmarna Period. '' OLD TESTAMENT HISTORY] influence could have entered indirectly; and until one can determine how much is specifically Babylonian the analogies and parallels cannot be made the ground for sweeping assertions. The influence of a superior power upon the culture of a people cannot of coursé be denied; but history proves that it depends upon the resemblance between the two peoples and. their respective levels of thought, and that it is not necessarily either deep or lasting.’ A better case might be made for Egypt; yet notwithstanding the presence of its colonies, the cult of its gods, the erection of temples or shrines, and the numerous traces of intercourse exposed by excavation, Palestine was Asiatic rather than Egyptian. Indeed Asiatic influence made itself felt in Egypt before the Hyksos age, and later, and more strongly, during the XVIIIth and following Dynasties, and deities of Syro-Palestinian fame ‘(Resheph, Baal, Anath, the Baalath of Byblos, Kadesh, Astarte) found a hospitable welcome. On the whole, there was everywhere a common foundation of culture and thought, with local, tribal and national develop- ments; and it is useful to observe the striking similarity of religious phraseology throughout the Semitic sources, and its similarity with the ideas in the Egyptian texts. And this becomes more instructive when comparison is made between cuneiform or Egyptian’ sources extending over many centuries and ‘particular groups of evidence’ (Amarna letters, Canaanite and Aramaean inscriptions, the Old Testament and later Jewish literature to the Talmud), and pursued to the customs and beliefs of the same area to-day. The result is to emphasize (a) the inveterate ‘and indissoluble connexion between religious, social and political life, (b) the differences between the ordinary current religious conceptions and specific positive developments of them, and (c) the vicissitudes of these particular growths in their relation to history.? There is reason to believe that the religion of Palestine in the Amarna age was no inchoate or inarticulate belief; like the material culture it had passed through the elementary stages and was a fully established though not, perhaps, a very advanced organism. There were doubtless then, as later, numerous local deities, closely connected with local districts, differing perhaps in name, but the centre of similar ideas as regards their relations to their worshippers. Com- mercial and political intercourse had also brought a knowledge of other deities, who were worth venerating, or who were the survivors of a former supremacy, or whose recognition was enforced. It is particularly interesting to find in the Amarna letters that the supremacy of Egypt meant also that of the national god, and the loyal Palestinian kings acknowledge that their land belonged to Egypt’s king and god. In accordance with what is now known to be a very widespread belief, the kingship was a semi-divine function, and the Pharaoh was the incarnation of Amon-Re. This would bring a greater coherence of worship among the chaos of local cults. The petty kings naturally recognize the identity of the Pharaoh, and they hail him as their god and identify him with the heads of their own pantheon. Thus he is called—in the cuneiform letters—their Shamash or their Addu. The former, the sun-deity, god of justice, &c., was already well known, to judge from Palestinian place-names (Beth-Shemesh, &c.). The latter, storm or weather god, or, in another aspect, god of rain and therefore of fertility, is specifically West Asiatic, and may be equated with Hadad and Ramman (see below). He is presumably the Baal who is associated with thunder and lightning, and with the bull, and who was familiar to the Egyptians of the XIXth and XXth Dynasties in the adulations of their divine king. He is probably also ‘‘ the lord of the gods ” (the head of a pantheon) invoked in a private cuneiform tablet unearthed at Taanach.? Besides these gods, and others whose fame may be inferred (Dagon, 1 Much confusion can be and has been caused by disregarding#()) and by supposing that the appearance of similar elements of thought or custom implied the presence of similar more complete organisms (e.g. totemism, astral religion, jurisprudence). Cf. p.182,n.4. ~ 2See, most recently, Ungnad’s translation in H. Gressmann, Ausgrabungen in Pal. u. d. A. T. (Tubingen, 1908), p. 19 seq. The title ‘‘ lord of heaven ’’—whether the Sun or Addu, there was a Religion. PALESTINE 6047 Nebo, Nergal, &c.), there’ were the closely-related goddesses Ashira and Ishtar-Astarte (the Old Testament ‘Asherah and Ashtoreth). Possibly the name Yahweh (see JEHovAH) had already entered Palestine, but it is not prominent, and if, as in the case of certain other deities, the extension of the name and cult went hand-in-hand with political circumstances, these must be sought in the problems of the Hebrew monarchy.’ At an age when there were no great external empires to control Palestine the Hebrew monarchy arose and claimed a premier place amid its neighbours (c.' 1000). “How the small’ Rise of the rival districts with their petty kings were united Hebrew into a kingdom under’ a’ single head is a disputed Momarchy. question; the stages from ‘the half-Hittite, half-Egyptian land to the independent Hebrew state with its national god are an unsolved problem. Biblical tradition quite plausibly represents a mighty invasion of tribes who had come from Southern Palestine and Northern Arabia (Elath, Ezion-geber)—but primarily from Egypt—and, after a series of national “ judges,” established the kingship. But no place can be found for this conquest, as it is described, either before the '‘‘ Amarna ” age (the date, following 1 Kings vi. 1) or about the time of Rameses II. and Mineptah (see: Exod. i. rz); and if the latter king (c. 1244) records the subjugation of the people (? or land) “Israel,” the complicated history of names does not guarantee the absolute identity of this “‘ Israel’ either ‘with the pure Israelite tribes which invaded the land or with the intermixed people after this event (see Jews: §§ 6-8). Whatever may have been the extent of this invasion and the sequel, the rise and persistence of an inde- pendent Palestinian kingdom was an event which concerned the neighbouring states. Its stability and the necessary furtherance of commerce, usual among Oriental kings, depended upon the attitude of the maritime coast (Philistia and Phoenicia), Edom, Moab, Ammon, Gilead and the Syrian states; and the biblical and external records for the next four centuries (to 586) fre- quently illustrate situations growing out of this interrelation. The evidence of the course of these crucial years is unequal and often sadly fragmentary, and is more conveniently noticed in connexion with the biblical history (see Jews: §§ 9-17). A conspicuous feature is the difficulty of maintaining this single monarchy, which, however it originated, speedily became two rival states (Judah and Israel). These are separated by a very ambiguous frontier, and have their geographical and political links to the south and north respectively. The balance of power moves now to Israel and now to Judah, and tendencies to internal disintegration are illustrated by the dynastic changes in Israel and by the revolts and intrigues in both states. As the power of the surrounding empires revived, these entered again into Palestinian history. As regards Egypt, apart from a few references in biblical history (e.g. to its interference in Philistia and friendliness to Judah, see PHILISTINE), the chief event was the great invasion by Sheshonk (Shishak) in the latter part of the roth century; but although it appears to be an isolated campaign, contact with Egypt, to judge from the archaeological results of the excavations, was never intermittent. The next definite stage is the dynasty of the Israelite Omri (¢.2.), to whom is ascribed the founding of the city of Samaria. The dynasty lasted nearly half a century, and is contemporary _with the expansion of Phoenicia, and’ presumably therefore with some prominence of the south maritime coast. The royal houses of Phoenicia, Israel and Judah were united by inter- marriage, and the last two by joint undertakings in trade and war (note also 1 Kings ix. 26 seq.). Meanwhile Assyria was gradually establishing itself westwards, and a remarkable confederation of the heirs of the old Hittite kingdom, “kings of the land of Hatti” (the Assyrian term for the Hittites) was formed to oppose it. Southern Asia Minor, Phoenicia, Ammon, the Syrian Desert and Israel (under Omri’s son “‘ Ahab the Israelite’) sent: their troops to support Damascus which, in spite of the repeated efforts of Approach ; of Assyria. tendency to identify them—was perhaps known in Palestine, as it certainly was in Egypt and among the Hittites. 3 See S. A. Cook, Expositor, Aug. 1910, pp. I11-127. ''608 Shalmaneser,! was evidently able to hold its own from 854 to 839. The anti-Assyrian alliance was, as often in west Asia, a temporary one, and the inveterate rivalries of the small states are illustrated, in a striking manner, in the downfall of Omri’s dynasty and the rise of that of Jehu (842-c. 745); in the bitter onslaughts of Damascus upon Israel, leading nearly to its annihilation; in an unsuccessful attack upon the king of Hamath by Damascus, Cilicia and small states in north Syria; in an Israelite expedition against Judah and Jerusalem (2. Kings xiv. 13 seq.);. and finally in the recovery and extension of Israelite. power—perhaps to Damascus—under Jeroboam II. In such vicissitudes as these Palestinian history proceeds upon a much larger scale than the national biblical records relate, and the external evidence is of the greatest importance for the light it throws upon the varying situations. Syria could control the situation, and it in turn was influenced by the ambitions of Assyria, to whose advantage it was when the smail states were rent by mutual suspicion and hostility. It is possible, too, that, as the states did not scruple to take advantage of the difficulties of their rivals, Assyria played a more prominent part in keeping these jealousies alive than the evidence actually states. Moreover, in the light of these moves and counter- moves one must interpret the isolated or incomplete narratives of Hebrew history.2. The repeated blows of Assyria did not pre- vent the necessity of fresh expeditions, and later, Adad-Nirari ITI. (812-783). claims as tributary the land of Hatti, Amor, Tyre, Sidon, ‘“‘the land of Omri’’ (Israel), Edom and. Philistia. Israel at the death of Jeroboam was rent by divided factions, whereas Judah (under Uzziah) has now become a powerful kingdom, controlling both Philistia and the Edomite port. of Elath on the gulf of ‘Akaba. The dependence of Judaean sovereignty upon these districts was inevitable; the resources of Jerusalem obviously did not rely upon the small district of Judah alone. If Ammon also was tributary (2 Chron. xxvi. 8, xxvii.), dealings with Israel and perhaps) Damascus could probably be inferred. A new period begins with Tiglath-Pileser IV. (745-728): pro- and anti-Assyrian parties now make themselves felt, and Predomi- When north Syria was taken in 738, Tyre, Sidon, nanceof Damascus. (under. Rezin), ‘‘ Samaria’ (under Assyria. Menahem) and a queen of Aribi were among the tribu- taries. It is possible that Judah (under Uzziah and Jotham) had come to an understanding with Assyria; at all events Ahaz was at once encircled by fierce attacks, and was only saved by Tiglath-Pileser’s campaign against Philistia, north Israel and Damascus.. With the siege: and fall of Damascus. (733-32) Assyria gained the north, and its supremacy was recognized by the tribes of the Syrian desert and Arabia (Aribi, Tema, Sheba). In 722 Samaria, though under an Assyrian vassal (Hoshea the last king), joined with Philistia in revolt; in 720 it was allied with Gaza and Damascus, and the persistence of unrest is evident when Sargon in 715 found it necessary to transport into Samaria various peoples of the desert. Judah itself was next involved in an anti-Assyrian league (with Edom, Moab and Philistia), but apparently submitted in time; nevertheless a decade later (7o1), after the change of dynasty in Assyria, it participated in a great but unsuccessful effort from Phoenicia to Philistia to shake off the yoke, and suffered disastrously.* With the crushing blows upon Syria and Samaria the centre of interest moves southwards and the history is influenced by Assyria’s rival Babylonia (under. Marduk-baladan and_ his successors), by north Arabia and by Egypt. Henceforth there is little Samarian history, and of Judah, for nearly a century, few political events are recorded (JEws: § 16). Judah was under Assyrian supremacy, and, although it was in- volved with Arabians in the revolt. planned by Babylonia 1 Recently found to be the third of that name (H. W. Hogg, The Interpreter, 1910, p. 329). 2So e.g. in references to Ammon, Damascus and Hamath, and in Judaean relations with Philistia, Moab and Edom. 3 See art. HEZEKIAH. A recently published inscription of Sen- nacherib (of 694 B.C.) mentions enslaved. peoples from Philistia and Tyre, but does not name Judah. PALESTINE [OLD TESTAMENT HISTORY (against Assurbanipal), it appears to have been generally quiescent. At this stage disturbances, now by Aramaean tribes, now by Arabia, combine with the new rise of Egypt and the weakness of Assyria to mark a turning-point in the world’s history. Psammetichus (Psamtek) I. (663-609) with ne eee his Greeks, Carians, Ionians and soldiers from Pales- ~?” tine and Syria, re-established once more an Egyptian Empire, and replaced the fluctuating relations between Palestine and the small dynasts of the Delta by a settled policy. Trading intercommunication in the Levant and the constant passage to and fro of merchants brought Egypt to the front, and, in an age of archaic revival, the effort was made to re-establish the ancient supremacy over Palestine and Syria. The precise meaning of these changes for Palestinian history and life can only incompletely be perceived, and even the significance of the great Scythian invasion and of the greater movements with which it was connected is uncertain (see ScyrTuta). At all events, Egypt (under. Necho, 609-593) prepared to take advantage of the decay of Assyria, and marched into Asia. Judah (under Josiah) was overthrown at Megiddo, where about nine centuries previously the victory of Tethmoses (Thutmose) III. had made Egypt supreme over Palestine and Syria. But Egypt was now at once confronted by the Neo-Babylonian or Chaldean Empire (under Nabopolassar), which, after annihilating Assyria with the help of the Medians, naturally claimed a right to the Mediterranean coast-lands. The defeat of Necho by Nebuchad- rezzar at Carchemish (605) is one of the world-famous battles. Although Syria and Palestine now became Babylonian, this revival of the Egyptian Empire aroused hopes in Judah of deliverance and led to revolts (under Jehoiachin and Zedekiah), in which Judah was apparently not alone.*. They culminated in the fall of this kingdom in 586. Henceforth the history of Palestine is disconnected and fragmentary, and the few known events of political importance are isolated and can be supplemented only by infer- ences from the movements of Egypt, Philistia or Phoenicia, or from the Old Testament. According to the Chaldean Nabonidus (553) all the kings from. Gaza to the Euphrates assisted in his buildings, and the Chaldean policy generally appears to have been favourable towards faithful vassals. Cyrus meanwhile was rising to lead the Persians against Media. After a career of success he captured Babylonia (553) and forth- with claimed, in his famous inscription, the submission of Amor. For the next 200 years Palestine remained part of the new Persian Empire which, with all its ramifications on land and on sea, embraced the civilized world from the Himalayas to the Levant, until the advent of Alexander the Great (see Jews: § 19). Very gradually the face of history underwent a complete change. Egypt had resumed its earlier connexions with the Levantine heirs of the ancient Aegeans, the old empires of the Nearer East had practically exhausted themselves, and Palestine passed into the fresh life and thought of the Greeks. (See below, p. 617.) In any consideration of the internal conditions in Palestine it must be observed that there is a continuity of thought, custom and culture which is independent of political changes saternai and vicissitudes of names. With the establishment. Conditions. of an independent monarchy Palestine did not enter ae en into a new world. Whatever internal changes sees ensued between the “‘ Amarna”’ age and tooo B.c., they have not left their mark upon the course of culture illustrated by the excavations. These still indicate communication with Egypt and the north (Syria, Asia Minor; Assyria and the Levant not excluded), and even when a novel culture presents itself, as in certain graves at Gezer, the affinities are with Cyprus and Asia Minor (Caria) of about the 11th or roth century.’ The use of 4Cf, Jer. xxvii. 2.seq., and the history of the Egyptian Hophra (Apries, 588-569). tewvd.b ® At present it is difficult as regards Palestine to distinguish Aegean influence (direct and indirect) from that of Asia Minor generally. Only after the old Gretan (Minoan) culture had passed its zenith and was already decadent does it suddenly appear in Cyprus (H. R. Hall, Proc. Soc. Bibl. Arch. xxxi. 227). Babylonian Empire. '' '' ''MEMOIRS OF THE AMERICAN MUSEUM OF NATURAL HISTORY. PART III— CRANIOMETRY OF THE EQUIDA. By Henry FAIRFIELD OSBORN. CONTENTS. Paar Introduction : ; : s ; : ; : : . : : i : ; ‘ ‘ ; : Se Dolichocephaly, brachycephaly, cytocephaly . : : j ‘ : ; : j : : : ; Be Indices, ratios, absolute measurements . : : i : : ; ; ; : ; : 3 : 58 Growth stages of the horse skull . : : : : : : ; : : : : ; ; : 60 I. Craniometric Systems : : : ; | : ; ; : : : : : : : 61 Franck, 1875, page 62 Salensky, 1902, page 66 Branco, 1883, ts 64 Ewart, 1907, ~ OG Nehring, 1884, “65 Bradley, 1907, i 6S Tscherski, 1892, 205 : Osborn, 1912, eS OS II. Distinctions, horses, asses, zebras ; : : 3 oe : : : : : : : ; 88 Authors: Branco, 1883, page 88 Salensky, 1907, : page 89 Dugés, 1898, on OO Chubb, 1911, 7.90) Occiput-vertex Angle, horses, asses, zebras. ; : : ; : ; : i ; : é i 90 Naso-frontal Sutures Be a as ; : : : : : : : ; ; ; : : 92 III. Cytocephaly, bending of face on cranium . : i : : ‘ ‘ : : : : ; : 92 Authors: Riitimeyer, 1882, page 93 Lankester,1902, page 93 Flower, 1891, fe Ge Ewart, 1907, Ce Of Cytocephaly in the Artiodactyla. ; : i ; : : : é ‘ : : : ; 94 Palato-cranial angle in Horses, asses, zebras . ‘ : ; : : : ; 4 : é : : 96 IV. Craniometry and odontometry in paleontology . ; : : ; : ; ; ; } ; ‘ 96 INTRODUCTION. A promising as well as most difficult problem in modern mammalogy is the phylogeny of the Equide. Skull measurements and proportions represent one of many systems of cross- tests of affiliation and phyletic relationship which may be applied. In 1902 the author pointed out the manifold effects of dolichocephaly and brachycephaly in the mammals.! Among the horses, as among the Oligocene titanotheres (see Figs. 1, 2), progressive brachycephaly or dolichocephaly sharply distinguishes certain phyla. Ewart 1‘ Dolichocephaly and Brachycephaly in the Lower Mammals.’ Bull. Amer. Mus. Nat. Hist., Vol. XVI, Art. vir, Feb. 3, 1902, pp. 77-89. {57} ''58 OSBORN, CRANIOMETRY OF THE EQUID. especially has pointed out the importance of the bending down of the face upon the cranium in the horses, a transformation to which Osborn has applied the term cytocephaly.' It is also very important to distinguish (see Fig. 2) between prodpic dolichocephaly (Osborn), in which the face is elongated, and opisthopic dolichocephaly (Osborn), in which the cranium is elongated. A line drawn through the postorbital process divides the face from the cranium. The chief changes of skull proportion in Ungulates thus come under the following heads: Brachycephaly, general broadening of the skull as whole. Dolichocephaly, general lengthening of the skull as a whole. Proopic dolichocephaly, elongation of the facial region. Opisthopic dolichocephaly, elongation of the cranial region. Cytocephaly, upward or downward flexure of the facial upon the basicranial axis. The index is a far better method of expressing these transformations than any system of Earlier students of the skull of the horse, like Franck (1875) relied It is obvious that among domesticated as among wild horses varia- absolute measurements. principally on percentages. tions in size due to age, sex, favorable or unfavorable environment, vitiate any system of absolute measurements. Non-blending or pure inheritance indices in the Skull of the Horse, Ass, Mule. Ass 46.9-49.9 Mule 40.8-43.6 Horse 40 .4—-44.1 Ass 15.6-17.6 Width of skull * 100 lic Index: 1. Cephalic Index Basilar length Diastema X 100 2. Diastema Index: 3. Cranio-facial Index: Basilar length of skull Length of cranium X 100 Mule 18.6-21.9 Horse 18 .2-23.0 Ass 56.3-61.0 Mule 48.9-51.8 Length of face Horse 5.3-49.9 96 .0-104 .2 : ; Ass ical d t f orbit X 100 4. Orbital Index: ee oe 5 ae aa Mule 78.7- 99.1 orizontal diameter Bone 842-03 5 Transverse diameter of M? < 100 Ass 15 .2-16.0 5. Molar Index: ee i : Mule 14.2-14.9 Total length of entire molar series Hone 13 9-15 7 Ass 52.5-60.0 Mule 61 .0-66.5 Horse 64 .0-76.5 6. Occiput-vertex angle Index: Angle between vertex of skull and line connecting most posterior points of occipital crest with condyles i. e., nearly all horse skulls will stand when set up on end, some mule skulls (one out of four), no ass skulls Ass 93.8-111.7 Mule 95.5-110.3 Horse 72.8- 86.5 Distance from palate to posterior end of vomer X 100 Vomer Index: - Distance from vomer to foramen magnum The indices, or proportions between various parts, are less influenced by environmental modification but are largely influenced by age; thus the facio-cranial index alters rapidly as the horse attains maturity. 1‘The Continuous Origin of Certain Unit Characters as Observed by a Palxontologist.’ Amer. Naturalist, Vol. XLVI, April, May, 1912, pp. 185-206, 249-278. ''OSBORN, CRANIOMETRY OF THE EQUID. 59 It also appears that the angle between the face and the cranium alters greatly during growth. That the index deserves more general use among mammalogists as of strong specific value is shown in the above table of comparison of the principal indices in the skulls of the ass , of the horse ”, and of the mule, taken from a recent paper by Osborn." Ge. Brach. Mes. & £9 Beck Mes. Dol. Fig. 1. Brachycephaly, mesaticephaly, dolichocephaly. Upper: as observed in the human cranium. Lower: as observed in the skull (7. e., cranium and face) of the titanotheres. The age of the animal measured is extremely important in its effect on all systems of measure- ment of both the skull and the teeth. The indices of brachycephaly and dolichocephaly, and the bending of the face on the cranium, or cytocephaly, as above noted all change with age. This last statement is demonstrated at the close of this paper. It is, therefore, extremely desirable to establish among the Equide a series of growth stages. The accompanying table, prepared under the writer’s direction by Messrs. 5. H. Chubb and J. W. Gidley, from the very large series of preparations by Mr. Chubb in the American Museum, is an attempt to establish a series of eleven age stages, based upon the all-important succession and wear of the teeth. 1 Amer. Naturalist, May, 1912, pp. 272, 278. ''60 OSBORN, CRANIOMETRY OF THE EQUID. Age Stages of the Horse. Milk Teeth and beginning of Permanent Teeth. Stage I.— A few days old. Milk series fully formed but unworn. Stage II.— About 10 months old. Summit of first permanent Molar just appearing above the bone. Milk Premolars all somewhat worn. First and second Incisors somewhat worn. Stage IIJ.— 12 or 13 months old. Summit of first permanent Molar at height of milk series, but unworn. First and second Incisors worn; third Incisor at height of others, but unworn. Stage IV.— About 18 months old. Summit of second permanent Molar just appearing above the bone. Milk Premolars all well worn. Third Incisor beginning to wear. Stage V.— About 2 years old. Summit of second permanent Molar at height of grinding series, but unworn. Milk Incisors all worn. Beginning of shedding stage. Stage VI.— About 33 years old. Summit of third permanent Molar just appearing above the bone. Shedding of last milk Premolars. Second and third Premolars beginning to wear. Permanent Teeth only, in Progressive Stages of Wear. Stage VII.— About 4 years old. Summit of third permanent Molar and fourth permanent Premolar at height of grinding series, but unworn. First permanent Incisor worn. : Stage VIII.— About 5 years old. Permanent teeth all in use. Fourth Premolar and third Molar but little worn. First and second Incisors worn; third beginning to wear. Stage IX.— Between 15 and 20 years old. All the teeth well worn. Upper Incisors have the external dental cavities (‘cups’) greatly’ reduced. Lower Incisors, external dental cavities (cups) have disappeared. Stage X.— Between 25 and 30 years. Second Premolar worn to near fangs. Tooth pattern of grinding teeth more or less obliterated. Incisors worn down short, their angle of meeting acute, and external dental cavities (‘cups’) entirely worn away. Stage XI.— Between 30 and 40 years. Premolars and first Molar worn to near fangs. Tooth pattern of grinding teeth more obliterated. Incisors worn down very short and their angle of meeting very acute. YOUNG OR IMMATURE HorsE WITH Mitk AND PERMANENT TEETH (Adolescent stages) ADULT OR Mature Horst WITH EFFECTIVE PERMANENT TEETH (Mature stages) Otp HorsrE WITH TEETH BECOMING LESS EFFECTIVE (Senescent stages) ''OSBORN, CRANIOMETRY OF THE EQUID. 61 I. CRANIOMETRIC SYSTEMS, 1875-1912. The present preliminary review of previous cranio-metric systems of the Equide is chiefly based upon the notes of the writer’s research assistant, Mrs. Johanna K. Mosenthal. It is historical, critical, and constructive. The diagrams are the work of Mr. Erwin 8. Christman. The systems reviewed are the following: Franck, 1875 Salensky, 1902 Branco, 1883 Ewart, 1907 Nehring, 1884 Bradley, 1907 Tscherski, 1892 Osborn, 1912 The system of measurements introduced by each of these authors is clearly set forth by means of a uniform set of diagrams and explanatory keys prepared under the writer’s direction by Mr. Erwin 8. Christman. In each table the common usage of a certain measurement by other authors is indicated by round brackets (). There is also inserted, in square brackets [ ], the usage and terminology proposed by Osborn in the present paper. Rangifer NN Cytocephaly () 2 a YD m Opisthopic : Brontother. Dol. N N ‘ Fig. 2. Dolichocephaly and Cytocephaly in the Ungulates. Upper: Rangifer, with slight facio-cranial deflection. Bubalis, with strong facio-cranial deflection. Middle: Opisthopic dolichocephaly in Brontotherium; proopic dolichocephaly in Equus; Lower: Mesaticephalic ancestors of Brontotherium (Eotitanops) and of Equus (Eohippus). The Osborn terminology thus indicated [] includes: (1) the adoption from other authors of what appear to be the most natural and really significant measurements, (2) the adoption of ''62 OSBORN, CRANIOMETRY OF THE EQUID. the terminology which has been most generally used rather than the introduction of a new termi- nology, (3) some new measurements of the skull and teeth which appear to be especially appli- cable to extinct as well as to recent horses. It is noteworthy that all previous craniometric systems have been based on recent horses : it is also noteworthy (p. 96) that some of the measurements and indices which are applicable to recent horses are not generally applicable to extinct horses, owing to accidents of preservation. Sources of Craniometric Error. There are two grand sources of error in measurements of the cranium and teeth in existing domestic horses: first, polyphyly, or the mixture of ancestral stocks, or breeds; second, coenogenesis, or progressive changes due to age. The ‘breed mixture’ is a factor almost impossible to avoid because, as shown especially by the researches of Ewart, most of our common horses of the present day contain the blood of four or five wild and remotely ancestral stocks. To this so-called blood mixture, representing actually the persistence of certain distinctive ‘unit characters,’ are probably due many of the so-called variations in the skulls of horses. ‘The only standard existing horse is the Przewalsky. Franck’s System, 1875.! (Text Fig. 3, page 70.) Materials. ‘The materials used in the pioneer treatise of Franck were clearly insufficient, and although most of his “points of distinction between different existing and prehistoric breeds of horses” are borne out by his measurements, the differences are too slight to carry much weight because they are based on so small a number of individuals, namely: Noric horses (2 not typical, according to Tscherski). Arab horses (2 not typical, according to Tscherski). Feldmochinger (1 not typical, according to Tscherski). Neolithic horses (from Pile-dwellings). Greek ponies. Nw W RP & All of the twenty-five measurements of Franck are computed in terms of percentage to the basilar length (see B, Fig. 3, 111). No measurement is given to determine the relative facial and cranial length or facio-cranial index, although one of the main differences between various races of horses lies in the proépic dolichocephaly. Of the eight or nine points of comparison between the skull of the horse and of the ass given by Franck, Tscherski, and Salensky, we find only those afforded by Franck’s ‘vomer index’ valuable. However, most of the important measurements taken by later authors are to be found in Franck excepting the facio-cranial above noted. Franck’s Noric Horse is equivalent to the ‘ Nordic’ horse of other authors, the ‘Forest Horse’ of Ewart, the occidental, or western, horse, as distinguished from the oriental, or eastern, horse. 1 ‘Win Beitrage zur Rassenkunde unserer Pferde,’ Landwirthschaftliche Jahrbiicher, 1875, Vol. IV. '' OSBORN, CRANIOMETRY OF THE EQUID. Franck’s Craniometry of Oriental and Occidental Horses." Note: — Measurements in percents, computed with basilar length taken as 100. Measurements Noric _ Arab 1. Length from foramen magnum to point between I, I a 100 (57.4) 100 (50.8) 2. From foramen magnum to posterior end of palatal suture 46.6 47.2 3. From foramen magnum to beginning of vomer 25.2 : 27.6 4. From posterior end of palatal suture to point between I, I 53.9 Done 5. From posterior end of palatal suture to beginning of vomer 22.0 2075 6. Width between maxillary crests at their origin 32.3 30.9 7. Width between canines Jel O 9.6 8. Greatest width of skull 39.3 40.1 9. Greatest width between orbital processes 40.8 | 41.7 10. Greatest distance between pterygoid processes of palatal bones 14.0 13.8 11. Length of molar row of upper jaw, not including P! 33.5 33.8 12. Greatest distance between M’, M’ 2551 28.2 tae Distance between P?, P?, anterior end 14.4 13.5 14. Straight line from middle of transverse process of occipital bone to point between I, I : 108.8 108.5 15. From middle of transverse process of occipital bone to tip of nasal bones (with tape measure) 98.8 97.3 16. From middle of transverse process of occipital bone to tip of nasal bones (straight line) 97.0 95.8 17. From middle of occipital bone to point between orbits 35.7 36.5 18. From middle of occipital crest to end of nasal process of frontal bone 49.3 51.9 19. Median length of nasal bones 50.7 46.5 20. Straight line between tip of nasal bones and I, I 22°6 22.2 21. Greatest breadth of cerebral portion of skull, above zygomatic processes of temporal bones 19.2 Des 22. Smallest breadth of skull capsule on small pterygoid foramina 1): 12 23. Greatest width on parietal bump | 16.6 18.2 24. Width between supraorbital foramina | 29.4 300k 25. Width between infraorbital foramina | 18.1 16.3 Noric is average of 5 individuals. Arab is average of 4 individuals. 1 Taken from Franck’s table in ‘Hin Beitrag zur Rassenkunde unserer Pferde,’ Landwirthschaftliche Jahrbiicher, Vol. IV, 1875. ''64 OSBORN, CRANIOMETRY OF THE EQUID. Franck’s conclusions. All modern horses are derived from two original races, ‘oriental’ and ‘occidental,’ which are respectively exemplified by the following types: Oriental. Occidental. Type: Arab. Type: Noriec. ‘Persian. “ Pinzgauer. = Sareek. “Flanders. © Kussian. *. Old Norman. “Hungarian. “Palfrey of knights. “ Feldmochinger. ‘“‘ Ardennes. ““ Neolithic horse (one). “Luxemburg. Ponies of Greece, China, Persia, etc. The testimony of philology goes to show that the Aryans possessed horses while they were still in Asia and that the horse migrated into Europe with them. The Arab horse is a branch of the ‘oriental stock’ that has been improved by breeding and that has spread over all countries. The occidental, or western, or ‘Noric’ horse may be identical with or descended from the E£. robustus Pomel of the Swiss Dwellings. The earliest mention of this animal is in the account by Rogers of the horses of Dacia and Panonia. The chief distinctions between these two great races are the following: Oriental. Occidental. Brachycephalic. Dolichocephalic. Profile straight or concave. Facial proportion developed at the expense of cranial. Of small size and swift motion. Profile rounded or sheep-like. More closely related to the ass. Size large, movements heavy and slow. Branco’s System, 1883.! (Text Fig. 4, page 72.) Materials. Branco’s object is to show the relation of Equus andium of the Upper Pliocene or Lower Pleistocene of South America to other Equide. The materials on which he based his thirty-two measurements of modern horses are insufficient. They consist of the following: 1 zebra. 4 asses (2 very aged). 3 Arab horses, the same individuals used by Franck Greek ponies, individuals used by Franck. Shetland ponies. 4 Pinzgaur horses. 5 foetal and young stages, ranging from 1 month to 23 years. Methods. Of Branco’s thirty-two measurements twenty are represented in the accompany- ing table. It will be observed that he does not introduce the facio-cranial index. While his measurements and indices are important in proving his points, they are of little general value as data by which we may distinguish the various breeds of horses, or distinguish the horses, asses, and zebras. Many of the measurements are so difficult to take with precision that there is great room for error, which is increased by the fact that the measurements are mostly in small numerals. Measurement No. 5, 7. e., the length of the maxillary behind m*, has little value because age probably has as much to do with this measurement as species or breed. 1 ‘Uber eine Fossile Saugethier Fauna von Punin bei Riobamba in Ecuador,’ Palaiont. Abhandl. Dames u. Kayser, Vol. I, No. 22, 4to., Berlin, 1883. : ''OSBORN, CRANIOMETRY OF THE EQUID. 65 From the foetal and young stages studied Branco constructs some interesting tables showing progressive changes, but since the breeds of the five individuals selected are not indicated, we cannot be certain that the differences are not due to breed as well as to age, since breed-variation in horses is so great. In short, Branco’s system stands apart from all the rest; he takes a different series of measurements, and there are no references to his memoir in the other later authors. His conclusions as to the affinities of H. andiwm are: (1) that it is nearer to H. caballus than E. asinus or E. burchelli; (2) that the trend of development in all South American Equide is more or less away from that of Z. caballus and the Diluvial forms of the Old World. Nehring’s System of 1884.' (Text Fig. 5, page 74.) Materials. Pleistocene horses of northern Europe. Methods. Nehring’s system of measurement is one of the foundations of modern crani- ometry; his measurements are less numerous, more simple, and more to the point than those of his predecessors; they bring out the essential and racial rather than the detailed characters. They especially form a basis for the systems of Tscherski and Salensky. Nehring’s system of indices is, however, faulty and misleading because he departs from the method used by anthropologists. Instead of dividing the lesser measurement by the greater, as in computing the cephalic index of the human skull wherein anthropologists divide the great- est width by the greatest length, Nehring reverses this system and divides the greatest length by the greatest width. For example, Nehring’s Index I. = basilar tenth x __ length of vertex x 100 Index Il a frontal width anterior segment of ocular line x_ 100° Index III = ~ posterior segment of ocular line The obvious disadvantage of Nehring’s system of indices is that the broad-skulled forms (brachycephaly) give the lowest figures while the narrow-skulled forms (dolichocephaly) give the highest figures, these indices being just the reverse of those obtained in the human skull. Unfortunately Nehring’s Index I, or Frontal Index, is used by Tscherski and Salensky. Similarly the ‘Preorbital’ and ‘Postorbital’ indices and Index III, the ‘Ocular Index,’ are not satisfactory as representing the proportions of the face and of the cranium or of the eye, since no allowance is made for the width of the skull. These indices have been criticised by both Tscherski and Salensky although, owing doubtless to the weight of Nehring’s authority, they are adopted by both these authors. Tscherski’s System, 1892.’ (Text Fig. 6, page 76.) The object of this very exhaustive monograph is to show the relation of the Post-Pliocene horses of northern Asia to other fossil and recent Equide and determine ultimately the origin of the modern horses. ~ 1 ‘Pogsile Pferde aus deutschen Diluvial-Ablagerungen und ihre Beziehungen zu den lebenden Pferden. Ein Beitrag zur Geschichte des Hauspferdes,’ Sonderabdr., Landwirthschaftl. Jahrbucher, 1884. : 2 ‘Beschreibung der Sammlung posttertidrer Saugethiere,’ Mém. Acad. Imp. d. Sci. St. Petersbourg, Sér. 7, Vol. XL, No. 1, 1892, pp. 1-511. ''66 OSBORN, CRANIOMETRY OF THE EQUIDZ. Materials. Measurements were taken of a very extensive series of Post-Pliocene horses. The measurements of the skull of £. przewalskii are based on a single young individual, yet they correspond quite closely to those given in Salensky’s subsequent memoir and point to the same conclusions. Only two zebras and one Arab horse are measured, but the data on his own materials are supplemented from the work of other authors. Post-Pliocene Eurasian horses, numerous specimens. E. przewalskit, 1 skull, young. E. ?2burchelli, 2 skulls. Arab horse, 1 skull. Methods. ‘Tscherski’s system as a whole is based on those of Franck and Nehring, from whom he adopted many features. The measurements are most exhaustive and the tables on account of the very multiplicity of measurements and confused arrangement, do not bring out very clearly the essential distinctions between the races. The age of the horses measured is not indicated. Nor can we adopt the facio-cranial index of Tscherski because he measures the face from the anterior border (9) of the orbit instead of measuring it from the posterior border, thus: oe) : : . Length of face to anterior border of orbit x 100 Tscherski’s facial index: Baglaeacneth of GEall (See Addendum, facing this page.) Salensky’s System of 1902.! (Text Fig. 7, page 78.) Salensky’s fine memoir was called forth by the necessity of determining the phyletic rela- tionships of the wild Asiatic horse EL. przewalskit. Materials. The materials are more extensive and important than those used by any pre- vious author, including sixty-four skulls, representing the following species: 9 E. przewalskit (4 young, 5 adult) 15 Pleistocene horses from Russia and Siberia 6 Domesticated asses, H. asinus 11 Kiang’s, Equus kiang 18 Onagers, Equus onager 1 Mountain zebra, Equus zebra Grevy’s zebra, Equus grevyi Burchell’s zebra, Equus burchelli Chapman’s zebra, E. chapmani mule 1 1 1 1 Methods. Of the thirty-five measurements of Salensky twenty-four are important as repre-_ sented in Fig. 7. Salensky’s measurement system is chiefly based on that of Nehring. Salensky, however, gives absolute measurements only, which renders it difficult to draw inferences from his large tables. The following criticisms of his measurements may be made. The length of face (30) to the inter-nasal suture is inadequate because the face should be measured back to the frontal lines (F F). The author follows Nehring in adopting his Indices I, II, III, in which the greater measure- ment is divided by the lesser instead of the lesser by the greater. The five measurements given 1 Salensky, W. ‘Equus przewalskii Poljak.’ Wissensch. Resultate, N. M. Przewalski nach Centrale-Asien Unternommenen Reisen auf Kosten einer von Seiner Kaiserlichen Hoheit dem Grossfiirsten Thronfolger Nikolai Alexandrowitsch. Published by Kaiserlichen Akad. d. Wissenshaften. Zool. Theil. Bd. 1, Mammalia, Abth. 2 Ungulata, Lieferung 1. 76 pp.4pll. St. Petersburg, 1902. ‘Prjevalsky’s Horse,’ Translation by Capt. M. Horace Hayes and O. Charnock Bradley, with an introduction by J. Cossar Ewart, 8vo., London, 1907. ''ADDENDUM. (To follow line 16, p. 66.) The use of the basilar length of skull as a divisor is an advance upon Nehring, because the long-faced forms were given the highest figures and the short-faced forms the lowest figures. Thus the mode of reckoning may be adopted but not the mode of measurement of the face, because the face should certainly include the orbit. Tscherski’s conclusions. Tscherski classifies the Equide into three great groups. Eastern, or Oriental. Western. 1. Broad-forehead type, 7. ¢., brachycephaly, typified 3. Narrow-forehead type, typified by the Forest Horse. by the Arab. 2. Medium-forehead type. The Post-Pliocene horses of Siberia belong to several races. They vary in size almost as much as the present races of domesticated horses in Europe. All have skulls of the median- forehead type, with a dentition, however, which resembles that of the long-headed occidental, or Norie Horses. Most of them have hoofs of medium or great breadth. The materials do not justify the assumption that among the Siberian fossil horses can be found forms which are iden- tical with the Post-Pliocene horses of Europe. Thus the facts do not contradict the view of Madame Pavlow that these Siberian horses of medium-forehead type are of Eastern stock, arising from such Tertiary Indian horses as H. namadicus and E. sivalensis. '' ''OSBORN, CRANIOMETRY OF THE EQUID”. 67 to determine the shape of the nasal bones do not bring out the essential differences between the horse and the ass very clearly. Salensky’s conclusions. There is no attempt to distinguish between the various original breeds of horses. Many important points of difference between the horses and the Asiatic and African asses are brought out incidentally. H. przewalski1 is found to belong to a special and sepa- rate type which, however, is more closely allied to the domestic horse than to the ass. Ewart’s System, 1907.! (Text Fig. 8, page 80.) Ewart’s system was called forth by the necessity of analyzing the skulls found in a Roman fort at Newstead. His epoch-making successive contributions in this and other papers to the study of recent and prehistoric horses have pointed the way to the clear distinction of the original stocks of horses. His system of measurements is clearly brought out in the important paper of 1907 above cited. Materials. The twelve Newstead skulls are measured, although the conclusions reached in the system of the author are based upon examination of many other types. Methods. The eight measurements given in the table bring out the chief differences in the eeneral proportions of the Newstead skulls under observation in a most telling manner. They serve this purpose, but a more extensive system of measurements is needed to bring out the essential points of difference, for example, between the horses, asses, and zebras. The frontal width measurement is that adopted by all previous authors and by Osborn. The cranium length (F’S) is also that adopted by Osborn. Ewart’s frontal index (7. e., frontal width ~ 4° does not give as true a conception of the relations of the width and length of the skull as the ‘cephalic index’ of Osborn and other authors (1. ¢., frontal width X 10°) basilar length Ewart has not put into his table any exact measurements of the facio-cranial angle, which constitutes one great point of difference between the three original stocks of horses which he dis- tinguishes. Conclusions. The Newstead skulls include “beside cross-bred animals” three very distinct types of horses which may be referred to three original stocks from which the modern domesti- cated horses sprang, namely, the Forest, the Steppe, and the Plateau. All three of these original stocks may well have existed in a pure and wild state at the end of the first century A. D. The descendants of these three stocks are now distributed as follows: Forest variety. Steppe variety. Plateau variety. (Broad-headed.) (Long-headed.) (Medium-to narrow-headed.) Northwest of Europe. Spain. Britain (native horses). Norway. Treland. Shetland and Faroe Isles. Scotch highlands and islands. Mexico. Southern Mexico. Iceland. South America. Jamaica. Asia. Austria. Java. Korea, ete. (= Ridgeway’s large-headed E. caballus.) The Arab belongs to the narrow-headed or Plateau Variety, the existing Arabs being mingled with Steppe and Forest blood. It is not necessary at this point to give Ewart’s more recent phyletic conclusions since the object of the present communication is chiefly craniometric. 1 ‘Qn Skulls of Horses from the Roman Fort at Newstead, near Melrose, with Observations on the Origin of Domestic Horses.’ Trans. Roy. Soc. Edinburgh, Vol. XLV, pt. 3, No. 20, 1907, pp. 555-587. ''68 OSBORN, CRANIOMETRY OF THE EQUIDA. Contribution of Bradley, 1907.! (Text Fig. 9, page 82.) Bradley’s contribution was based upon a comparison of the Przewalsky with the Forest type of Ewart as well as with the Celtic and Iceland ponies. In several respects it marks a departure from all previous systems of measurement, and this is regrettable because it prevents compari- son with all other tables although serving the author’s immediate purpose of bringing out several of the important differences between the types examined. The chief points brought out by Brad- ley’s observations and measurements may be summarized as follows: The actual width of the cranium of EH. przewalskii is less than an examination of this animal during life would lead us to suppose. It has a long, narrow face in contrast with the short, broad face of the Iceland pony and the intermediate face of the Celtic pony. Thus the face of E. przewalskii forms a considerable portion of the total length of the skull. The orbit of Z. prze- walskii has an elongate form and is placed relatively far back in the head. The premaxillary width in #. przwalskii is relatively less than in either the Celtic or the Iceland ponies. System of Cranial Measurement, Osborn, 1912. (Text Fig. 10, page 84; Text Fig. 11, p. 86.) The system here proposed is synthetic; (1) it retains so far as possible the measurement standards of Franck, Branco, Nehring, Tscherski, Salensky, Ewart, Bradley, in order that students may avail themselves of the absolute measurements given in the craniometric tables of these authors; (2) it contains nine measurements which are of es pecial value in distinguishing horses, asses, and zebras; (3) it contains measurements which will be of value in the comparison of fossil and recent animals and will be expressive even where proportions of the skull only are preserved. The absolute measurements from which the indices are calculated are the following, made in each instance by calipers or orthogonal projection: 1. Vertex length - 7. Cranial length 2. Basilar length 11. Occiput height 3. Frontal width 15. Diastema length 5. Facial length 16. Muzzle width 17. Dental length. The indices are obtained by the combination of the above actual measurements as follows: 4. Cephalic index, ratio of width to length. 6. Facio-cephalic index, ratio of face to total length. _ -8. Cranio-cephalic index, ratio of length of cranium to basilar length. 12. Orbital index, ratio of the length to the height of the orbit. 13. Vomerine index, expressing posterior extension of vomer. 18. Dental index, ratio of superior grinding series to total length of skull. 19. Molar index, ratio of breadth of a single grinder to length of entire molar-premolar series. In each instance the index is obtained by dividing the lesser diameter by the greater. 1‘Craniometrical Observations on the Skull of Equus prjevalskii and other Horses.’ Proc. Roy. Soc. Edinburgh, Vol. X XVII, Pt. I (No. 8), 1907, pp. 46-50. ''OSBORN, CRANIOMETRY OF THE EQUID. 69 The angles believed to be most significant are the following: o; 10. IEE ai. Palatal angle, elevation of posterior border of palate. Palato-cranial angle, approximately representing deflection of face on cranium. expressing the backward prolongation of the occiput above the condyles. Occiput-vertex angle, -cranial the deflection of the face on the cranium. Facio-cranial angle, expressing more accurately than the palato The manner of taking these measurements, indices, and angles is very clearly shown in the accompanying key (p. 85) and in Figs, 10, 11, (Text continued on page 88.) ''OSBORN, CRANIOMETRY OF THE EQUIDA. 70 FRANCK Fig. 3. Franck’s system, 1875. ''ee 10. ET. 12. 13. 14. 16. oi 18. 19, 20. al. 23. 24, 20. OSBORN, CRANIOMETRY OF THE EQUID. ie! Kry to FRANCK’s SYSTEM OF 1875. Length from foramen magnum to point between I’, |= BASILAR LENGTH, O.] (as measure also by Branco, Nehring, Tscherski, Salensky, Osborn). Fig. 3, III, B-I. From foramen magnum to posterior end of palatal suture (as measured also by Tscherski). Fig. 3, III, B-P. From foramen magnum to beginning of vomer (as measured also by Branco, Nehring, Tscherski, Salensky, Osborn). fie 3, Hl BEY. From posterior end of palatal suture to point between I’, [= PALATAL LENGTH, O.| (as measured also by Tscherski, Bradley, and Osborn,— fossil Equidee). Fig. 3, III, P-L. From posterior end of palatal suture to beginning of vomer (as measured also by Branco, Nehring, Tscherski, Salensky, Osnom). Fie. 3, 1; PY. Width between mawillary crests at their origin [= FACIAL-MAXILLARY WIDTH, O.] (as measured also by Nehring, Tscherski, approximately as measured by Salensky and Osborn,— fossil Equide). Fig. 3, II, 6. Width between canines (as measured by Franck only). Fig. 3, II, 7. Greatest width of skull (as measured also by Tscherski, Bradley, approximately as measured by Osborn,— fossil Equide). Fig. 3, III, 8 (between outer ends of joint surfaces for lower jaw). Greatest width between orbital processes [= FRONTAL wipTH, O.]| (as measured also by Branco, Nehring, Tscherski, Salensky, Ewart, Bradley, Osborn). Fig. 3, II, F-F. Greatest distance between pterygoid processes of palatal bones (as measured by Franck only). Fig. 3, III, 10. Length of molar row of upper jaw, not including p' (as measured also by Nehring, Tscherski, Salensky, Osborn). Fig. 3, ‘Hi; D=D. Nehring, Tscherski, Salensky: measure both along wearing surfaces and along alveoli. Greatest distance between M?, M? (as measured by Franck only). Fig. 3, III, 12. Distance between P?, P?, anterior end [= PatataL wipts (2), O.] (as measured also by Tscherski, Osborn,— fossil Equide). Fig. 3, II, 13. Straight line from middle‘of transverse process of occipital bone to point between I, I [= VERTEX LENGTH O.] (as measured also by Nehring, Tscherski, Salensky, Ewart, Osborn). Fig. 3, I, S—I. From middle of transverse process of occipital bone to tip of nasal bones (with tape measure) (as measured by Franck only). Fig. 3, I, 15. From middle of transverse process of occipital bone to tip of nasal bones (straight line) (as measured by Franck only). Hye. 3, 1, 16. From middle of occipital crest to point between orbits (?). Ewart, Osborn, Bradley: cranial length is taken from middle of line connecting posterior borders of orbits to occip- ital crest. From middle of occipital crest to end of nasal process of frontal bones (as measured also by Tscherski, Salensky). Fig. 3, i, 18. Median length of nasal bones (as measured also by schersks, Salensky). Fig. 3, II, 19. Straight line between tip of nasal bones and I’, (as measured by Franck only). Fig. 3, I, 20. Greatest breadth of cerebral portion of skull, sous zygomatic processes of ene bones (as measured also by Tscherski, Salensky, Ewart). Fig. 3, II, 21. Smallest breadth of skull capsule on small pterygoid foramina (as measured by Franck only). Fig. 3, III, 22. Branco: measures width of cranium at bases of styloid processes of petrosum. Tscherski: measures greatest width of head behind orbits at mastoid processes. Greatest width on parietal bump (as measured also by Tscherski, Salensky, Ewart). Fig. 3, I, 23. Width between supraorbital foramina (as measured by Franck only). Fig. 3, II, 24. Tscherski: measures width of forehead between points taken in middle of length of upper orbital edge. Width between infraorbital foramina (as measured by Franck only). Fig. 3, II, 25. ''72 OSBORN, CRANIOMETRY OF THE EQUIDZ. Fig. 4. Branco’s system (principal measurements), 1883. BRANCO ''10. tle 12. 13. 14. 15. 16. 17, 18. 19. 20. OSBORN, CRANIOMETRY OF THE EQUID. 73 Kry to Branco’s SysteM oF 1883. Lower edge of foramen magnum to alveolar border of premaxillaries, between I', I. [= BAstLaR LENGTH, O.] (as meas- ured also by Franck, Nehring, Tscherski, Salensky, Osborn). Fig. 4, II, B-I. Lower edge of foramen magnum to middle of posterior end of vomer (as measured also by Franck, Nehring, Tscherski, Salensky, Osborn). Fig. 4, III, B-V. Middle of posterior end of vomer to middle of posterior border of palate tas measured also by Franck, Nehring, Tscherski, Salensky, Osborn). Fig. 4, III, V—P. Diastema from anterior end of P? to posterior end of I? |= D1asTEMA LENGTH, O.] (as measured also by Nehring, Tscher- ski, Salensky, Osborn). Fig. 4, I, D-I?. Length of maxillary behind M? (as measured by Branco only). Fig. 4, I, 5. Horizontal line from posterior point of occipital crest to point in a vertical line with most anterior part of anterior orbital edge (as measured by Branco only). Fig. 4, I, 6. Vertical line from alveolar border of maxillaries, immediately behind M?, to plane of upper profile line of skull [= Depth OF SKULL (2) (as measured also by Tscherski, Salensky, Osborn,— fossil Equide). Fig. 4, I, 7. Vertical line from highest point of upper orbital edge to plane of upper profile line of skull (as measured by Branco only). dig. 4; I, 8. Vertical line from plane of internasal suture to upper angle of suture of lachrymal and maxillary (as measured by Branco only). Fig. 4, I, 9. Vertical line from plane of internasal suture to upper point of suture of premaaillary and maxillary (as measured by Branco only). Vertical distance from lower edge of zygomatic arch to upper (as measured by Branco only), at median point of lower orbital edge. Fig. 4, I, 11. Lower edge of foramen magnum to highest point of occipital crest (as measured also by Nehring, Tscherski, Salensky). Hie, 4, 11; B-S: Vertical diameter of orbit; horizontal diameter of orbit (as measured also by. Tscherski, Salensky, Ewart, Bradley, Osborn). Fig. 4, II, Ov Oh. idth of posterior border of orbit (as measured also by Tscherski at its narrowest point). Fig. 4, I, 14. Width of cranium (as measured by Branco only, at bases of styloid processes of petrosum). Fig. 4, III, 15. Franck: measures smallest breadth of skull capsule on pterygoid foramina. Tscherski: measures greatest width of head behind orbits at mastoid processes. Width of premazillaries at outer posterior borders of I, I’ [= Muzzue wits, O.] (as measured also by Franck, Nehring, Tscherski, Salensky, Bradley, Osborn). Fig. 4, II, I?-I?. Distance between most prominent points of orbital processes of frontals [= FRonTAL wiprs, O.] (as measured also by Franck, Nehring, Tscherski, Salensky, Ewart, Bradley, Osborn). Fig. 4, II, F-F. Height of horizontal ramus of lower jaw at posterior border of Ms |= Drptu oF JAw, O.] (as measured also by Osborn,— fossil Equide). Fig. 4, 5. : Height of horizontal ramus of lower jaw between Ps and M, [= Depru oF sAw, O.] (as measured also by Tscherski and Osborn,— fossil Equide). Fig. 4, 6. Height of horizontal ramus of lower jaw‘between P2 and P3 (as measured by Branco only). Fig. 4, 7. ''a4 OSBORN, CRANIOMETRY OF THE EQUIDZ. ee ee eras een ont alee ios ee eee ale oe ee ee Il NEHRING Fig. 5. Nehring’s system (principal measurements), 1884. ''10. id. 12. 13. 14. 15. 16. OSBORN, CRANIOMETRY OF THE EQUID. 75 Key to Neurine’s PrincrpAL MEASUREMENTS, 1884. Length of vertex (as measured also by Franck, Tscherski, Salensky, Ewart, Osborn) from edge of occipital crest along external upper surface of skull to base of middle incisors. Fig. 5, I, S-I. ~ Basilar length (as measured also by Franck, Branco, Tscherski, Salensky, Osborn) along base of skull from lower border of foramen magnum to base of middle incisor teeth. Fig. 5, HI, B-I. Frontal width (as measured also by Franck, Branco, Tscherski, Salensky, Ewart, Bradley, Op between posterior margins of orbital cavities. Fig. 5, II, F-F. Index I [= Fronvau INDEX, O.] (as taken also by Branco, Tscherski, Salensky); basilar length (2) X 100 + frontal width (3). Ewart: Frontal index is frontal width x 100 + facial length. Index IT (as taken also by Tscherski, Salensky); length of vertex (1) X 100 + frontal width (3). Posterior segment of ocular line (as measured also by Tscherski, Salensky) from middle of occipital crest to most external point on posterior border of orbital cavity. Fig. 5, I, 6. Anterior segment of ocular line (as measured also by Tscherski, Salensky) from most external point on posterior border of orbital cavity to base of middle incisors. Fig. 5, I, 7. Index IIT — Ocular Index (as taken also by Tscherski, Salensky); anterior segment of ocular line (7) X 100 + poste- rior segment of ocular line (6). Facial width [= FACIAL-MAXILLARY WIDTH, O.] (as measured also by Franck, Tscherski; approximately as measured by Salensky, Bradley, Osborn,— fossil Equidse) between zygomatic ridges of superior maxillary bones. Fig. 5, II, 9. Width of muzzle (as measured also by Franck, Branco, Tscherski, Salensky, Osborn) at posterior alveolar borders of tPF. Wig. 5,11, 2-1. Franck’s vomer index (as measured also by Franck, Branco, Tscherski, Salensky, Osborn); distance from posterior border of palate to middle of posterior edge of vomer (PV) X 100 + distance from same point on vomer to anterior border of foramen magnum (VB). Fig. 5, III. Height of skull at occipital crest [= Occirut netcnt, O.] (as measured also by Branco, Tscherski, Salensky, Osborn) ; length of perpendicular from middle of occipital crest to base of mandible. Fig. 5, I, S~A. Length of diastema, I°-P? (as measured also by Branco, Tscherski, Salensky, Osborn). Fig. 5,1, Length of series of wpper cheek-teeth, along wearing surfaces (as measured also by Franck, Branco, Tscherski, Salensky, Osborn). Fig. 5, LH, D-D. Along alveoli (as measured also by Tscherski, Salensky). Fig. 5, I, D'-D'. Length of mandible (as measured also by Tscherski, Salensky, Osborn,— fossil Equide) from condyle to base of central incisors. Fig. 11, 1. Height of mandible (as measured also by Tscherski, Beige Osborn,— fossil Equidee) from condyle to base. Fig. MM 2: ''76 OSBORN, CRANIOMETRY OF THE EQUID. Fig. 6. Tscherski’s system (principal measurements), 1892. TSCHERSKI ''bo 10. ik 12. 13. 14. 15. 16. iy, 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. OSBORN, CRANIOMETRY OF THE EQUID. i Key to Tscurerski’s PrincipaAL MEASUREMENTS, 1892. Length of base [= BaAsILAR LENGTH, Osborn] (as measured also by Franck, Branco, Nehring, Salensky, Osborn) from point between central incisors to lower edge of foramen magnum. Fig. 6, III, I-B. Cranial length (greatest length) [= VERTEX LENGTH, Osborn] (as measured also by Franck, Nehring, Salensky, Ewart, — Osborn) from point between central incisors to occipital crest). Fig. 6, 1, 1-3. Greatest breadth of forehead [= FRONTAL WIDTH, Osborn] (as measured also by Franck, Branco, Nehring, Salensky, Ewart, Bradley, Osborn) at posterior borders of orbits. Fig. 6, II, F-F. Frontal index (as measured also by Branco, Nehring, Salensky); length of base (1) X 100 + greatest breadth of forehead (3). Ewart, Bradley: Frontal index is frontal width X 100 + facial length. Nehring: Index II is length of vertex X 100 + frontal width. Transverse convexity of forehead — Index (as measured by Tscherski only), measuring between middle points of upper orbital edges, are X 100 + cord. Fig. 6, I, 5. Longitudinal convexity of forehead — Index (as measured by Tscherski only), measuring from line connecting middle points of upper orbital edges to occipital crest, are X 100 + cord. Fig. 6, II, 6. Greatest breadth of head behind orbits (as measured by Tscherski only) at mastoid processes. Pig. 6; LE, 7 Franck: measures smallest breadth of skull capsule on pterygoid foramina. Branco: measures width of cranium at bases of styloid processes of petrosum. Posterior section of eye-line (as measured also by Nehring, Salensky) from middle of occipital crest to most external point on posterior border of orbital cavity. Fig. 6, I, 8. Anterior section of cye-line (as measured also by Nehring, Salensky) from point between central incisors to most ex- ternal point on posterior border of orbital cavity. Fig. 6, I, 9. Eye index (as measured also by Nehring, Salensky) anterior section of eye-line (9) X 100 + posterior section of eye- line (8). Length of face (as measured by Tscherski only), from point between central incisors to nearest point of anterior orbital edge. Fig. 6, II, 11. Salensky: Length of face is measured from middle incisors to posterior end of internasal suture. Ewart, Bradley, Osborn: Facial length is measured from median incisive border to middle of line connecting poste- rior borders of orbits. Facial index (as measured by Tscherski only) length of face (11) X 100 + length of base Eb) Salensky: Index IV is length of vertex X 100 + length of face (see 11, note, above). Ewart, Osborn: Facio-cranial index is facial length (see 11, note, above) X 100 + cranial length. Convexity of nasals — Index (as measured by Tscherski only) measuring smallest width of nasal bones behind infra- orbital foramina, arc X 100 + cord. Fig. 6, II, 13. Breadth of snout at maxillary crests [= FactAL-MAXILLARY WIDTH, O.] (as measured also by Franck, Nehring; approx- imately as measured by Salensky, Bradley, Osborn,— fossil Equidee). Fig. 6, III, 14. Breadth of snout at points between P* and M. (as measured by Tscherski only). Fig. 6, III, 15. Breadth of snout at anterior ends of alveoli of P? [= Patavau wrpts (2), O.] (as measured also by Franck, Osborn,— fossil Equide). Fig. 6, III, 16. Breadth of snout at posterior ends of alveoli of I? [= Muzzie wiptu, Osborn] (as measured also by Franck, Branco, Nehring, Salensky, Osborn, Bradley). Fig. 6, III, 17. Length of diastema between I? and P? [= D1asTEMA LENGTH, Osborn] (as measured also by Branco, Nehring, Salensky, Osborn). Fig. 6, I, —-D. : Length of dentition, P?-M® along wearing surfaces (as measured also by Franck, Branco, Nehring, Salensky, Osborn) ; along alveoli (as measured also by Nehring, Salensky). Fig. 6, III, D-D. Perpendicular height of face at posterior alveolus of M? |= Drpru oF SKULL (2), Osborn] (as measured also by Branco, Salensky, Osborn,— fossil Equide). Fig. 6, I, 20. Perpendicular height of face at posterior end of anterior nasal opening (as measured by Tscherski only). Fig. 6, I, 21. Height of head |= Occrput nEicHt, Osborn] (as measured also by Branco, Nehring, Salensky, Osborn), perpendicular from occipital crest to base of mandible. Fig. 6, I, S-A. Length of lower jaw (as measured also by Nehring, Salensky, Osborn,— fossil Equidee) from median incisive point to posterior, outer edge of head of condyle. Fig. 11, 1. Height of ascending ramus (as measured also by Nehring, Salensky, Osborn,— Fossil Equide) from head of joint perpendicularly to plane of table. Fig. 11, 2. Width of incisive portion [= Wiprr or sympuysis (2), Osborn] (as measured also by Salensky, Osborn,— fossil Equi- de). Vig. 11, 3: Narrowest width [= Wipru or sympuysis (1), Osborn] (as measured also by Osborn,— fossil Equide). Fig. 11, 4. Height of lower jaw in region of space between P, and M, |= DEprtu oF Jaw, Osborn] (as measured also by Branco, Osborn,— fossil Equide). Fig. 11, 6. ''OSBORN, CRANIOMETRY OF THE EQUID. Fig. 7. Salensky’s system (principal measurements), 1902. SALENSKY '' }. 9 N 10. ae Lé. 18. 19. 20. 22. 23. 24. 20. 26. 27. OSBORN, CRANIOMETRY OF THE EQUIDA. 19 SALENSKY’sS PrincipAL MEASUREMENTS, 1902. Frontal width (as measured also by Franck, Branco, Nehring, Tscherski, Ewart, Bradley, Osborn), between posterior margins of orbital cavities. Fig. 7, H, F-F. Basilar length (as measured also by Franck, Branco, Nehring, Tscherski, Osborn) from base of middle incisors to lower border of foramen magnum. Fig. 7, III, I-B. Length of vertex (as measured also by Franck, Nehring, Tscherski, Ewart, Osborn) from base of middle incisors to edge of occipital crest. Fig. 7, I, I-S. Posterior ocular line (as measured also by Nehring, Tscherski) from occipital crest to most external point on posterior border of orbital cavity. Fig. 7, I, 4. Anterior ocular line (as measured also by Nehring, Tscherski) from base of middle incisors to most external point on posterior border of orbital cavity. Fig. 7, I, 5. Index I [= FrontTAu INDEX, Osborn] (as measured also by Branco, Nehring, Tscherski, Osborn) basilar length x 100 + frontal width. Ewart: Frontal index is frontal width X 100 + facial length. Index IT (as measured also by Nehring, Tscherski) length of vertex X 100 + frontal width. Index ITI (as measured also by Nehring, Tscherski) anterior ocular line (5) X 100 + posterior ocular line (4). Facial width [= FAcIAL-MAXILLARY WIDTH, Osborn] (as measured also by Osborn,— fossil Equide, approximately as measured by Nehring, Tscherski at maxillary crests). Fig. 7, I, 9. Length of series of upper cheek teeth along wearing surfaces (as measured also by Franck, Branco, Nehring, Tscherski, bom). Pag. 7, 11 D_D: ; along alveoli (as measured also by Nehring, Tscherski). Fig. 7, I, D'-D'. Distance from margin of foramen magnum to vomer (as measured also by Franck, Branco, Nehring, Tscherski, Osborn). His, 7, 1 Bey. Distance from vomer to palatine bone (as measured also by Franck, Branco, Nehring, Tscherski, Osborn). Fig. 7° Ti VP: Nasal width of upper jaw [= Mvuzz_e wipru, Osborn] (as measured also by Franck, Branco, Nehring, Tscherski, Bradley, Osborn) at posterior alveoli of I’. Fig. 7, I, P-IP. Length of diastema, I°—P? (as measured also by Branco, Nehring, Tscherski, Osborn). Fig. 7, I, ’-D'. Height of skull [= Occreut HEtGHT, O.]| (as measured also by Branco, Nehring, Tscherski, Osborn) perpendicular from occipital crest to base of mandible. Fig. 7, I, S-A. Length of cranium (as measured by Salensky only) from posterior end of internasal suture to occipital crest. Fig. 7, Hi, N-S: Ewart, Osborn: Cranial length is measured from middle of line connecting posterior borders of orbits, to occipital crest. Length of face (as measured by Salensky only) from middle incisors to posterior end of internasal suture. Fig. 7, Il, I-N. Tscherski: Length of face is measured from point between central incisors to nearest point on anterior orbital edge. Ewart, Bradley, Osborn: Facial length is measured from median incisive border to middle of line connecting poste- rior borders of orbits. Antero-posterior diameter of orbital cavity (as measured also by Branco, Tscherski, Ewart, Osborn, approximately as measured by Bradley). Fig. 7, I, Oh. Vertical diameter of orbital cavity (as measured also by Branco, Tscherski, Ewart, Osborn, approximately as measured by Bradley). Fig. 7, I, Ov. Width of cranium (as measured by Salensky only) between superior margins of external auditory meati. Fig. 7, III, 20. Greatest width of cranium (as measured by Franck, Tscherski, Ewart) above zygomatic processes of temporal bones. Fig, 7, bl; 21. Least width of cranium (as measured also by Tscherski) behind orbital cavities. Fig. 7, I, 22. Height of occipital bone (as measured also by Branco, Nehring, Tscherski) from lower edge of foramen magnum to occipital crest. Fig. 7, I, SB. Height of face |= Deprun o¥ sku. (2), Osborn] (as measured also by Branco, Tscherski, Salensky, Osborn) from base of M? to middle of internasal suture. Fig. 7, I, 24. Index IV (as measured by Salensky only) length of vertex (3) X 100 + length of face (17). Tscherski: Facial index is length of face (see 17, note, above) X 100 divided by basilar length. Ewart, Osborn: Facio-cranial index is facial length (see 17 note above) X 100 + cranial length, (see 16, note, above). Width of nasal bones in their posterior part (as measured by Salensky only) at posterior and most external point on naso-lachrymal suture. Fig. 7, II, 26. Width of nasal bones in middle of their length (as measured by Salensky only) at anterior and most external point on naso-lachrymal suture. Fig. 7, II, 27. ''80 28. 29. 30. ol: OSBORN, CRANIOMETRY OF THE EQUIDZ. Width of nasal bones in their anterior part (as measured by Salensky only) at point of contact of intermaxillary and nasal bones. Fig. 7, II, 28. Length of nasal bones (oblique) (as measured by Salensky only) at highest point of naso-frontal suture. Fig: 7, 35. 29: Length of nasal bones (median) (as measured also by Franck, Tscherski. Fig. 7, I, 30. Length of lower jaw (as measured also by Nehring, Tscherski, Osborn,— fossil Equidee) from condyle to base of central incisors. Fig. 11, 7. Height of lower jaw (as measured also by Nehring, Tscherski, Osborn,— fossil Equide). Fig. 11, 2. Width of lower jaw |= WiprH or sympuysis (2), Osborn] (as measured also by Tscherski, Osborn,— fossil Equide). Piz, 11; 8, Length of series of lower cheek tecth along wearing surfaces (as measured also by Tscherski, Osborn,— fossil Equide) ; along bases (as measured also by Tscherski). Fig. 11, 8. Length of diastema, I3~P: (as measured also by Tscherski, Osborn,— fossil Equide). Fig. 11, 9. Fig. 8. Ewart’s system, 1907. ''10. OSBORN, CRANIOMETRY OF THE EQUIDZ2. 81 Ewart’s PrincipaL MEASUREMENTS, 1907. Total length [= VERTEX LENGTH, Osborn] (as measured also by Franck, Nehring, Tscherski, Salensky, Osborn) from occipital crest to alveolar point, i. e. to wedge-like piece which projects between upper central incisors. Fig. 8, TS. : Facial length (as measured also by Bradley, Osborn) from central incisors to a line connecting posterior borders of orbits. Fig. 8, I, S-F’. ; Tscherski: Facial length is taken from point between central incisors to nearest point on anterior orbital edge. Salensky: Facial length is taken from central incisors to posterior end of internasal suture. Frontal width (as measured also by: Franck, Branco, Nehring, Tscherski, Salensky, Bradley, Osborn) between outer margins of orbits. Fig. 8, I, F-F. Cranium length (as measured also by Osborn) from occipital crest to line connecting posterior borders of orbits. Fig. 61 5. e Salensky: Cranial length is taken from posterior end of internasal suture to occipital crest. Cranium width (as measured also by Franck, Tscherski, Salensky) at widest part. Fig. 8, I, 5. Frontal index (as measured also by Bradley) frontal width (3) X 100 + facial length (2). Branco, Nehring, Tscherski, Salensky, Osborn: Frontal index is basilar length X 100 + frontal width. Cephalic index (as measured by Ewart only) cranium width (5) X 100 + cranium length (4). Orbital index (as measured also by Branco, Tscherski, Salensky, Osborn, approximately as measured by Bradley) vertical diameter of orbit X 100 + horizontal diameter. Fig. 8, I, O-0 . Angle of deflection of face (line) [= PALATO-CRANIAL ANGLE (LINE), Osborn] (as measured also by Osborn) distance from middle of posterior border of palate to a line connecting median incisive border and foramen magnum. Angle of deflection of face [= approximately PALATO-CRANIAL ANGLE, Osborn] (approximately as measured by Osborn) between line of palate and basicranial axis. ''82 OSBORN, CRANIOMETRY OF THE EQUIDZ. Fig. 9. Bradley’s system, 1907. BRADLEY ''10. a. Te 13. 14. 15. 16. ET, 18. £9, 20. 21. 22. 23. OSBORN, CRANIOMETRY OF THE EQUIDA. 83 Brapiey’s PrincreaL MEASUREMENTS, 1907. Cranium length (as measured by Bradley only) from the opisthion to the interorbital point, 7. e., to the middle point of a transverse line on a level with the anterior border of the post-orbital process of the frontal bone. Fig. 9, I, X-F’. Condylar breadth of cranium (as measured also by Franck, Tscherski, approximately as measured by Osborn,— fossil Equide) transverse diameter from the outermost part of the condyles on the squamous temporal bones. Fig. 9, 2. Maximum parietal breadth of cranium (as measured by Bradley only) between points on the parieto-squamosal suture. Pic. 9, 11, 3. Cephalic index A (as measured by Bradley only): condylar breadth (2) X 100 + cranium length (1). Cephalic index B (as measured by Bradley only): maximum parietal breadth (8) & 100 + cranium length (1). Facial length (as measured also by Ewart, Osborn) from the alveolar point to the interorbital point, 7. e., to the middle point of a transverse line on a level with the anterior border of the postorbital process of the frontal bone. Fig. 9, I-II, I-F’. Facial breadth (approximately as measured by Salensky, Osborn,— fossil Equidee) between the most distant points on the sutures separating the malar and maxillary bones. Fig. 9, II, 7. Facial index (as measured by Bradley only) facial breadth (7) X 100 + facial length (6). Frontal breadth (as measured also by Franck, Branco, Nehring, Tscherski, Salensky, Ewart, Osborn) maximum breadth of skull at posterior borders of orbits. Fig. 9, I, F-F. Frontal index (as measured also by Ewart) frontal breadth (9) X 100 + facial length (6). Cranio-facial length (as measured by Bradley only) from the opisthion to the alveolar point. Fig. 9, II, I-X. Cranio-facial index (as measured by Bradley only) length of face (6) X 100 + cranio-facial length (11). Palate length (as measured also by Franck, Tscherski, Osborn,— fossil Equide. Fig. 9, III, I-P. Palate width (as measured also by Osborn,— fossil Equide). Fig. 9, III, 14. Palatine index (as measured by Bradley only) palate width (14) X 100 + palate length (13). Maximum diameter of orbit (approximately as measured by Branco, Tscherski, Salensky, Ewart, Osborn). Fig. 9, I, Oh. Minimum diameter of orbit (approximately as measured by Branco, Tscherski, Salensky, Ewart, Osborn). Fig. 9, I, Ov. Orbit index (approximately as measured by Branco, Tscherski, Salensky, Ewart, Osborn) minimum diameter of orbit (17) X 100 + maximum diameter (16). Distance from opisthion to supraorbital foramen (as measured by Bradley only). Fig. 9, I, 19. Distance from supraorbital foramen to alveolar point (as measured by Bradley only). Fig. 9, I, 20. Index (as measured by Bradley only) distance from opisthion to supraorbital foramen (19) X 100 + distance from supraorbital foramen to alveolar point (20). Maximum width of premaxilla [= Muvzz.e wipts, Osborn] (as measured also by Franck, Branco, Nehring, Tscherski, Salensky, Ewart, Osborn) measured from the bases of the third incisor teeth. Fig. 9, I, P’-P. Index (as measured by Bradley only) facial length (6) X 100 + width of premaxille (21). ''84 OSBORN, CRANIOMETRY OF THE EQUID. Fig. 10. Osborn’s system, skull, 1912. OSBORN ''10. 1s OSBORN, CRANIOMETRY OF THE EQUID. 85 Key To CrantAL MEASUREMENTS, OsBorRN, 1912. Vertex length (from median incisive border to middle of occipital crest). Fig. 10, I, I-S. As measured also by Franck, Nehring, Tscherski, Salensky, Ewart. Basilar length (from median incisive border to anterior edge of foramen magnum). Fig. 10, III, I-B. As measured also by Franck, Branco, Nehring, Tscherski, Salensky. Frontal width (at posterior borders of orbits). Fig. 10, II, F-F. As measured also by Franck, Branco, Nehring, Tscherski, Salensky, Ewart, Bradley. * Cephalic index (frontal width X 100 + basilar length). Ewart, Bradley: Frontal index is frontal width X 100 + facial length. Nehring, Tscherski, Salensky: index II is length of vertex X 100 + frontal width. —__— — As measured by Osborn only. Facial length from median incisive border to middle of line connecting posterior borders of orbits). Fig. 10, I, I-F’. Tscherski: facial length is measured from point between central incisors to nearest point on anterior orbital edge. Salensky: facial length is measured from incisors to posterior end of internasal suture. As measured also by Ewart and Bradley. Facio-cephalic Index (facial length X 100 + basilar length). Tscherski: facial index is length of face (see 5, note, above) X 100, divided by basilar length. Salensky: index IV is length of vertex X 100 + length of face (see 5, note, above). As measured by Osborn only. Cranial length (from middle of line connecting posterior borders of orbits to middle of occipital crest). Salensky: cranial length is measured from posterior end of internasal suture to occipital crest. As measured also by Ewart. Cranio-cephalic Index (cranial length X 100 + basilar length). Palatal angle, line (vertical distance from middle of posterior border of palate to a line [dots] connecting median incisive border and foramen magnum). Fig. 16, P—P?. As measured also by Ewart. Palato-cranial angle (angle between basicranial line, taken outside, and basifacial line, 7. e., line passing through median incisive border and middle of posterior border of palate). Fig. 16, B-B', I-P!. Lankester: a vertical longitudinal section of skull is made. Angle is taken between basicranial line, which follows cut surface of bones, and “palatine horizontal,” which connects anterior border of premaxillaries and foramen magnum (see Fig. 16, lower, I-B). Ewart: angle is measured between line of palate and basicranial axis. As measured by Osborn only; approximately as measured by Lankester, Ewart, and others. * Occiput-vertex angle (angle between vertex and line connecting most posterior points of occipital crest and condyles). Mig. 10,3, 1S C. As measured by Osborn only. (* Denotes importance in comparing horses with asses and zebras.) ''86 12. 13. 14. 15. 16. ly; 18. Us 20. yale OSBORN, CRANIOMETRY OF THE EQUID2. * Occiput height (length of perpendicular from middle of occipital crest to base of mandible). Fig. 10, I, S-A. As measured by Branco, Nehring, Tscherski, Salensky. * Orbital index (vertical diameter of orbit X 100 + horizontal diameter). Fig. 10, I, Ov, Oh. i. As measured also by Branco, Tscherski, Salensky, Ewart, approximately as measured by Bradley. * Franck’s vomer index (distance from posterior border of palate to middle of posterior edge of vomer X 100 + distance from same point on vomer to anterior edge of foramen magnum). Fig. 10, III, P-V, V-B. As measured also by Franck, Branco, Nehring, Tscherski, Salensky. * Convexity of nasofrontal suture (distance from posterior end of internasal suture to middle of line connecting two most posterior points of frontonasal suture). Fig. 10, II, N-N. As measured by Osborn only. * Diastema length P—P?. Fig. 10, I, -D. As measured by Branco, Nehring, Tscherski, Salensky. * Muzzle width (at posterior alveolar borders of I’). Fig. 10, I, [?-I’. As measured also by Franck, Branco, Nehring, Tscherski, Salensky, Bradley. Molar-premolar series total length (upper) p*-m? (length of wearing surface). Fig. 10, III, D—D. Nehring, Tscherski, Salensky measure both wearing surface and length along alveoli. As measured also by Franck, Branco, Nehring, Tscherski, Salensky. * Dental Index (Molar-premolar series total length (upper) X 100 + basilar length). * Molar Index (transverse diameter of m? X 100 + molar-premolar series total length). Fig. 10, III, M-M, D-D. As taken by Osborn only. Facio-cranial angle (from optic foramen to middle of incisive border as compared with basi-cranial line). ''OSBORN, CRANIOMETRY OF THE EQUID. 87 Fig. 11. Osborn’s system, lower jaw, 1912. System of Measurements of Lower Jaw, Osborn, 1912. Branco. : (1) Height of horizontal ramus at posterior border of Ms [= Depru or saw, Osborn] (as measured also by Osborn,— fossil Equide). Fig. 11, 5. (2) Height of horizontal ramus between Ps and M, [= Depvu oF saw, O.] (as measured also by Tscherski and Osborn,— fossil Equide). Fig. -11, 6. (3) Height of horizontal ramus between P2 and Ps (as measured by Branco only). Fig. 11, 7. Nehring. (4) Length of mandible (as measured also by Tscherski, Salensky, Osborn,— fossil Equidz) from condyle to base of central incisors. Fig. 11, 1. (5) Height of mandible (as measured also by Tscherski, Salensky, Osborn,— fossil Equidee) from condyle to base. Fig. 11, 2. Tscherski. : (6) Length of mandible (as measured also by Nehring, Selensky, Osborn,— fossil Equidee) from median incisive point to posterior outer edge of head of condyle. Fig. 11, 1. (7) Height of ascending ramus (as measured also by Nehring, Salensky, Osborn,— fossil Equids) from head of joint perpendicularly to plane of table. Fig. 11, 2. (8) Width of incisive portion [= WipTH oF sympHysis, (2) Osborn] (as measured also by Salensky, Osborn,— fossil Equide). Fig. 11, 3. (9) Narrowest width [= WiptH oF sympuysis (1), Osborn] (as measured also by Osborn,— fossil Equide). (10) Height in region of space between Ps and M,{= Depru or saw (1), Osborn] (as measured also by Branco, Osborn,— fossil Equide). Fig. 11, 6. Salensky. (11) Length of lower jaw (as measured also by Nehring, Tscherski, Osborn,— fossil Equide) from condyle to base of central incisors. Fig. 11, 1. (12) Height of lower jaw (as measured also by Nehring, Tscherski, Osborn,— fossil Equide) from condyle to base. Bie 1): 2. (13) Width of lower jaw |= Wiorx oF sympnysis (2), Osborn] (as measured also by Tscherski, Osborn,— fossil Equidee) at posterior alveoli of Is. Fig. 11, 3. (14) Length of series of lower cheek teeth along wearing surfaces (as measured also by Tscherski, Osborn,— fossil Equi- dee); along bases (as measured also by Tscherski). Fig. 11, 8. (15) Length of diastema, I3-P2 (as measured also by Tscherski, Osborn,— fossil Equide). Fig. 11, 9. ''88 OSBORN, CRANIOMETRY OF THE EQUID. II. DISTINCTIONS BETWEEN HORSES, ASSES, AND ZEBRAS. Attention may be called to some of the distinctions between the skulls of horses, asses, and zebras as observed by the following authors: Branco (1883), Dugés (1898), Salensky (1907); with annotations by 8. H. Chubb and J. K. Mosenthal, whose observations are enclosed in square brackets [ |: Branco, 1883: Orbital dimensions. Horse, longitudinal diameter greater than vertical. Ass, longitudinal and vertical diameters nearly alike; vertical may be greater than horizontal. [Very prominent, brow projecting laterally.] Distance from m to a vertical line from the anterior border of orbit. Horse, not so great as in Ass. [Increases greatly with age.| Distance from 7 to 7°. Horse, greater relatively than in Ass. Proportion of transverse and anteroposterior dimensions of Molars and Premolars. Horse, transverse relatively less than in Ass. Shape of mandible. There are two types of mandibles: (1) the horizontal ramus has a straight lower border, (2) the horizontal ramus has an indentation (Gefiissausschnitt) in a line with m3, which makes the ramus less high at this point. Horse, both types occur. Ass, generally has the latter type. Fig. 12. Lower jaws of domestic ass (above) and horse (below), showing characteristic differences in lower border of horizontal rami (ass, Amer. Mus. spec. 15675; horse, Amer. Mus. spec. 16613). '' OSBORN, CRANIOMETRY OF THE EQUIDZ. 89 Dugés, 1898: : Ass. 1. Well marked longitudinal convexity of forehead [valid]. 2. Face short and high [valid]. 3. Orbit subtriangular (its posterior border is straight) [see notes]. 4. Vertical line, from occipital crest to plane of base of mandible, passes well behind condyles [see notes]. Occipital crest is prominent. 6. Line of Lesbre (through anterior end of maxillary crest and point just above external auditory opening) passes through occipital crest. Ou 7. Upper molars lack the internal fold, ‘pli caballin’ of Lesbre. Horse. 1. Forehead straight. 2. Face long and relatively narrow. 3. Orbit subcircular [see notes]. 4. Vertical line (as above, No. 4) passes between condyles or touches them. Occipital crest less prominent and continues curve of occiput. S> Or Line of Lesbre (as above, No. 6) passes behind (below) occipital crest. ‘Pli caballin’ very distinct. [Absence of ‘pli caballin’ extremely rare except in very old subjects. | Note: drawings from Lesbre, 1892, Bull. Soe. Anthropol., Lyon, No. I. Cornevin, Lesbre, Piétrement, Gaudry, cited in Supplementary Note to Dugés article. au Salensky, 1907: FraNnck’s INDEX. (1) distance from posterior edge of vomer to posterior edge of palate; (2) distance from posterior edge of vomer to foramen magnum. (1) is greater in ass. (2) is greater in horse. [Not characteristic for Asiatic asses. Varies with species and within species of zebras. | [Franck gives 9 characteristic differences between ass and horse skulls. Salensky and Tscherski consider them insufficient, with the exception of the above.] Shape of nasals. Horse, nasals narrow gradually anteriorly (wedge-shape). Ass, nasals narrow rapidly anteriorly (club-shape). Horse, naso-frontal sutures are two fairly high arches. Ass, naso-frontal sutures are two almost straight lines. Horse, posterior ends narrow anteriorly. Ass, posterior ends increase in width anteriorly. [Not characteristic for Asiatic asses.] Naso-lachrymal suture. Horse, a straight line, nearly parallel with long axis of skull. Ass, lachrymal border concave. [Not characteristic for Asiatic asses.] Position of infraorbital foramen. Horse, nearer naso-maxillary suture than in Ass. [By no means constant.] Post-orbital process of frontal. Horse, three-sided. Ass, oval and compressed. [In Kiang narrow.] Height of skull at occiput, mandible included. Horse, lower than in Ass. Length of diastema. Horse, longer than in Ass. [Even shorter in Kiang.| [Variable in Asiatic Asses. ] Shape of mandible. Horse, inferior border smooth and straight. ''90 OSBORN, CRANIOMETRY OF THE EQUID. Ass, inferior border curved and furnished with prominences; inferior border thicker than in horse; between horizontal and vertical rami a characteristic indentation. [Very important.] S. H. Chubb, 1911. Mule: Much closer to horse 2 than to ass co’. Occipital crest horse-like, only slightly modified. Nasal-very close to horse though convexity in posterior third very slight. Well arched in central transverse section, though not quite so high as in horse. Diastema very little shorter than in horse. Lachrymal hardly differs from horse. Kiang in very many points approaches horse rather than ass. Horse: Strong convexity in posterior third of nasals (profile aspect), not present, however, in Arabs. Highly arched in central transverse section, coming low to meet maxillaries. Ass: Nasals, naso-premaxillary suture very short and high; premaxilla inserted well into nasal. Kiang shows the opposite extreme. Suture is very low. Nasal process coming down to meet premaxilla, the horse being somewhat intermediate in this point. a B IE ae . =s ~ SS B 3 > C J oe. Zs ep E 7 oo = A Ar a oo ' ce 3 \, ' / mets aves 1s Se \ ihe es = e Uc ™ Ne ~~ e fae —— y mae SO . I, 17 ee ee 7 ) | SSSA ype d 3 mpd ' ' 1 BH SS is — tears EL \ B 1 oe ce ae *e i zebra (fine line), and domestic ass (dotted line), showing differences in Fig. 13. Outlines of typical skulls of horse (heavy line), The occiput. The skulls are drawn so that the lines connecting the condyles and the median incisive border in each ease coincide. individuals represented are of approximately the same age (Horse, Amer. Mus. spec. 16613, Zebra, Amer. Mus. spec. 35153, Ass, Amer. Mus. spec. 15675). OcciputT VERTEX ANGLE. The outlines of the crania of the horse (A), the zebra (B), the ass (C) in Fig. 13 expresses the fairly constant differences in the proportions and profiles of these three types. It is espe- cially designed to express the occiput-vertex angle, in which the horse is seen to be more per- pendicular, the zebra intermediate, and the ass more retrocumbent. This is set forth in the following measurements in a series of horses, asses, and zebras, in which it appears that the occiput vertex angle in the horse varies from 76.5° to 64°, while the ass varies from 60° to 52.5°. The averages are as follows: Horse, average, not including E. przewalskit = 70.33° Zebra, average 61.35° Ass, average = 55D: ANGLE oF OccrpuT AND VERTEX (OCCIPUT-VERTEX ANGLE). Horses: Pony (Texas, A. M. No. 166613) Stage 8-5 years 16.5. Horse (Indian Territory, A. M. No. 19173) « 5. * To Horse (A. M. No. 16275) {OID (4.5 '' OSBORN, CRANIOMETRY OF THE EQUID. 91 Horse (New Mexico, A. M. No. 15753) Stage 9-1 years Toe Arab horse (Chubb No. 70) . 8-8 “ aa Horse (New York, A. M. No. 14131) [ 8-10 “ 69° Horse (Wyoming, A. M. No. 16279) ea 600: Horse (New York, A. M. No. 14132) s 9-15 “ 65.5 Horse (A. M. No. 15093) & Slo 64.° E. przewalskii (hybrid, Chubb No. 71) 2 8-5. 66.° Horse, average, not including E. przewalskti = 70.33° Zebras: Burchell’s Zebra (A. M. No. 35167) Stage 8-5 years 65.5" Burchell’s Zebra (A. M. No. 35153) 9-15 “ 63° Burchell’s Zebra (A. M. No. 14096) « ol 62.5" Burchell’s Zebra (Chubb No. 47) eo Gr Burchell’s Zebra (Chubb No. 45) i 64 “ 61° Burchell’s Zebra (Chubb No. 46) : - fon 58.5" Grevy’s Zebra (Chubb No. 65) © Oe 58° Zebra average = 61.35° Asses: Mexican Burro (A. M. No. 13982) Stage 5-2 years 60° E. asinus from Soudan (A. M. No. 295) : 4-14 “ Be E. asinus from Jamaica (A. M. No. 15675) eS 8-5. Sao E. asinus from Soudan? (A. M. No. 139) ‘* »11-over 30 years 52.57 Ass average = 55.75° Line of occiput ii ; ‘ ASS ZEBRA HORSE Averages Fig. 14. Diagrams showing differences in occiput-vertex angle (see Osborn’s system, No. 10) of skulls of horses, zebras and asses. Measurements were made on the skulls of ten horses of various breeds and ages, seven zebras (chiefly HZ. burchelli), and four domestic asses. The upper diagram indicates the largest and the smallest angle in each series of measurements. The lower represents the averages of all the measurements. ''92 OSBORN, CRANIOMETRY OF THE EQUID. Naso-FRONTAL SUTURES. The accompanying diagrams exhibit the extreme variations in the forms of the naso-frontal suture in the horse (left) and in the ass (right). The middle figure represents the Grevy’s zebra in which the fronto-nasal suture is seen to correspond closely with that of the ass although the elongate and narrow cranial proportions correspond with those of the horse. Fig. 15. Skulls of horse (left hand), domestic ass (right hand), and of zebra (center), showing characteristic differences in shape of nasal bones. Note unlikeness of fronto-nasal sutures and naso-lachrymal sutures in the two skulls (horse, Amer. Mus. spec. 16613; zebra, EL. grevyt, Chubb No. 65; ass, Amer. Mus. spec. 13982). Pe OCTPaALY, THE BENDING OF THE FACE ON THE CRANIUM Among the many authors who have more or less directly contributed to this subject are: Riitimeyer, 1882; Flower, 1891; Lankester, 1902; Ewart, 1907. The whole literature and philosophy of this upward and downward deflection of the face on the cranium has not as yet been thoroughly reviewed by anyone. It appears that there is a general functional adaptation expressed in this deflection in which the following principles appear: (1) in young animals the palatal and cranial lines are more nearly in the same plane; (2) in certain animals the deflection increases rapidly with age; (38) a horizontal and upward deflection is generally characteristic of primitive browsing types; (4) the downward deflection of the face and palate is highly characteristic of certain grazing types. '' OSBORN, CRANIOMETRY OF THE EQUID. 93 2 a = I a CRANIO OSBORN B PALATAL ANGLE Cd] tannesteR B > Fig. 16. Cytocephaly, modes of measuring cranio-palatal angle. Lower: Lankester’s system. Upper: Osborn’s system. Ritimeyer (1882! ) makes the following observations as to the deer (p. 12): “The very much extended, almost cylindrical form of the skull is characteristic for Cervide. This is due chiefly to the uniformly horizontal axis of the skull, which rarely shows the deflection so conspicuous in most of the Cavicornia, either from their youth or appearing in the course of their later growth and development. The straightness and cylindrical shape of the skull is most pronounced in the adult deer; but from the beginning the cranial portion of the skull is longer and flatter relatively than in the Cavicornia.” Lankester (1902) published a series of more extended observations suggested by his studies of Okapia.2 In this paper are given figures of vertical longitudinal sections of skulls of giraffe, elk, waterbuck, domestic goat. For the facial plane a line is drawn connecting the anterior border of the premaxillaries and the foramen magnum. This line is called the ‘palatine hori- zontal.’ His chief observations are as follows: The skull of the Okapi, p. 285, is not only astonishingly long in the postorbital region, but it presents a great peculiarity in the fact that this region is in the same horizontal plane as the preorbital region. In fact, the basicranial axis and the basifacial axis of the Okapi’s skull appear to lie nearly (not quite) in one plane. This is a primitive and unusual character. It is more or less retained by the genus Alces and others of the Cervidee, where the planes of the two axes form a very open angle; but in the Bovide a much less obtuse angle is formed... . . The whole of the brain-case or postorbital region of the skull of the Bovidee appears to be bent down as on a joint formed across the junction of the cranial and facial portions of the skull. There is good ground, p. 286, for connecting the presence of the deflection of the cranial cavity, above noted, with the mechanical conditions arising from the use of horns having the position and direction of those found in the Bovide and the Giraffe. Even in the hornless females of certain Antelopes the deflection is maintained. The absence or small size of the deflection is almost certainly to be regarded within the group Pecora as a primitive character, and in the horizontality of the skull or tendency to coincidence of the basicranial and basifacial axis-planes the Okapi bears evidence of (1) being relatively primitive, (2) not having had horn-bearing forebears, (3) of being remote from the cavicorn stock. 1 Riittimeyer, L. ‘Studien zu der Geschichte der Hirschfamilie. I Schiidelbau.’ Verh. d. Naturforsch. Gesell. Basel, Bd. VII, pp. 3-61. Year on separate, 1882; on bound vol. of Verh. Nat. Ges., 1885. 2Lankester, E. Ray. ‘On Okapia, a new Genus of Giraffide, from Central Africa.’ Trans. Zool. Soc. London, Vol. XVE Pt: Vi, August, 1902. ''94 OSBORN, CRANIOMETRY OF THE EQUID. Ewart (1907) first, to our knowledge, applied cytocephaly as a means of distinguishing various phyla of horses. His principal observations are as follows: The extent to which the face is bent down on the cranium is made evident when outlines of photographs of side views of skulls are placed one above the other, and when the angle formed by the line of the palate with the basi-cranial axis is given. The elk and sheep illustrate the two types of skull. The difference in these two forms is perhaps accounted for by the fact that the elk is a short-necked forest form adapted for feeding on shrubs and trees, 7. ¢., for holding the head in a nearly horizontal position, while the sheep is a denizen of the mountains, adapted for holding the head when feeding in a nearly vertical position. There are excellent reasons for believing that a bent skull greatly facilitates feeding on very short herbage. Ina sheep when feeding the grass is pressed by the sharp-edged lower front teeth against the hard pad attached to the upper jaw; and then the head is as a rule jerked rapidly forwards, with the result that the grass is severed partly by cutting and partly by tearing. In the Steppe variety of the horse the head instead of being invariably jerked forwards, is sometimes moved forwards, sometimes backwards, but more frequently from side to side. If, as Lankester in his paper ‘On Okapia’ suggests, the deflection in Bovide and the Giraffe is connected with the use of horns, it should doubtless be regarded as due to the downward bending of the cranium on the face. If the deflection is connected with grazing, with feeding on short herbage close to the ground, it might be more accurate to regard it as due to the bending of the face on the cranium. In the Forest variety the hard palate is nearly parallel with the basi-cranial axis, but even in such skulls the face is not in line with the cranium, for when a line is carried through the basi-cranial axis it emerges a considerable distance above the level of the incisors. In members of the Forest variety the face at birth is nearly as bent downwards as in adults of the Steppe variety. The reason may be that in animals which suck standing the necessary pressure on the mammary gland is more easily exerted on this account. Thus in the young giraffe the face is as much deflected as in fullgrown sheep and goats. But though the deflection may facilitate sucking it was probably originally acquired to facilitate grazing, which during the first year is very dfficult on account of the long legs and the short neck. In the Steppe horse the face gradually unbends in the first year, so that the skull of a 15 months Prjevalsky resembles that of an adult Forest horse. During the second year the face begins to bend down again. Thus it repeats during its growth one of the most striking characters of its remote forest-haunting ancestors. Moreover, since the bending of the face has probably been effected very gradually (since forests and marshes were abandoned for a free life on open plains) it follows that the Steppe variety branched off from the common stem at a very remote period. In Neohipparion of the Miocene, Hipparion of the Pliocene, and E. scotti of the Pleistocene, the face is strongly bent down on the cranium. CYTOCEPHALY IN THE ARTIODACTYLA. Extremes of Palatal Deflection. The accompanying figure (Fig. 17, bottom of diagram) illustrates the extreme range of adaptation as observed in the various Artiodactyla. The tran- sition from the more horizontal to the more oblique position may be illustrated in the following genera. Caribou, Rangifer, adult. Moose, Alces, young. Giraffe, Giraffa. Barbary sheep, Ammotragus tragelaphus. Hartebeest, Bubalis. The Aoudad or Barbary wild sheep, Ammotragus (Ovis) tragelaphus, presents an extreme example of the progressive deflection of the palate, on the cranium with advancing age as shown in Fig. 17; also in the following succession of measurements: At birth (average) 25.5° Male (horns 3 in.) 30° Male (horns 3 in.) 47° Male (adult) 5a0° '' OSBORN, CRANIOMETRY OF THE EQUID. 95 HORSE I E PRIEVLSAY I ZEBRA OONKEY = HORSE Equines - various 3-15 rears L (AVERAGE) Zebras (Burchell3) AT BIRTH (AVERAGE) Bast- alatal line MALE (HORNS th) MALE (HORN 3 IN) Ovis tragelaphus MALE (aoucr) RANGIFER ALCES-ADULT ALCES - YOUNG I GIRAFFA ovis EVBALIS - I y Artiodactyla + various Fig. 17. Cytocephaly. Palato-cranial Angle in various Ungulates. Horses, Asses, Zebras. Various. Burchell’s Zebras. Growth stages, deflection increasing with age, 1-2 days to 15 years. Aoudad, Barbary Wild Sheep. Growth stages, deflection increasing with age. Reindeer, moose, giraffe, sheep, hartebeest. Deflection characteristic of browsing (Rangifer, Alces, Giraffa) and grazing (Ovis, Bubalis) types. ''96 OSBORN, CRANIOMETRY OF THE EQUID. PaLATO-CRANIAL ANGLE IN Horsss, ASSES, AND ZEBRAS. ADULTS. The existing common domesticated horses are polyphyletic, so that, as shown in Fig 17, I (top), they exhibit a range of deflection, or cytocephaly, which falls both above and below the elements presented by the Przewalsky, the zebra, and the ass. The averages are as follows: Horse, not including E. przewalskii, average = 19.23° Zebra, chiefly E. burchelli, ue — 2242 The range of variation is shown in the following table: Horse: Percheron (Am. Mus.) Stage 8-+-about 10 years 10.° Arab (Am. Mus.) ie 8-about 10 years Loe Horse (New York, Am. Mus. No. 14132) - 9-15 “ 165° Horse (Am. Mus. No. 16278) = 9-15 “ Le Horse (New Mexico, Am. Mus. No. 15753) eee 18.5° Horse (Indian Territory, Am. Mus. No. 19173) os 20,5 Horse (Wyoming, Am. Mus. No. 16279) oe Oa 20-7 Horse (New York, Am. Mus. No. 14131) “8+ -about 10 years 21.° Horse (Am. Mus. No. 16275) - 9-15 213 Horse (Texas, Am. Mus. No. 16613) fe 8-5 years 225° Horse (Am. Mus. No. 15093) S 9-15 “ a ole E. przewalskit (Chubb, hybrid) - Ba. 7. Lo Zebras: Burchell’s Zebra (Am. Mus. No. 14096) Stage 3-1 years 19.° Grevy’s Zebra (Am. Mus.) . 5+-3 “ a1 Burchell’s Zebra (Am. Mus. No. 35153) s 9-15 “ 23° Burchell’s Zebra (A. M. No. 35167) - Ey So DA: Burchell’s Zebra (Am. Mus. No. 45) G5 20. Asses: Donkey (Mexico, Am. Mus. No. 13982) Stage 5-2 years 205° Donkey (Jamaica, Am. Mus. No. 15675) . 5 " D3.e Progressive Deflection with Age in Zebras. In a small collection of zebra skulls in the American Museum there appears to be evidence of progressive deflection of the palate on the cranium with advancing age as represented in Fig. 17, I, and in the following table of measurements, which, however, are less reliable because taken from different species. : Zebra, Burchell type (Chubb, No. 20) Stage 1 36 hows old = 155- Zebra (Am. Mus. No. 14096) oe Sonontis > — 19" Zebra, Grevy’s type (Chubb, No. 65) as 3 years = 21 Zebra, Burchell type (Am. Mus. No..35153) eS = 2a Zebra (Chubb, No. 47) is About 13th year = 22.° Zebra, Burchell (Am. Mus. No. 35167) nes ; Se Zebra (Chubb, No. 45) “ 36 About 4th year «= 25." IV. CRANIOMETRY AND ODONTOMETRY IN PALZONTOLOGY. In fossil skulls the indices lose value because the slightest degree of crushing or distortion seriously disturbs an index. Nevertheless the indices and ratios should be used wherever obtain- able. Since fossil skulls and dental series are rarely complete or perfect, the paleontologist requires an additional series of detailed measurements of parts of the skull not needed by the zoologist. '' OSBORN, CRANIOMETRY OF THE EQUID. 97 1. Measures of Equal Value in Recent and in Fossil Skulls where obtainable. 1. Vertex length 11. Occiput vertex angle 2. Basilar length 12. Occiput height 3. Frontal width 13. Orbital index 4. Cephalic index 14. Vomerine index 5. Facial length 15. Naso-frontal suture 6. Facio-cephalic index 16. Diastema length 7. Cranial length 17. Muzzle width 8. Cranio-cephalic index . 18. Molar-premolar series length 9. Palatal index ; 19. Dental index 10. Palato-cranial angle 20. Molar index 21. Facio-cranial angle. 2. Jaw Measurements. Identical with those in recent jaws. 3. Additional Measurements Desirable in Fossil Skulls. 22. Zygomatic width, width across zygomatic arches. This measurement in titanotheres and many other Ungulates is taken as a means of computing the general ratio between the width and length of the skull. 23. Palatal length, along the middle line from the incisive border to the posterior median line of the palate. 24. Palatal width. a. Between M! and M! of the opposite ae wT and P* ee: s c. At narrowest point of the palate. 25. Condylar width, i. e., width across the two occipital condyles at their widest part. 26. Cranial depth, a. ‘Taken opposite posterior alveolus of M'. b. Taken in a line opposite posterior alveolus of M?. 4. Measurements of Dental Serves. The entire system of measurements of the superior and inferior dental series embraces: (1) the height and width of the crowns of each of the incisor teeth; (2) the vertical, antero-posterior, and transverse diameters of the crowns of the canine teeth; (3) the diameters and indices, antero-posterior, transverse, and vertical, of each of the grinding teeth. The entire series of measurements desirable in the grinding teeth is as follows: 18. Molar-premolar series (upper), P?-M® (measured along the middle line of the wearing surfaces). 19. Dental index, as in recent skulls. 20. Molar series, total length (upper), measured along middle line of the wearing surfaces. 26. Premolar series total length (lower), Ps—Pa. - 27. Antero-posterior diameters of Ps, Ps, Ps, Mi, Mz, Ms. 28. Transverse diameters of P2, Ps, Ps, Mi, Ma, M3 (measurements of lower teeth taken in same way as those of upper in every case). 21. Premolar series total length (upper), P?-P! (measured along middle line of wearing surfaces). 22. Antero-posterior diameters of P®, P?, P!, M', M?, M? (measured along middle line of wearing surfaces). 23. Transverse diameters of P?, P’, P!, M', M?, M? (greatest diameter taken exactly at right angles to middle line of tooth). Approximately the same as Gidley, who measures across mesostyle and posterior half of protocone. 24. Molar-premolar series total length (lower), P2—-M3. 25. Molar series total length (lower). The index of each grinding tooth, that is, the ratio of the transverse to the antero-posterior diameter, is extremely important as indicative in certain mammals of the dolichocephaly and brachycephaly of the skull. In certain titanotheres, for example, we obtain “brachycephalic indices’ of each of the grinders where the width equals or exceeds the length of the grinder; also ‘dolichocephalic indices’ of single tecth where the length exceeds the width of the grinder. In the horses it is extremely important to observe a fact pointed out eae oe aa 1 Gidley, J.W. ‘Tooth Characters and Revision of the North American specie of the Genus Bee Bull, Amer. Mus. Nat. Hist., Vol. XIV, Art. rx, May 31, 1901, pp. 91-142. ''aS. OSBORN, CRANIOMETRY OF THE EQUID. and overlooked by Ewart, namely, that the ratios of the two diameters change with the degree of wear. Gidley observes: “Thus in the little worn condition of these teeth in a young horse especially before the teeth have worn to that stage where the transverse diameter is greatest, the antero-posterior diameter is always greater than the transverse. As the crown wears away, the antero-posterior diameter diminishes and a stage is reached where the two diameters are about equal, then, as the antero-posterior becomes still more shortened the transverse exceeds it.” Gidley’s observations on Equus are so important in the practice of odontometry that it appears desirable to reprint his statement as a whole (op. cit., pp. 96-103). (I). Effects of Wear on the Proportions of the Teeth. 1. The Teeth Taken Individually. Unlike the degree of complexity of the enamel foldings, the corresponding diameters are affected differently by wear in different teeth of the molar-premolar series. The same general rule for the change in ratio of the antero-posterior to transverse diameter may be applied to the intermediate teeth p* to m? inclusive, but the most anterior and posterior teeth (p? and m’) are affected differently, in this respect, from the intermediate teeth of the series and from each other as well. a. Laws Governing the Changes of Diameters of the Tooth Crowns. There seems to be no exception to the following laws for the changes of diameters of the tooth crowns as they are worn away by use. TaBLE I. MEASUREMENTS ON THE TRITURATING SURFACES OF THE UPPER TEETH OF EQUUS CABALLUS AND EQUUS ASINUS. APPROXIMATE} CATALOGUE DIAMETERS IN MM.1 Duscnrerion. AGE, NuMBER 2 3 4 1 2 3 Dp Dp | Oe Te ool pA 0 ace | ( Antero-posterior.......... 40 32 30 © | 30 30 29 Large draught Horse 9 5 years 16274 (ETAMSVEISC: oe ee. 27 28.5 127 5-27 26.5 2 2125 | Antero-posterior, Protocone) 11 14.5 | 15 13.5 | 15.5 | 16 nl eAMUELO-pOSb. 22 es are 36.5 | 29 28 25:9} 209. 122925 = . ‘ tt See 14131 Pe MIBVOISOs ke ea | 25.5 | 28 Oi, 26.5 | 26 22 Antero-post., Protocone...| 10. 13.5 | 14 13) alo 16 F Pare ibe 98.5 | 29.5 | 985 24 95 og R . e oe Sere 15+ “ 289 SRGAMSVErSe 0 2 23 Beas 28 26° 125.7) 2225) Antero-post., Protocone.../ 9.3 | 13 13 11.54) 12) 53 shes Antero-post.............. |-40 | 20.5 | 80: 127. 4 By te . ot it No number Tete cic | 28 30 28.5 | 28 26.57} 23 Antero-post., Protocone...| 10 14 "| 14 | 13 138.5 | 18 pee eee ( Anterospost... 86. | 26 | 25. }21087 2a ee 2 20 = years 16277 SUPANSVErSe. 2 eG 24 260.1 Qe Qa Zo 22.5 Antero-post., Protocone...| 9 10 dt 10 12h os | ao ANGerO=pOsts. hse 33:0 | 25.5 | 25.5 1-22 5 | 28-5326 5 a ore |. 20 16275 ATANSVerse, 7 ee | 23 26.3 | 26 25 25 23 Z Antero-post., Protocone...| 10 12 13,021 13-59) I 14 5 Loaned by Amitero=pOSt.3. 6) ao20 127.0 | 28 25 25.5 | 20.5 2 | Thoroughbred 2 6 . CR, Knight) 4: @ransverse..<. 2. i. re. 25 2625' | 26.5 125.0 1°25 2135 Antero-post., Protocone...| 8.5 | 13 13.5 | 13-9 | 13.5) 15 ANCErOspOSt~«. 2.0... 35 28 26 24 24.5 | 28.5 Texas Pony on 7 o No number sPrans Verse. 23 25.0 |) 25 25-2 224 21.5 Antero-post., Protocone...| 8 10.54) 10.7 |) 10752 105: etl ( Antero-post. 62055. 33 26 25 22 23 23 oo Domestic Ass 9 5 years 15675 DV ANISVCISCs) 2 hate he OAR | 25 24 2229 | 2275) | 2180 R Antero-post., Protocone...| 6.5 | 9.5 | 10.5} 8.5 | 9.5 | 10.5 5 Antero-post. 222. 31 OA 5 10235 1221 5 | te 5 |e 2 | Mexican Burro J One No number SETANSVECISO.) eas 23 24.3 | 24 23.3 | 2118 Antero-post., Protocone...| 6.5 | 8.5 | 9 9 | 9 10.5 1 In every case the transverse diameters were measured across from the exterior ridge of the mesostyle to the exterior wall of the posterior lobe of the protocone, exclusive of cement. [Total length of three of the above skulls are as follows: No. 16274, 604 mm.; No. 14131, 573 mm.; Texas Pony, 515 mm.] '' OSBORN, CRANIOMETRY OF THE EQUID. 99 (1) The antero-posterior diameters of the grinding surfaces of all the intermediate teeth are greatest at the stage when the tooth has just fully come into use, that is, when about one-half of an inch, or less, of the crown has been worn away ; from this point the antero-posterior diameter diminishes very rapidly for a short distance and then continues to diminish more gradually to the roots of the tooth. (2) The antero-posterior diameter of the first premolar (p?) remains about the same for the whole length of the crown, except that sometimes it narrows slightly near the roots. (3) The antero-posterior diameter of the last molar (m3), however, is relatively small at first, and increases continually as the tooth is worn away. (4) When the teeth first come into use the transverse diameters of all the teeth of the series are quite narrow, owing principally to the rapid incurving of the ectoloph; this diameter increases very rapidly for about one-half to three-fourths of an inch, but from this point to the roots of the teeth the transverse diameters of p* to m? inclusive remain about the same, diminishing slightly near the roots; p? gradually diminishes while m increases in transverse diameter as the crown wears away. (5) The antero-posterior diameter of the protocone in all the teeth of the series remains the same for the whole length of the crown. (6) The antero-posterior or long diameter of the incisors diminishes with age while the transverse diameter increases. b. Effect of Wear on the Relative Measurements of Tooth Crowns. (1.) Ratio of the antero-posterior to the transverse diameter. It will be seen from the foregoing that owing to the very slight variation of the transverse diameters of the crowns of p’ to m? inclusive, for almost their entire length, and to the great shortening of their antero-posterior diameters, the ratio of these diameters in these teeth is very different in old and in young individuals of the same species. Thus in the little worn condition of these teeth in a young horse, especially before the teeth have worn to that stage where the transverse diameter is greatest, the antero-posterior diameter is always greater than the transverse. As the crown wears away, the antero-posterior diameter diminishes and a stage is reached where the two diameters are about equal, then, as the antero- posterior becomes still more shortened, the transverse exceeds it. In every series this variation in ratio seems always to be most advanced in m! and m2. This is evidently due not only to the order in which the teeth of the horse come into use, whereby the first to appear would at a given stage be most worn, but also, as is shown by an examination of Table I (p. 98), because the range of reduction of the antero-posterior diameters is greater in the molar than in the premolar teeth. M1! is always the most advanced, as it comes into use before any of the others of the permanent set." (2.) Ratio of antero-posterior diameter of the protocone to the antero-posterior diameter of the crown. The antero-posterior diameter of the protocone, being, like the transverse diameter of the crown, practically unchanged through wear, also holds to the ever-changing antero-posterior diameter of the crown in the old and much worn tooth a very different relation from what it did when the tooth first came into use. Thus, it may happen that in a little worn tooth the antero-posterior diameter of the protocone is much less than half that of the entire crown, but may become greater than half this diameter when the tooth has become much worn in consequence of this shortening of the antero-posterior - diameter of the crown. 2. On the molar-premolar series as a whole. The shortening of the antero-posterior diameters of all the other teeth in the series, except p’, is not nearly compensated by the lengthening of this diameter in m’, hence it results that the series, as a whole, becomes much shortened and the teeth from behind crowd forward toward p? which retains the same relative position in the skull, so that m* shows the greatest displacement and the discrepancy in length is all taken from the posterior end of the series. The gap which would other- wise be left in the maxillary bone behind m* becomes gradually filled in with a new growth of bone, as the teeth shift for- ward, leaving a flattened ridge which is continuous with the rugose prominence or ridge which marks the posterior extension of the maxillary bone beyond m’; hence the length of this posterior extension of the maxillary depends principally upon the age of the horse. In passing from the young to the old stage, there is also a marked change in the relative position, with respect to the molar teeth, of the anterior projection of the maxillary ridge, the post-palatal foramina, and the anterior projection of the post-palatal notch,— all appearing relatively more posterior in the old individual... . b. Effect of individual variability on dimensions of the teeth— It has been shown under the topic of age variations that the transverse diameters of all the superior molars and premolars, except p? and m°, the antero-posterior diameters of the protocones of all the teeth and the antero-posterior diameter of p* are measurements which change but slightly for 1The order of appearance or eruption of the permanent teeth of the large species from Texas (Z. scotti) is the same as Owen has given for E. caballus, and is: first, m1, second, m2, third, p?, fourth, p*, fifth, p14, sixth, m%. This is probably the order in all other fossil species of this genus. ''100 OSBORN, CRANIOMETRY OF THE EQUIDZ. much the greater length of the crown; hence, unless specimens of the same age are taken for comparison, it is in these measurements that one should look for evidence of individual variability. Careful measurements of the teeth of more than ten specimens of E. caballus have led to the following conclusions: (1) The transverse diameters of the corresponding teeth for p* to m? inclusive are remarkably constant, especially in skulls of nearly the same size; the greatest difference in a certain series of four skulls of large draught horses examined not exceeding 2.5 mm., and in another series of three skulls belonging to animals about the size of carriage horses being less than 1 mm. (See table of measurements, p. 98). The transverse diameters of m? of the large series vary only .8 mm., while in the small series all the transverse measure- ments for this tooth are the same. ‘The greatest difference in the transverse diameter of m?, including both series of skulls, is only 1.5 mm., and adding a skull of the Texas pony to the list the extreme difference between the transverse diameter of m? of this whole lot of skulls, ranging in size from the large draught horse to the small Texas pony, is only 2.5 mm. It will be seen by reference to the table of measurements (p. 98) that the average variation of the corresponding transverse diameter for all these teeth is very small considering the great difference in size of the animals represented. It seems reasonable that much greater variations of the comparatively constant characters of the teeth would be found in FE. caballus than in the extinct species, since in this species domestication and breeding have caused such a very wide range in size and proportions of the individuals; hence, when, in two lots of fossil horse teeth, the difference between the transverse diameters of corresponding teeth is on the average greater than that between the large and small varieties of E. caballus, it would seem that the teeth of the two lots could scarcely belong to the same species, and although the character of size, alone, could hardly be considered sufficient ground for establishing a species, yet where this difference exists, it seems reasonable to expect that when skulls which represent such two lots of teeth are known, other differences will be found which will clearly mark them as distinct species. It has been shown that the antero-posterior diameter of the protocone is very little affected by wear; Table I (p. 98), makes it clear, however, that the range of individual variability of this diameter is very great, and cannot be depended upon as a distinguishing character even in corresponding teeth of individuals of the same size. BIBLIOGRAPHY. Franck, L. 75. Ein Beitrag zur Rassenkunde unserer Pferde. Landwirtschaftliche Jahrbiicher, IV, 1875. RUtmeyer, L. ’82. Beitriige zur Geschichte der Hirschfamilie. I. Schidelbau. Verhandl. Naturforsch. Ges. Basel, Pt. VII, Vol. I, 1882. Branco, W. and Reiss, W. ’83. Uber eine Fossile Siugethier Fauna von Punin bei Riobamba in Ecuador. Paléiont. Abhandl. Dames u. Kayser, Vol. I, no. 22, 4to, Berlin, 1883. NeEsrRInG, A. ’84. Fossile Pferde aus deutschen Diluvial-Ablagerungen und ihre Beziehungen zu den lebenden Pferden. Ein Beitrag zur Geschichte des Hauspferdes. Sonderabd. Landwirtschaftliche Jahrbiicher, 1884. TscHerskl, J. D. 92. Beschreibung der Sammlung posttertiérer Séugethiere. Mém. Acad. Imp. Sci. St. Pétersbourg, Ser. 7, Vol. XL, mo. 1892) pp, toll. Osporn, H. F. 02. Dolichocephaly and Brachycephaly in the Lower Mammals. Bull. Amer. Mus. Nat. Hist., Vol. XVI, Art. vii, Feb. 3, 1902, pp. 77-89. LANKESTER, E. R. 702. On Okapia, a New Genus of Giraffide, from Central Africa. Trans. Zool. Soc. London, Vol. XVI, Pt. VI, August, 1902. SALENSKy, W. 702. Equus przewalskii Poljak. Wissensch. Resultate, N. M. Przewalski nach Centrale-Asien Unternommenen Reisen auf Kosten einer von Seiner Kaiserlichen Hoheit dem Grossfiirsten Thronfolger Nikolai Alexandro- witsch. Published by Kaiserlichen Akad. d. Wissenschaften. Zool. Theil. Bd. I Mammalia, Abth. 2 Ungu- lata, Lieferung 1. 76 pp. 4 pll. St. Petersburg, 1902. 07. Prjevalsky’s Horse. Translation by Capt. M. Horace Hayes and O. Charnock Bradley, with an introd. by J. C. Ewart. 8vo, London, 1907. Ewart, J. ©: 07. On skulls of Horses from the Roman Fort at Newstead near Melrose, with Observations on the Origin of Domes- tic Horses. Trans. Roy. Soc. Edinburgh, Vol. XLV, Pt. III, no. 20, 1907, pp. 555-587. Brapuny. O, C. : : 07. Craniometrical Observations on the Skull of Equus prjevalskii and other Horses. Proc. Roy. Soc. Edinburgh, Wool XOXV Bt Nes; 1907, pp: 46-00. '' '' '' '' '', the Law of the Four | Factors of Evolution BY HENRY FAIRFIELD OSBORN, LL.D., Sc.D. RESEARCH PROFESSOR OF ZOOLOGY IN COLUMBIA UNIVERSITY, CURATOR EMERITUS OF VERTEBRATE PALZONTOLOGY IN THE AMERICAN MUSEUM OF NATURAL HISTORY S : ’ PHILADELPHIA 1912 '' '' PREFACE. During the last three years the author has made three preliminary statements of the law which is elaborated in this contribution. It belongs to an extended treatise which the author has had in preparation since 1905. It is regrettably incomplete in failing to give concrete examples from nature and from experiment of each of the series of subordinate principles set forth. It seemed best, how- ever, to publish the central principle, or Law of the Inseparable Factors, in its present form rather than to delay longer for time and opportunity to publish the concrete examples. The author takes advantage of the present opportunity to cite from his purely biological contributions,’ begun in 1889 and continued to 1912, which form a progressive and more or less connected or consistent interpretation of the evo- lution phenomena as observed by a paleontologist. 1 See full bibliography on page 307. 275 '' '' TETRAPLASY, THE LAW OF THE FOUR INSEPARABLE FACTORS OF EVOLUTION. By Henry FarrFietp Ossorn, LL.D., Sc.D. CavsaATION or Causality (Lat. causa, derived perhaps from the root cav- as in caveo, and meaning something taken care of; corresponding to Gr. «/r/a), a philosophical term for the operation of causes and for the mental conception of cause as operative throughout the universe. The word ‘“‘cause”’ is correlative to ‘‘effect.”” Thus when one thing B is regarded as taking place in consequence of the action of another thing A, then A is said to be the cause of B, and B the effect of A. . . . The senses can say only that in all observed cases B has followed A, and this does not establish necessary con- nection. ... J. S. Mill argues that, scientifically, the cause of anything is the total assemblage of the conditions that precede its appearance, and that we have no right to give the name of cause to one of them exclusively of the others. ... (3) Efficient cause (4pxh Tijs xwjoews), the alcohol which makes a man drunk, the pistol-bullet which kills, This is the cause as generally understood in modern usage. . . . Vera causa is a term used by Newton in his Principia, where he says, ‘‘ No more causes of natural things are to be admitted than such as are both true and sufficient to explain the phenomena of those things’’; vere cause must be such as we have good inductive grounds to believe do exist in nature, and do perform a part in phenomena analogous to those we would render an account of. (11th Edition Encyclopedia Britannica, pp. 557-558.) Cause and effect are not two but one. That they are inseparable is indeed recognized by the relativity of the very terms themselves . . . in content they are absolutely identical. It is only in form that they can be distinguished, and then we may speak of the one as determining, and of the other as determined. (Welton, Manual of Logic, vol. II, p. 25.) CONTENTS. Page Introduction. . 0. 8. Od 278 How the Problem is Approached: .0))...222 CUS (ee 278 Reasons for the Historical Method of Analysis...................... 280 Special Action of Each of the Factors.) 2) 0000009 Oe 284 1. Environment.:..025000000.00 UP a 284 2. Ontogeny... oc cence cell. RI a ee 285 3. Heredity, including Variation j 9:66. io esae a ee 291 4° Beleetion: 200 SO ee OO ee 294 Coincident, Selection iui) .6 ccc udn, enemas. 1. ee ae 295 Interaction ‘of the Pour Factots. 0. ea ee 296 1. Initiation and Genesis. . ...... 0:56. cies oi 300 2. Initiation in vironment) ey 301 3. Yattiation ih Ontogeny.) se ee 302 4. Initiation and Genesis in Heredity... 3804 5. Genesis as observed in Palwontology 0 ee 304 Summary. 20 DP oe ee eee 306 Appendix by William K, Gregory’. 25057. 0. 1077) A a) 307 Previous Papers by the Author Developing this Subject................ 307 277 ''278 THE FOUR INSEPARABLE FACTORS OF EVOLUTION. INTRODUCTION. How THE PROBLEM Is APPROACHED. The total assemblage of the conditions that precede the appearance of ‘“‘characters”’ in living animals is the subject of this contribution. To continue the paraphrase of John Stuart Mill’s argument, we have no right to give the name of cause to one of these conditions exclusive of the others. Mi§ill’s doctrine that the cause is the ‘‘sum total of the conditions, positive and negative, taken to- gether, the whole of the contingencies of every description, which being realized, the consequent invariably follows,” is the doctrine on which the Law of the Four Inseparable causes or Factors of Evolution is founded. Out of the analysis of all the series of conditions which accompany evolution it appears that these phenomena fall into four groups which center respectively around the conditions that we term Environment, Ontogeny, Heredity, and Selection. In the causes and effects resident in these complexes of conditions are to be discovered the origins of new characters and the transformations of existing characters. This contribution to biology is associated with the name of Joseph Leidy, one of the most distinguished members of this Academy, through the fact that he was one of the first to make known the extinct family of perissodactyl quad- rupeds now popularly termed the Titanotheres. In attempting to monograph this family from very abundant materials, the author of the present memoir found that these materials presented a rare opportunity of contributing what appear to be new data on the two chief phenomena of evolution above mentioned, namely: the origins of new characters, the transformations of existing characters. To understand these origins and transformations the author was led to the above-mentioned analysis of all the evolutionary phenomena and conditions.’ This analysis resulted in the conclusion that all these phenomena are comprised, as indicated above, under four relations or complexes of causes and effects. These relations are external and internal as follows: External; ENVIRONMENT = the sum of all external conditions. Internal: ONTOGENY = the sum of all conditions of individual, bodily, or somatic develop- ment. Internal: Hrrepity = the sum of all germinal or blastic forces and conditions. ; External: SELECTION = the sum of all competitive relations between individuals in the struggle for existence. The true conception of individual animals or plants, of varieties, of species, is that they are all complexes of the relations of these four sets of conditions, which for the sake of simplicity may be called Factors, although the word factor does not exactly express our meaning. A factor is defined as ‘‘one of several - circumstances, elements, or influences which tend to the production of a given 1 Osborn, H. F., The Ideas and Terms of Modern Philosophical Anatomy, Science, N.S8., vol. XXI, No. 547, June 23, 1905, pp. 959-961. Jour. of Philosophy, Psychology and Scientific M ethods, vol. II, No. 17, Aug. 17, 1905, pp. 455-458 (condensed). '' THE FOUR INSEPARABLE FACTORS OF EVOLUTION. 279 result.” Thus, for example, it is claimed that “light is the most powerful factor amongst all the agents which influence life upon the earth.’ Thus Wallace in 1889 wrote: ‘The importance of natural selection as the one invariable and ever-present factor [italics our own] in all organic change and that which alone has produced the temporary fixity combined with a secular modification of species.” There is a perpetual balance between these four factors or complexes of con- ditions, comparable to the balance of living nature as a whole, such that any disturbance of any one factor produces a disturbance in all the other factors. This recalls to our memory one of Herbert Spencer’s definitions of life, namely, that life is a continuous adjustment of all internal relations to all external re- lations. This coincides also with John Stuart Mill’s doctrine of cause cited above as the “sum total of the conditions, positive and negative, taken together.” This conception of a multiplicity of conditions antecedent and consequent must be in the mind of every biologist who seeks to determine the causes of the origins and of the transformations of characters. This conception of inseparable relations has more than a theoretical, it has a highly practical bearing in paleontology and in experimental zoology. Granted that new characters aswell as transformations of existing characters must have origins or beginnings, in what part of the complexes of conditions do they first appear? What is invariably antecedent, and what is invariably con- sequent? Where is the initiation of the chain of conditions which leads to the genesis of new characters? This is the question which the author set to himself in the examination of every new character and of every transformation of char- acter discovered among the fossil titanotheres. Was there evidence of a dis- turbance in the balance of conditions and where did this disturbance originate? This method of analysis by which the answer shall be attained appears to be the right one, but the answer itself isnot yet. The first step is the recognition of the universal law of the inseparable actions and reactions of the four factors, a law which we propose to designate as Tetraplasy, from 7érpé (four) and Adcom (to form, mould, shape). After several years of reflection and analysis of the evolution of the titano- theres, the author first presented this law in an address before the Department of Zoology at Columbia University November 3, 1905. It was not published,’ however, until after the Boston meeting of the Seventh International Zoological Congress of August, 1907. In December, 1908, the subject was presented before 1 Osborn, H. F.: Evolution as it Appears to the Paleontologist. Address before the Seventh International Zoological Congress, Section of Paleozoology, Boston, Aug., 1907. Science, N.S., vol. XXVI, No. 674, Nov. 29, 1907, pp. 744-749. Proc. Seventh International Zool. Congr., Boston Meeting, Aug. 19-24, 1907, Cambridge, Mass., 1912, pp. 733-739. The Four Inseparable Factors of Evolution. Theory of their Distinct and Combined Action in the Transformation of the Titanotheres, an Extinct Family of Hoofed Animals in the Order Perissodactyla. Science, N.S., vol. XXVII, No. 682, Jan. 24, 1908, pp. 148-150. Biological Conclusions Drawn from the Study of the Titano- theres, Science, N. 8., vol. XX XIII, No. 856, May 26, 1911, pp. 825-828. ''280 THE FOUR INSEPARABLE FACTORS OF EVOLUTION. the American Society of Zoologists at the New Haven meeting and published a few weeks later under the title The Four Inseparable Factors of Evolution, Theory of their Distinct and Combined Action in the Transformation of the Titanotheres, an Extinct Family of Hoofed Animals in the Order Perissodactyla. REASONS FOR THE HistToricAL MretHop or ANALYSIS. The reason perhaps that this law is connected with the study of an extinct group of animals is that the hypothesis of the simultaneous operation of several factors on different groups of characters could only suggest itself to a pale- ontologist working upon a very complex organism, in which an almost countless number of characters are simultaneously evolving. A gross representation of what is observed as to ‘‘characters”’ in the study of a number of different organisms through successive phyletic stages is seen in the following scheme. o Cerdrogiteniot 3 Progressive Steps. a A 5 AC 2° Sra ee Be Line aoa oC DA ons 458° 62 81 Of s 2 t OF Go 12 Heal 42, 3.24, 5 LL a: 2 bows Ko 2.3 Gel 23 4 1M This representation is gross because (1) the progressive and retrogressive steps instead of being sharply defined, as indicated by numerals 1, 2, 3, 4, and so on, are delicately intergraded; (2) the number of characters is very much larger than indicated above. It is estimated that there are at least 560 independent “‘characters’’ evolving simultaneously in the grinding teeth alone. The paleontologist is in a peculiarly favorable position to observe both the origins of new characters and the transformations of existing characters as visible modes of evolution: the paleontologist enjoys the unique advantage denied to all other zoologists of being present before the birth, at the birth, during the progress, into the rise, decline and death of new characters and organs. The genesis of single characters is apparently a simple, concrete problem. When and how does a new cusp arise on a grinding tooth? When and how does a new horn arise on a skull? When and how does a broad skull change into a long skull, and vice versa? In seeking, however, to explain the conditions. or causes which give rise to these things it is necessary for the palzeontologist to remind himself constantly of the following important distinction: '' THE FOUR INSEPARABLE FACTORS OF EVOLUTION. 281 New hereditary characters appear in the body. New hereditary characters reside in the germ. The paleontologist is only observing appearances. From the laws governing these appearances he must deduce what is going on in the germ. It is necessary for him, therefore, to secure a broad biological foundation for his observation and reasoning and to see how far the phenomena which he observes, or thinks he observes, square with the phenomena observed by special students of ontogeny, selection, and heredity. The body may be distinguished as the soma and the germ as the blastos. 'To designate new characters which apparently make their first appearance in the soma we may use Weismann’s term somatogenic; to designate those which so far as we can perceive make their first appearance from the germ we similarly use Weismann’s term blastogenic. This distinction corresponds with the two great and more or less contempo- raneous movements in living things. The first movement (ontogeny), which has been known and observed from very ancient times, is that of the developing organism, its growing and func- tioning body or soma; this movement is called ontogenesis, or individual genesis. These are the visible expressions of the forces, potentialities, predispositions and reactions to environment derived from the germ, or heredity-substance. The second movement (heredity), which has been realized as entirely distinct only in modern times, is in the forces, potentialities, and predispositions of the germ, or blastos. Thus the two kinds of movement and genesis of single characters in all living things may be clearly distinguished as Blastogenesis, -genic, Somatogenesis, -genic. The soma (ontogeny) is relatively visible and measurable. It enjoys contact and combat with all the forces of nature, consequently some new characters may arise in the soma. It is also the carrier or vehicle of the blastos, it is the environ- ment of the blastos, or intermediary between the outer environment and the blastos. The forces of the blastos are invisible; in the mammals they are re- motely buried and removed from environment; the blastos (heredity) carries along all characters from generation to generation; it modifies characters; it loses characters; it gains characters. Whereas certain new characters which arise in the soma are merely transient and may not appear in the blastos, we have reason to believe that all new characters which arise in the blastos appear in the soma. Quite marvelous and inconceivable to us is the manner in which the infinitesimally minute blastos can carry the infinitely grand and diverse char- acters of the soma of such animals, for example, as Balenoptera, Elephas, Homo. Still more marvelous and inconceivable is the evolution of the blastos, its incessant change, its gain and loss of characters, always and continuously in the main current of adaptation, or of fitting the soma to its activities and its environ- ''282 THE FOUR INSEPARABLE FACTORS OF EVOLUTION. ment, although there may be a dysteleology of side currents, of lagging characters in which the blastos does not, so to speak, keep pace with the soma, as well as of neutral characters which do not appear to be adaptive. This mutability, or evolution movement, or whatever it is that is in progress in the blastos is the chief and most difficult matter either to explain or to form any conception of. Emphasis may be laid on the words adaptive need or necessity because there is distinct evidence in palzeontology that the blastos is not evolving alone through its internal forces independently of the external forces of ontogeny and of en- vironment, but that there is some harmony or form of interaction, the nature of which we do not understand. The appearances which the paleontologist observes in the origins of character and the transformations of character are simply modes of evolution. It is, how- ever, from the analysis of these modes of evolution as well as by experiment that we must ultimately discover the causes or factors, if, indeed, they are discoverable. These modes of evolution present themselves under such widely different aspects to different classes of observers that we are reminded of the fable of the ele- phant and the nine blind men, who formed nine entirely different opinions as to the nature of the animal through judging wholly by the respective parts with which the hands of each happened to come in contact. The stout opinions of these nine blind men have their more or less clear counterparts in modern bio- logical investigators, who are in reality studying different aspects merely of the same great phenomena. There is first the standpoint of the natural philosopher of the Herbert Spencer type; then that of the systematic zoologist and field natur- alist or botanist; of the breeder, horticulturalist, and fancier, typified by Mendel, De Vries, and Bateson, working in an entirely different field of facts; of the anthropologists and anatomists who deal again with a distinct class of phenom- ena; akin to this is the work of the student of animal mechanics. Then there are the observers of geographical distribution and segregation, such as Gulick and Ortman. In the opposite extreme the observers of cell structure, such as Nageli, Boveri, and Wilson are in a circumscribed field of their own. The experimentalist is on the borderland between the field naturalist, the student of cell structure, and the student of animal mechanics. The biological statistician has his own field of observation. The paleontologists have divided into two groups, those who have directly observed the origin and transformation of character, like Waagen and Hyatt, and those who have entered into mechanical explanations, like Cope. The subjective and objective elements in observation and induction may be expressed thus: Differences in the , : (2. the er oo ern x yon a ee x apparent modes | _ Resultant inductions SG Roel és atic observed of change in each and conclusions. ae ad of these objects | It is a familiar psychic process that predisposition for a certain theory causes corresponding aberration of vision; facts which agree with one’s theory are seen ''THE FOUR INSEPARABLE FACTORS OF EVOLUTION. = 283 and collected more readily than those which do not.