Baluchitherium, ~From the Oligocene of Mongolia

Home , Aceratherium, Indricotheriinae, Paraceratherium

Further Notes on the Gigantic Extinct Rhinoceros, Baluchitherium, ~from the Oligocene of Mongolia

BY WALTER GRANGER AND WILLIAM K. GREGORY.

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

VoL. LXXII, ART. I, pp. 1-73

'j- New York

I I88se4 Augut 21, 1936

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VOLUME LXXII, 1936

Article I.-FURTHER NOTES ON THE GIGANTIC EXTINCT RHINOCEROS, BALUCHITHERIUM, FROM THE OLIGOCENE OF MONGOLIA1

BY WALTER GRANGER AND WILLIAM K. GREGORY

PLATES I TO IV; TEXT FIGURES 1 TO 47

CONTENTS PAGE SKULL...... 2 VERTEBRAE ...... 8 RIBs...... 41 LIMBS AND FEET ...... 41 RELATIONSHIPS ...... 54 RESTORATION ...... 65 LIST OF SPECIMENS OF Baluchitherium AND RELATED FORMS FROM MONGOLIA, 1922-1930. ,O0 LITERATURE CITED ...... 72

The present article is designed to extend and supplement the results set forth in Novitates No. 78, 1923, by the late Henry Fairfield Osborn and to describe the "Baluchitherium" material obtained in the Baron Sog and Houldjin beds of Oligocene age by the Central Asiatic Expe- ditions in 1922-1930. While nothing like complete associated skeletons was found, there were enough partial associations of limb bones with vertebrae or jaws to justify a revised " paper restoration" of the skeleton of this animal, together with notes on its relationships and chief structural features. Our new restoration (Fig. 47) was first published in very brief articles in Novitates (April 1, 1935) and Natural History (April, 1935). 1 Publications of the Asiatic Expeditions of The American Museum of Natural History. Con- tribution number 135. SKULL It has already been remarked by Osborn (1923) that "the skull o Baluchitherium grangeri, while of enormous size ... is relatively primi tive in structure," and that "this is a primitive Eocene and Lowe Oligocene form of skull grown large." In this connection it is also note worthy that the pair of downwardly directed upper incisors in coopera tion with the forwardly directed pair of conical lower incisors, have everi appearance of being homologous with the corresponding teeth in Trigo nias osborni and therefore of representing i2 and ii, respectively.

Fig. 1. Baluchitherium grangeri Osborn. 'lType skull. Amer. Mus. No. 1865( X 1/1o natural size. Osborn (1923, p. 6) has also suggested that the enlarged upper inci sors functioned as defensive tusks; we feel, however, that their primar, function was to assist in the sudden jerking loose of shrubs by downwar movements of the head and neck, since they are well placed to act thu: as picks and levers, while the skull (Fig. 1) is braced to resist such stresse through its strong rostrum, down-curved zygomata and greatly empha sized basi-occipital eminence. The cheek teeth (Fig. 2) are also of primitive rhinoceros type, bu slightly hypsodont and frequently well worn. They are rather small fo such huge animals, so that the food was probably not very silicious 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 3

Close relationships on the one hand with Cooper's Paraceratherium and on the other with Borissiak's Indricotherium are indicated (see page 54 below). In general the skull of Baluchitherium is remarkable for its great length and dorso-ventral lowness; while its occiput (Fig. 3) above the condyles is extremely narrow as compared with that of the titanotheres or even of recent rhinoceroses. This contrast is associated with the absence of horns at the front end of the head, the use of which, especially in the titanotheres, generated enormous oblique stresses on the occiput and necessitated a huge development of lateral crests to withstand the pull of the immensely thick neck muscles. In Baluchitherium, on the contrary, the occipital muscles although doubtless very strong were extended vertically rather than transversely. Near the upper border of the occiput there was a very large and deep median pit for the ligamen- tum nuchae. Thus the skull seems to have been normally suspended from above and, as will presently be shown, the characters of the neck vertebrae do not suggest to us that the animal normally held its head above its back or that it browsed on branches above its head. While the occiput itself is very narrow, the occipital condyles are extremely wide. The huge size of the paroccipital processes of the ex- occipital in Baluchitherium implies great strength of the cephalo-humeral muscles. The channel for the ventral muscles of the neck on the under sides of the cervical vertebrae is remarkably wide and the tubera basi- occipitalia and associated median eminence of the basi-occipital are likewise immense. All this would enable the animal to make a powerful downward sweep with its upper incisors. In the 1928 collections are two immense occiputs, one of which (Fig. 3), as shown in the accompanying table of measurements (Table I), measures 34 cm. across the occipital condyles as compared with 31.5 cm. in the type skull of B. grangeri described by Osborn. This occiput is considerably too wide across the condyles for the atlas described by Cooper, but is apparently about the same size as the occiput figured by- Borissiak (P1. i, fig. 1). The jaw of B. grangeri as restored to fit the skull is slightly larger than the robust jaw No. 26166 (Fig. 4), which is associated with our largest humerus and radius. On the other hand, as noted above, the B. grangeri skull is somewhat smaller than our largest occiput (No. 26165). In any case the B. grangeri type skull (No. 18650) belongs with the next-to-the-largest animals and is undoubtedly much too large to go with the fore and hind feet (No. 21618) with which it is at present exhibited. ce

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C_, 6 Bulletin American Museum of Natural History [Vol. LXXII TABLE I.-COMPARATIVE SKULL MEASUREMENTS (IN CENTIMETERS) No. 18650 No. 26165 No. 26167 Boriss. B. grangeri (very large) Tab. I (type) Pmx to condyle...... 128.6 (H.F.O.) Maximum width across occipital condyle...... 31.5 29.5 34.1 Width across paroccipital processes...... 33.6 35 36.5 Height of occiput...... 33 35.5 38 p1-m3...... 40.3 Width across zygomata... 61.4 Length, condyle to middle of glenoid...... 35.5 34 34 (32 est.)

Fig. 3. Posterior part of skull of Balu- chitherium grangeri. Amer. Mus. No. 26165. X 1/io natural size.

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Fig. 4. Lower jaws of Baluchitherium grangeri. X 2/1i natural size. A, B. Large animal (size Grade II), No. 26166. C, D. Smaller animal (size Grade IV), No. 26172. 7 8 Bulletin American Museum of Natural History [Vol. LXXII

The foramina of the skull (Fig. 5) conform in general with the modern rhinoceros type but are more primitive in the separation of the alisphenoid canal from the foramen ovale. This is conformable with the shal- lowness and length of the skull. Similarly the pterygoid wings of the alisphenoid, together with the true pterygoid bones, are not deeply extended vertically as they are in modern rhinoceroses. These characters are also related with the relatively slight vertical extent of the cheek teeth. BRAINCAST.-An incomplete endocranial cast (Pls. I, II) was obtained from the type skull of Baluchitherium grangeri. It is more primitive than the brain of Rhinoceros indicus as figured by Owen (1852, P1. xix) and, as might be expected in so large an animal, the length from the frontal pole to the back of the cerebellum, measuring 125 cm., is only 8 per cent of the skull length, whereas the length of the brain of Rhinoceros sumatrensis as figured by Marsh (1884, p. 66) is 17.7 per cent of the skull length. VERTEBRAE Our collections include among others the following notable vertebrae: No. 26387 (874), sacrals, lumbars and several dorsals, associated with limb bones of moderate size; No. 26168, two enormous mid-cervicals of doubtful association, the largest vertebrae known in any land mammal; No. 26390 (914), a well-preserved axis, much too small to fit Cooper's atlas; No. 26392 (877), a seventh cervical of intermediate size; No. 26173 (731), a large first dorsal; No. 26169, a fourth (?) dorsal associated with femur, tibia, metatarsus. Close comparisons with Borissiak's plates, which figure the more or less incomplete remains of numerous vertebrae, together with the casts of the atlas and two neck vertebrae received from Professor Forster Cooper, supply data for a fairly satis- factory knowledge of the chief morphological features of the various regions of the vertebral column. As explained below (p. 65), since the bones preserved belong to adults of widely varying sizes, we have en- deavored to determine in the case of each bone what its size grade is, and for convenience we have recognized only four grades, hereafter desig- nated as size Grade I, II, III, IV, in descending order. The ATLAS (Fig. 6), as described and figured by Forster Cooper, is relatively longer and narrower than that of modern rhinoceroses. Its mid-length between the transverse planes of the anterior and posterior articular facets is 26.5 centimeters; its maximum width across the transverse process, 48.3 cm., is 1.81 times its mid-length, whereas in Rhi- o~ .0 c .I II* CQo

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Fig. 6 A, B. Atlas of Baluchitherium osborni. Size Grade II. X 2/i5 natural size. A Left side. B. Front surface. From cast. noceros sumatrensis the corresponding index is 2.86. The total width across the anterior cotyli (27.7) is, however, too small to receive the gigantic occipital condyles of our largest skull, which measure 34.5 cm. in diameter. If Cooper's atlas were widened to 130 per cent of its present width, its anterior cotyli would be 36 cm. wide and its length increased from 30 to 39 cm. This would be about the right size for the largest occiput. Hence we assign Cooper's atlas to size Grade II. The anterior articular facets or cotyli are relatively lower and the posterior articular facets of the atlas are spread out more widely than are those of the Sumatran rhinoceros, in accordance with the relatively greater width of the neck. Above the neural tunnel the spine has sub- sided into a low boss, while its anterior base, tied by ligament to the occiput, finally widens out into a forwardly projecting transverse bar which would greatly limit the dorsal extension of the occiput. 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 11

The AXIS (Fig. 7) is represented in our collection by a perfectly preserved specimen, No. 26390 (914), which in spite of its apparently gigantic size, is decidedly too small to fit Cooper's atlas and very small in comparison with our huge mid-cervicals. We therefore regard it as belonging to size Grade III. Its mid-length, from the anterior tip of the neural spine to a transverse plane touching its posterior zygapophysial facets, is 39 cm. as compared with 9 cm. in Rhinoceros sumatrensis. Its width across the posterior zygapophysial facets is 22.7 cm., which is 58

Fig. 6 C, D. Atlas of Baluchitherium osborni (continued). Size Grade II. X 2/lr natural size. C. Upper surface. After Cooper. D. Lower surface. After Cooper. .Iy I- N'

Fig. 7. Axis of Baluchitherium grangeri. Size Grade III. X 1/ natural size. A. Left surface. B. Upper surface.

12 Fig. 8. Fourth cervical vertebra of Baluchitherium grangeri. Size Grade I. X 1/5 natural size. A. Left lateral view. B. Upper surface. C. Anterior view. 13 61 I

Fig. 9 A, B. Sixth cervical vertebra of Baluchitherium grangeri. Size Grade I. Xl/6 natural size. A. Left lateral view. B. Upper surface.

14 Fig. 9 C, D. Sixth cervical vertebra of Baluchitherium grangeri (continued). Size Grade I. X 1/5 natural size. C. Anterior view. D. Posterior view. 15 16 Bulletin American Museum of Natural History IVol. LXXII per cent of its mid-length, whereas in Rhinoceros sumatrensis the width is 82 per cent of the length. Its height from the ventral keel to the posterior tip of the neural spine, 30 cm., is but 77 per cent of the mid-length, while in Rhinoceros sumatrensis the same measurement is 143 per cent of the mid-length. The width of the posterior cotylus or cup of the centrum is 18 cm., its height, 12.2 cm., so that it is 1.47 times as broad as it is high, whereas in Rhinoceros sumatrensis the width of the cup only equals the height. As compared with that of Trigonias osborni, the width across the posterior zygapophysial facets is relatively less, while the neural spine is long and very low. Thus the axis as a whole is relatively long, low and narrow but presents no feature that appears to be inconsistent with the

TABLE II.-COMPARATIVE CHARACTERS OF THE ATLAS Rhinoceros Baluchitherium Draught sumatrensis (Fig. 6) Horse Proportic ns Very short, very Long, not wide Very long, narrow wide Spine Prominent Absent Very low Anterior cotyli Large, high Very wide, tending Narrow, high, deepKly to be shallow concave Dorsal anterior bor- Much thickened, Thin, deeply der above cotyli widened, pro- notched duced forward (tied by ligament to occiput) Anterior cotyli, ven- Slightly produced Extremely produced tral border anteriorly ventrally on either side Transverse process Very wide, horizon- Elongate, wide, sub- E longat e, outer tal horizontal flange sharply de- curved Longitudinal branch Not perforating Typically perforat- Perforating trans- of tunnel for occipi- transverse proc- ing transverse verse process tal artery ess process but vari- able Posterior facets Produced postero- Produced postero- Not produced; high externally; mod- externally; shal- erate height low dorso-ventrally Median posterior Produced backward Present, abbreviate Truncate, well in ventral process (for beneath body of front of body of insertion of longus axis axis colli muscle) 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 17 hypothesis that Baluchitherium has been derived from a Trigonias-like ancestor. As compared with that of Borissiak's "Epiaceratherium," the axis of Baluchitherium is almost identical in everything but proportions, being much further evolved in absolute size and relative length (see p. 56 below). Further comparisons are set forth in the accompanying tables. TABLE III.-COMPARATIVE CHARACTERS OF THE Axis Rhinoceros Baluchitherium Draught sumatrensis (Fig. 7) Horse Proportions Short and high, Long, wit h low Very long and low with high spine spine Dens epistrophei Projecting, narrow The same Spout-shaped, with with oval dorsal vestige of dorsal facet for floor of oval facet neural tunnel Anterior articular Convex, oblique The same Flattened, trans- facets verse Median ventral notch Not developed The same Sharp and large beneath dens epistrophei Postarticular cup Slightly flattened Much wider than Narrow, deeply con- at sides, higher high cave than wide Mid-ventral posterior Not presenIt Not present Prominent keel Mid-ventral posterior Prominent Prominent Absent notch Base of anterior bor- Notched The same Perforated by large der of neural arch foramen Transverse process On plane with mid- The same Raised above floor of dle of centrum neural tunnel Post-zygapophyses Flat Slightly concave Concavo-convex Opposite post-zyga- Widely separated Strongly approxi- Strongly approxi- pophyses mated mated Spine, median poste- Pronounced The same Abortive rior keel

The mid-cervicals are represented in our collection by two vertebrae of colossal size (Figs. 8, 9), which at first sight look more like the vertebrae of sauropod dinosaurs than like those of even the largest land mammals, especially since they bear on each side deep pleurocoeles or rounded cavities which, as Forster Cooper has shown, extend under the floor of 18 Bulletin Am&erican Museum of Natural History [Vol. LXXII

TABLE IV.-COMPARATIVE CHARACTERS OF THE FOURTH CERVICAL VERTEBRA Rhinoceros Baluchitherium Draught sumatrensis (Fig. 8) Horse General proportions Very short and Long, low and wide Very long, narrow high Anterior art icular Compressed, much Wider than high Higher than wide facet (ball) higher than wide, very convex Spine Well developed Absent Sessile, flattened Facets of anterior zy- Flat, moderately in- S ightl y convex, Very large, concave, gapophyses cined to horizon- spreading steeper tal Facets of posterior Flat, moderately in- The same; wide Elongated, steeply zygapophyses clined to horizon- and spreading inclined, less di- tal vergent Lateral processes Stout, blade-like, Elongate, blade- Extremely elongate, with three low like, with two slender, directed lateral processes processes shaxply upward posteriorly Lateral cavities on Not developed Highly developed Not developed body of centrum Posterior articular Higher than wide Wider than high Subequal facet (cup) Median posteriorven- Absent Absent Pronounced tral keel the neural tunnel and are separated in the midline only by a thin vertical septum. We are inclined to regard these openings not as due to a mysterious adaptation for the lightening of the bones but rather as a growth response to the general principle of fenestration, which in many parts of the vertebrate skeleton results in the strengthening and concen- tration of bony tissue along the zones of greatest stress and in the opening up of areas where stresses are minimized, as in the temporal fenestrae and pelvic plates of extinct reptiles and mammals. "Judging from the three known cervical vertebrae," writes Forster Cooper (1923, p. 38) "the length of the neck of Baluchitherium was of much the same proportion as that of the horse. The vertebrae are totally unlike those of the Rhinoceros, but show some approximation in general shape and proportions to those of the horse, the only Perissodac- tyle to which any likeness can be found. There are, however, notable differences which may be explained as adaptations consequent upon the great weight of the skull. The points of resemblances are, however, of 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 19

TABLE V.-CGMPARATIVE CHARACTERS OF THE SIXTH CERVICAL VERT EBRA Rhinoceros Baluchitherium Draught sumatrensis (Fig. 9) Horse General proportions Short Longer Very long Descending flange Simple Subdivided into an- Very elongate terior and posterior descending processes Planesofanteriorzyg- Facing chiefly in- Facing more for- Facing upward and apophysial facets ward and upward ward slightly forward Shape of anterior zyg- Rounded, plane Convex oval Slightly concave, apophysial facets subcircular, very large Spine Fairly large Absent Long, low Inclination of planes Downward and Slightly downward Sharply downward of posterior zyg- slightly outward and outward and outward apophysial facets Shape and size of pos- Rounded Widened Long ovate, very terior zygapophys- large ial facets Descending processes Flange-litke, con- Divided into long Skidlike, elongate, tinuous, deep antero - external shallow; not or and postero-in- barely subdivided ternal processes by deep embay- ment Transverse processes Directed backward, Directed outward, Directed outward not widely pro- projecting widely and backward jecting laterally Ball of centrum Higher than wide Wider than high Higher than wide Lateral cavities in Not developed Highly developed Not developed body of centrum Transverse and verti- Subequal Wider than high Subequal cal diameters of posterior articular facet (cup) Concavity of cup Pronounced Shallow Extremely deep Forward inclination Moderate Increased Extreme of cup to midline Extension of ball on Moderate Slight Extreme ventral surface of centrum great interest as examples of convergence in the shape of bones as the result of (presumably) similar stresses and strains in necks of equal proportional length." 20 Bulletin American Museum of Natural History [Vol. LXXII

After rather prolonged comparative studies of all the material now available we find that, while the atlas and axis to a certain extent suggest the corresponding bones of a horse, the remaining cervical vertebrae present many significant differendes, as follows: (1) In Baluchitherium the upwardly facing facets of the anterior zygapophyses are inclined much more forward than in the horse; and both the anterior and posterior zygapophysial facets tend to be small and ovate, whereas in the horse they are very large and subcircular. (2) Cervicals 4-7 of Baluchitherium are all relatively much wider and shorter than in the horse. (3) The ball-and-cup facets of the centra in Baluchitherium are wider than high, the reverse being true in the horse. (4) The convex facet on the ball of the centrum extends ventroposteriorly beneath the centrum to a much less extent in Baluchitherium than in the horse, in which this arrangement permits extreme raising of the neck. (5) The descending flange in cervical 6 is sharply subdivided into two processes in Baluchitherium, while in the horse the subdivision is at most incipient. (6) The marked relative shortness of the seventh and sixth cervicals in Baluchi- therium correspondingly limits the upward reach of the neck above the shoulders. From the facts set forth in our comparative tables we conclude that, contrary to what was shown in previous restorations, the mighty jointed drawbridge which was the neck and skull of Baluchitherium was normally directed downward-suspended by the great ligaments and muscles of the neck and occiput and pivoted at the proximal end on the well-buttress.ed joints of the thorax. The form of the zygapophysial and centrum facets and the relative shortness of the centra indicate wide lateral movements and relatively much shorter reach above the shoulders than in the horse. In brief Baluchitherium was essentially a feeder on relatively low bushes. The principal measurements of the cervicals 4, 6 and 7 are as follows: C4 C6 C7 (No. 26168) Centrum, length between transverse planes touching ball and cup 36. cm. 29.6 18 est. Centrum, width of ball 19.5 19 18 est. Centrum, height of ball 12.5 14.6 11 + Extreme width across transverse process 47.5 58 47 Extreme width across posterior zygapophyses 34 41.5 37 Extreme width across anterior zygapophyses 34 46 44.5 Extreme height, posterior cup to roof of neural arch 23.5 26 23 Length (a.p.), neural arch midline 24.5 17.5 11 est. Extreme length between transverse planes, anterior and posterior zygapophyses 42.3 19361 Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 21

Cervicals 4 and 6 represent our size Grade I, while C 7 is referred to Grade III.

TABLE VI.-COMPARATIVE CHARACTERS OF SEVENTH CERVICAL VERTEBRA Rhinoceros Baluchither,ium Draught sumatrensis (Fig. 10 Horse General proportions Moderate Short, broad, low Long Transverse processes Small, not extend- Massive, exteending Slender, upturned, ing below base of well below base well above base of centrum of centrum centrum Planes of anterior zyg- Inclined slightly Inclined forward Inclined upward and apophysial facets upward and in- and upward inward ward Shape of anterior zy- Subcircular, plane Oval, convex Very large, ovate gapophysial facets and plane Spine Large and high Presumably low Low, delicate Inclination of poste- Chiefly downward Downward and Downward and out- rior zygapophysial and outward backward ward facets Shape and size of pos- Transverse ovoid Small and subeircu- Very large, subquad- terior zygapophys- lar rate ial facets Ball of centrum Higher than wide Decidedly wider Higher than wide than high Extension of ball on Moderate Moderate Extreme ventral surface of centrum Transverse diameters Not much greater Much greater Subequal across neural arches compared w i t h transverse diameter of posterior articular facet (cup) The FIRST DORSAL (Fig. 11) vertebra seems to be of about the right size to fit with the seventh cervical and to belong to an animal of size Grade III. The centrum is 20.5 cm. long and 17.5 wide (ball) and the maximum spread across the lateral processes is 46.5. As compared with the first dorsal of a large draught horse, the side view of our specimen is moderately short, relatively less elongate; the planes of its anterior zygapophysial facets face upward and forward, whereas in the horse they face upward and inward; the facets themselves are of moderate size, convex oval, while in the horse they are very large Fig. 10. Seventh cervical vertebra of Baluchitherium grangeri. Size Grade III. X 1/5 natural size. A. Left lateral view. B. Upper surface. C. Anterior view. 22 Fig. 11. First dorsal vertebra of Baluchitherium grangeri. Size Grade III. X 1/5 natural size. A. Posterior view. B. Anterior view. (For lateral view see Fig. 12A.) 23 24 Bulletin American Museum of Natural History [Vol. LXXII and flat oval. The spine is very long, instead of being short and directed forward as in the horse. The excavation along the back of the spine is extremely developed, whereas in the horse it is absent. The area of the posterior zygapophysial facet as compared with that of the anterior zygapophysial facet is reduced, while in the horse it is greatly reduced. The median keel on the inferior surface of the centrum is absent, in contrast with the well-developed state in the horse. Finally the pro- longation of the ball upon the antero-ventral surface of the centrum is moderate, while in the horse it is marked. In all these features except the excavation along the back of the spine the first dorsal of Baluchi- therium is nearer to that of the recent Rhinoceros sumatrensis than to that of the horse. The SECOND DORSAL (Fig. 12) of a very small animal (not otherwise described as to size) resembles the first dorsal except that it is less extended transversely and has a more backwardly inclined neural spine; the central ball is more circular, the rib facets larger. The FOURTH DORSAL (Fig. 13) differs from the first especially in the extremely small size of the facets of the anterior zygapophysis, in the pentagonal contour of the anterior articular surface of the ball of the centrum, in the smaller transverse process and in the higher position of the dorsal surface of the transverse process as compared with the body of the centrum. This vertebra differs from that of Rhinoceros sumatrensis in the deep vertical excavation of the posterior border of the neural spine and in the consequent wider separation of the posterior zygapophysial facets. From that of the horse it differs in the lesser development of the metapophyses above the transverse process, in the relatively much smaller and lower capitular facets on either side of the concave posterior facet of the centrum. The horse also lacks the deep posterior excavation of the neural spine and has a sharp ventral keel on the centrum. The FIFTH(?) DORSAL iS apparently represented in Borissiak's Tab. IV, figs. 4a, b, c. Here the spine was set farther back than in D 3, the metapophyses were high and prominent and the facets for the capitula were higher up, flanking the top of the posterior facet of the centrum. The chief differences from Rhinoceros sumatrensis are the greater size and erect position of the metapophyses. As compared with the same bone of the horse, the anterior and posterior concave facets for the capitula are less prominent and less deeply concave, while the metapophyses are less extended anteroposteriorly. For the EIGHTH(?) DORSAL also we must rely on Borissiak's plate D

Fig. 12. First and second dorsal vertebrae of Baluchitherium grangeri. X 1/5 natural size. A. Left lateral view of first dorsal, larger animal. Size Grade III. (Cf. Fig. 11.) B. Left lateral view of second dorsal, very small animal. C. Anterior view, very small animal. D. Posterior view, very small animal.

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Fig. 13. Fourth dorsal vertebra of Baluchitherium grangeri. Size Grade IV. X 1/5 natural size. A. Posterior view. B. Anterior view. C. Left lateral view. 26 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 27

(Tab. IV, fig. 5), where a vertebra of this vicinity is shown with a relatively short spine and with the entire transyerse process directed outward and upward. The facets of the anterior zygapophyses are nearly horizontal. There is a rather close general correspondence with D 8 of Rhinoceros sumatrensis. In the horse the spine is relatively much longer and the rib facets very large and cup-like.

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Fig. 14. Twelfth dorsal vertebra of Baluchitherium grangeri. Size Grade IV. X 1/5 natural size. A. Left lateral view. B. Posterior view. C. Anterior view. 28 Bulletin American Museum of Natural History [Vol. LXXII

Fig. 15. Thirteenth dorsal vertebra of Baluchitherium grangeri. Size Grade IV. X 1/6 natural size. A. Left lateral view. B. Posterior view. C. Anterior view.

In the TENTH(?) DORSAL (Borissiak, Tab. IV, fig. 6a, b, c) the large spine is somewhat sigmoid in side view. There is hardly any lateral projection of the transverse processes and the small rib facets lie lateral to the pedicles of the neural arch, although extending down on the uppermost levels of the posterior articular face of the centrum. The latter is deep vertically and narrow transversely. D 10 of Rhinoceros sumatrensis has a relatively shorter spine, much wider transverse pro- 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 29 cesses, higher metapophyses. In the horse, D 10 has much larger facets for the capitulum and relatively wider centra. The TWELFTH and THIRTEENTH DORSALS (Figs. 14, 15) are represented in our collection by two well-preserved vertebrae (No. 26387)

B Fig. 16. First lumbar vertebra of Baluchitherium grangeri. Size Grade IV. X 1/5 natural size. A. Left lateral view. B. Posterior view. C. Anterior view. A7

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Fig. 17. Second lumbar vertebra of Baluchitherium grangeri. Size Grade IV. X 1/5 natural size. A. Left lateral view. B. Posterior view. C. Anterior view.

30 Fig. 18. Third lumbar vertebra of Baluchitherium grangeri. Size Grade IV. X '/I natural size. A. Left lateral view. B. Anterior view. C. Posterior view. 31 32 Bulletin American Museum of Natural History [Vol. LXXII associated with lumbar and sacral vertebrae and limb bones. This animal belongs to our size Grade IV. In the thirteenth dorsal the top of the neural spine is 34 cm. above the level of the base of the centrum; length of tl4e centrum, 13 cm., maximum width of same, 16 cm. Both D 12 and 13 have much higher spines than the corresponding vertebrae of Rhinoceros sumatrensis but are in this respect not so different from the white rhinoceros. The metapophyses, however, are more conspicuous. The postzygapophysial facets are beginning to be subdivided by a median eminence into inner and outer planes corresponding to concave facets on the anterior zygapophyses, whereas in both Rhi- noceros sumatrensis and Ceratotherium simum they are flat. The neural spines retain shallow posterior excavations throughout their length as in Ceratotherium simum, whereas in Rhinoceros sumatrensis the very short spines are keeled posteriorly. The lumbars, first, second and third, are well-preserved bones (Figs. 16, 17, 18) of the same series of relatively small-sized animals. The principal measurements of these lumbars are as follows: Li L2 L3 Height to top of spine 37 cm. 41 cm. 42 cm. Length, centrum (side) 13 14.5 14.5 Max. width, centrum (post.) 16.5 15.5 17.5 The lumbars are characterized by their long, large spines, erect metapophyses, keeled centra and small anterior zygapophysial facets on LI, L2, and deeply concave facets on L3. In Rhinoceros sumatrensis the spines are short and antero-posteriorly longer, the metapophyses vestigial and the zygapophysial facets relatively larger. These conditions are accentuated in Ceratotherium simum. The transverse processes of the first lumbar, although short and truncate distally, are true ribs, articulating by a vestigial capitulum with the antero-superior corner of the centrum and by a much widened tubercu- lum with an oval facet on the lateral process. In our Rhinoceros sumatrensis the first lumbar rib is detached on the right side, leaving a large articular surface on the transverse process. Traces of this separateness of the rib are visible on Li of a large white rhinoceros but not on our draught horse. In the second and third lumbars the long upwardly arched transverse processes show no trace of having arisen as ribs. The fourth lumbar is missing. The fifth (?), as represented in Borissiak's Tab. V, Figs. 5a, b, c, has a large low transverse process and widely oval anterior 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 33

and posterior articular facets of the centrum. The spinie is fairly high, the metapophyses prominent and forwardly directed. There is a large oval facet for the sacral rib on the back of the transverse process. In Trigonias the last lumbar vertebra was free from the sacrum but in Rhinoceros sumatrensis it has become enlarged and integrated with the sacrum, forming indeed the largest and widest member of the sacral series. The same seems to have been true in Baluchitherium. The SACRUM (Fig. 19) is incompletely preserved in the same series (No. 26387). Of the five spines preserved the first and highest seems to represent the last lumbar (or lumbo-sacral), which is also represented by the very large flaring transverse process of the left side. The last spiine belongs to the first coccygeal or sacro-caudal vertebra. The spines are

L5 Si 1!32 j 3 34

Fig. 19. Fifth lumbar vertebra and sacrum of Baluchitheriumn grangeri. Size Grade IV. X 1/5 natural size. Left later al view. 34 BuUletin American Museum of Natural History [Vol. LXXlI

expanded and very rugose on top, not compressed as in Rhinoceros sumatrensis. An antero-posterior series of four pairs of large nerve exits on the under side lie between the transverse processes of the three true sacral vertebrae, the centra of which are flat beneath. There is a very large lateral articular surface for the ilium on the transverse process of the great lumbo-sacral, together with a much smaller iliac facet on the transverse process of the first true sacral and the minute iliac facets of the second and third sacrals. The lumbar and sacral regions of Baluchi- therium are widely different from those of a horse. In the draught horse there are six lumbars, of which the last three have the flattened form of lumbo-sacrals. Possibly the sixth represents an appropriated first sacral. The CAUDAL VERTEBRAE are not preserved but to judge from the relatively large size of the sacro-coccygeal vertebrae, the tail was not materially different except in absolute size from that of Trigonias. To sum up, the cervical vertebrae of Baluchitherium, in spite of their specializations for huge size, including greater relative width, loss of spines, development of pleurocoeles, downward and forward turning of anterior zygapophysial facets, etc., exhibit no peculiarities which are not apparently derivable from the far more primitive condition preserved in Trigonias and Allacerops ("Epiaceratherium") turgaicum. Thus to derive the atlas of Baluchitherium from that of Trigonias one would have to increase the length of the transverse processes faster than their width. At the same time the transverse diameters of the cotyli for the occipital condyles would have to be accelerated. The axis of Ba- luchitherium has evidently been derived from a form like that of Alla- cerops ("Epiaceratherium") merely by a differential lengthening of the bone as a whole. The fourth cervical of Baluchitherium is relatively longer, lower and much wider than that of Trigonias, the ball of the centrum is widened and the cup still more so; the spine, well developed in the primitive Trigonias, has disappeared and the anterior zygapophysial facets have been turned more forward. The transverse blade is more horizontal in position and tends to be subdivided into anterior and posterior divisions. Deep lateral cavities have developed on the centrum. The sixth cervical of Baluchitherium is relatively longer and wider than that of Trigonias and has lost its spine; its downwardly directed flange is sharply subdivided into anterior and posterior wings, its centrum is deeply excavated and the anterior zygapophysial facets face more forward, the posterior ones more backward, this favoring lateral 1936] Granger and Gregory, Baluchitherium from -the Oligocene of Mongolia 35 rather than vertical movements. The seventh cervical as compared with that of Trigonias is short, broad and low, with low spine; its very massive transverse process extends well below the base of the centrum. The centrum is extremely wide and the ball is somewhat more produced ( 'I'lit,,It','

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lu Fig. 20. Ribs of Baluchitherium grangeri. (Small animaL1, size Grade IV, No. 26387.) X 1/lo natural size. A. Sixth dorsal rib, left. Anterior view. B. Eighth dorsal rib, left. Anterior view. C. Eighth dorsal rib, left. Lateral view. D. Tenth dorsal rib, left. Lateral view. 36 Bulletin American Museum of Natural History [Vol. LXXII on the ventral surface than in Trigonias. The planes of the postzygapophysial facets are directed more downward and backward and not so much outward. The pedicles or pillars of the neural arches are massive and greatly widened. Thus the neck of Baluchitherium was capable of freer lateral movements than that in the Trigonias-like ancestor. The dorsal, lumbar and sacral vertebrae of Baluchitherium are like-

Fig. 21. Ribs of Baluchitherium grangeri (continued). (Small animal, size Grade IV, No. 26169.) X 1/lo natural size. A. Tenth doisal rib, right. Lateral view. B. Tenth dorsal rib, right. Anterior view. C. Twelfthdorsalrib, right. Anterior view. D. Twelfth dorsal rib, right. Lateral view. 1936] Granger and Gregory, Baluchitherium from the Olgiocene of Mongolia -37 wise all readily derivable from the corresponding parts of primitive subcursorial rhinoceroses and differ very widely from those of the horse. Among their more conspicuous features are the posterior excavations of the long neural spines of D 1-D4,the emphasis of the erect metapophyses, which are depressed or relatively feeble in recent rhinoceroses and the

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-, I. 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 41 relatively small size of the concave facets for the capitula of the ribs, perhaps indicating less extreme movements of the ribs in respiration. RIBS Several ribs (Figs. 20, 21) were found associated with two of the smaller animals (Grade IV), while others (Fig. 22) were found with No. 26166, one of the larger animals (Grade II). Careful comparisons with the ribs of the Indian rhinoceros reveal surprisingly close resemblances, both in general form and in details, so that we were able to determine approximately the serial number of the individual ribs. Dorsal ribs 6, 8, 10, 12, 14 and the first lumbar rib, as well as the various ribs figured by Borissiak, thus prove that the shape of the thorax as a whole was close to that of the Indian rhinoceros. However, on account of the very long radius and femur of Baluchitherium, its thorax would look decidedly smaller in proportion to the height of the whole animal.

LIMBS AND FEET Comparison of Baluchitherium limb bones with those of the highly specialized modern rhinoceroses establishes the fact that while Baluchi- therium has retained surprisingly many characters of the ancestral cursorial rhinoceros, it has also acquired a few " graviportal" characters with its great weight. Of these perhaps the most outstanding are: the lengthening of the femur, the shortening of the tibia, the widening of the astragalus and of the ungual phalanges of the middle digits. On the other hand, the usual shortening of the middle digits is conspicuously absent. In fact Baluchitherium alone among gigantic perissodactyls has widened and elongated the middle metapodials and phalanges, especially in the manus. In the vertebral column the chief graviportal characters are the widening of the neck vertebrae and the deep lateral excavation of the centra. Our comparative measurements and studies of the skeletons of Rhi- noceros sumatrensis, Rhinoceros unicornis and Ceratotherium simum indicate that in the modern rhinoceroses the legs have become relatively short and broad, while the body has increased greatly in bulk; the occiput has broadened as the nasal horns have developed. In Baluchi- therium, on the other hand, the limbs rapidly lengthened while the head became long and low and remained hornless. The SCAPULA iS best known from Borissiak's specimen (text fig. 2, p. 56), which reveals a rather slender straight spine, a transversely thick, 42 Bulletin American Museum of Natural History [Vol. LXXII expanded glenoid articular surface and a massive transversely thick process for the tendon of the biceps. The antero-posterior width of the bone is not known but Granger records a field measurement of 80 cm. for the "length" (height) of the scapula in a rather small individual (No. 26387) associated with vertebrae and limb bones. This length would be only 70 per cent of the length of the radius of the same individual, a surprisingly low figure. The shape of the scapula differs very widely in the various genera of fossil and recent rhinoceroses but in those that have very long dorsal spines, as in the white rhinoceros, the scapula, while massive, is greatly elongated vertically. Nevertheless the positive

Fig. 23. Left humerus of Baluchitherium grangeri. (Large animal, size Grade II, No. 26166.) X 1/lo natural size. A. Anterior view. B. Lateral view. 1936] Granger and Gregory, Baluchitheriumfrom the Oligocene of Mongolia 43 measurement recorded by Granger indicates a relatively short scapula; accordingly we have assigned to this bone in the restoration a contour of generalized rhinocerotic type, somewhat widened antero-posteriorly. Yet the scapula of Indricotherium as figured by Borissiak was remarkably thick transversely at the lower end; thus the bone doubtless played its full share both in supporting the huge body through the subscapularis

Fig. 24. Right radius of Baluchitherium grangeri. (Large animal, size Grade II, No. 26166.) X 1/10 natural size. A. Anterior view. B. Lateral view. 44 Bulletin American Museunm of Natural History [Vol. LXXII and serratus muscles and in forming part of the fulcrum for the neck anid skull through the superficial neck muscles. The HUMERUS (Fig. 23) is of enormous strength and thickness. Its length, 98.5 cm., in our largest specimen, which is known to belong to an animal of the second size rank (see p. 65), somewhat exceeds the dinien- sions recorded by Borissiak (pp. 58, 59). The immense, gently convex head faces chiefly upward and less backward than in more typical rhinoceroses. This implies only a gentle inclination of the scapula to the humerus in the standing pose. The prominent deltopectoral crest, tri- angular in cross-section, stiffens the bone on the anterior face. The

F'ig. 25. Part of right radius and ulna of Baluchitherium graeriger. (Size Grade IV, No. 26169.) X 1/io natural size. A. Lateral view. B. Anterior view. great tuberosity was almost sessile on top of the deltopectoral crest. The lesser tuberosity is massive but low, implying a powerful subseapu- laris muscle. Although the bicipital groove was not pronounced, the biceps itself must have beeni of enormous size, as we know from the great thiclkess of the bicipital process on the scapula and the promillence of the bicipital eminence on the radius. The humerus is much shorter than the ra(lius, possibly because Balu- 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 45 chitherium is a direct descendant of cursorial rhinoceroses, with relatively short humeri and long radii and ulnae. As shown by articulating the humerus with the radius, the angle between the two bones in the standing pose is distinctly greater than in Borissiak's restoration. The RADIUS (Fig. 24) is a very long bone flattened in front but of massive cross-section, the length reaching 122 ecn. in our largest specimen, as compared with 94.5 cm. in Parelephas imperator. The distal end is subrectangular in section, which gives great resistance to bending stresses. It is immediately derivable from the cursorial radius of "Epiaceratherium" (Borissiak, Tal). II, fig. 3).

Fig. 26. Third left metacarpal of Baluchitherium g rang e r i. (Large animal, size Grade II, No. 26166.) dt R X natural size. 1,/ 1/ioA. Anterior view. B. Lateral view.

The ULNA (Figs. 25, 38) is considerably longer than the radius, and has a prominent, well-rounded but not gigantic olecranon and relatively slender shaft. The MANUS, having been fully described by both Cooper and Borissiak, calls only for the note that one of our third metacarpals (Fig. 45 F), attaining an estimated length of 63.5 cm., exceeds any other hitherto recorded. The second largest middle metacarpal (Fig. 26) is fortunately associated with the complete radius and humerus described above. A relatively small and immature manus (No. 21618, Fig. 27) associated with a pes, emphasizes the tendenicy toward mono- dactylism already noted by Cooper and Borissiak. The PELVIS iS known chiefly from the specimen figured by Borissiak (p. 91), which according to him was one metre in length. As restored Fig. 27. Right manus of Baluchitherium gransgeri. (Small animal, size Grade IV, No. 21618.) X 1/5 natural size. 46 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 47 the dorsal crest of the ilium was not expanded in the usual way among gigantic animals. Since, however, Borissiak's figure shows that the blade of the ilium was badly broken in small pieces, it raises a doubt whether the dorsal border is correctly restored. Moreover, a pelvis of a very large ungulate provisionally referred to Baluchitherium was photographed by Forster Cooper (1923, p. 371, Fig. 2) in the field.

Fig. 28. Right femur of Baluchitherium grangeri. (Small animal, size Grade IV, No. 26169.) X '/lo natural size. A. Anterior view. B. Lateral view. 48 Bulletin American Museum of Natural History [Vol. LXXII

This specimen has the widely expanded dorsal crest of the ilium as in other large rhinoceroses. Accordingly in our restoration (Fig. 47) we have given the pelvis an expanded dorsal crest. Nor can we agree with Borissiak in giving the pelvis suich a nearly vertical position in the restoration of the skeleton, since the characters of the sacrum indicate that the inclination of the ilium to the backbone was not essentially different from that which is found in other rhinoceroses.

Fig. 29. Left tibia and fibula of Baluchitheriumb grangeri. (Small animal, size Grade IV, No. 26169.) X '/lo natural size. A. Anterior view. B. Lateral view.

Granger records a field measurement of a pelvis of 112 cm. length associated with a radiu.s of the same. longth and a femur of 128.5 cm. length. If we multiply the length of Granger's pelvis by 1.4 (the factor used to enlarge size IV to size I) it makes an estimated length of 156.8 cm. In our restoration of size I we have somewhat exceeded this esti- mate, but such an immense hind limb of distinctly rhinocerotic type would seem to require a large ilium with a spreading crest, especially Fig. 30. Right pes of Baluchitherium grangeri. (Small animal, size Grade IV, No. 21618.) X 1/5 natural size. 49 Bulletin American Museum of Natural History [Vol. LXXII since the ribs and vertebrae indicate a thorax and lumbar region closely approaching those of Rhinoceros indicus. The FEMUR (Figs. 28, 41) is represented by several good specimens in our collection, which show that the third trochanter is far less reduced than one would suppose from Borissiak's text figure 10, page 93. The great trochanter lies well below the level of the head. We cannot agree with Borissiak in making the shaft of the femur almost vertical in the restoration, as a direct fitting of associated bones shows that the knee was bent even in the standing pose. The TIBIA (Fig. 29), remarkably short and wide, is beautifully preserved in one specimen, associated with femur and fibula. Both field and laboratory measurements indicate that the tibia is about 66 per cent

4 BCt D Fig. 31. Middle metatarsals of Baluchitherium grangeri. (Small animals, size Grade IV.) X 1/lo. A. Right, anterior view, No. 26387. C. Left, anterior view, No. 26169. B. Right, inner view, No. 26387. D. Left, lateral outer view, No. 26169. of the length of the femur and is thus rather graviportal in proportions. The rectangular cross-section of its lower end is well adapted to resist the great bending stresses to which this bone was subjected. The FIBULA (Fig. 29) is well preserved in two of the smaller specimens. It has a slender shaft and expanded distal end. The PES (Fig. 30), having been fully described by Borissiak, requires little comment except that even in our somewhat immature specimen the tendency toward monodactyly is pronounced. The middle metatarsal (Fig. 31), preserved in two of the smaller specimens, is slightly shorter and relatively stouter than the middle metacarpal. In spite of the graviportal adaptations of the astragalus (Fig. 32), the cuboid facet of that II~~~~~~~~~~~~I

Fig. 32. Tarsal bones of Baluchitherium grangeri. A. Left calcaneum, anterior view, No. 26387. X 1/'. E. Left astragalus, anteriorview, No. 26973. X 1/s. B. Left astragalus, anterior view, No. 26387. X 1/5. F. Left astragalus, posterior view, No. 26973. X 1/4. C. Left astragalus, posterior view, No. 26387. X l/4. G. Left astragalus, posterior view, No. 5209. X V/4. D. Left calcaneum, anterior view, No. 26973. X 1/6. H. Left astragalus, anterior view, No. 5209. X 1A. 51 52 Bulletin American Museum of Natural History [Vol. LXXII bone is but little increased, in contrast to the conditions observed in the graviportal titanotheres. The associated FORE AND HIND FEET (Figs. 27, 30) of a single articu- lated skeleton, the rest of which had been unfortunately eroded away, give us some valuable correlations of measurements between the manus and pes, and since the limbs of the animal were preserved nearly in situ, they also afford an approximation to the distance between the fore and hind limbs. On the right side the distance between centers of the fore and hind feet was 229 cm., on the left, 254 cm., giving an average distance between fore and hind feet of 241 cm. As the legs were directed outward in the death pose, the distance between the two hind feet (152 cm.) is of little value. Other measurements of this small and somewhat immature specimen are given in Table VIII.

TABLE VIII.-MEASUREMENTS OF AssocIATED FORE AND HIND FEET (No. 21618) Radius, transverse distal end 25.5 Mtc. III, r. length 40-42 Carpus, transverse proximal end 23.5 Mtc. III, width prox. 13.5 Carpus, mid. height 16 Mtc. III, max. width (list. 14 Carpus, lateral height Width dist. phal. Mtc. III 14 Carpus, height cun. + une. 17.5 Width prox. " " " 12.5 Distance between centers, right Mtc. III, l. length 40-42 fore foot and right hind foot 229 Mtc. III width, prox. Distanice between centers, left Mtc. III max. width dist. fore foot and left hind foot 254 Mtc. II, length 36.5 Aveiage distance 241 Mtc. II, width Width between fore feet 135 Mtc. IV, length 35 Width between hind feet Tibia, width dist. end 19 (sprawled) 152 Tarsus, height, front (tibia to Average distance 143 prox. Mts. III) 16

No. 21618 No. 26387 No. 21619 Mts. III, r. length 39 40 Mts. III, width, prox. 12 14 Mts. III, width, distal 12 15 Max. width, Phal. III 14 Width, pr-ox. 14 Mts. II, length 33 Mts. IV, length 32 Calcaneum, length 26.5 28 Astragalus, width acr'oss condyle 16.5 17.3

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the limb bones in Baluchitherium, we may note the following striking results recorded in Table IX: (1) Allacerops ("Epiaceratherium") in most of its limb segment ratios tends either to be intermediate between Trigonias and Baluchitherium or to agree with either one or the other. (2) In Baluchitherium the scapula is relatively very short, as compared with the humerus, whereas in the cursorial Equus the scapula is long. (3) In Baluchitherium the radius is very long, both relatively and absolutely. (4) The middle metacarpal of Baluchitherium has shared to some extent in the elongation of the lower half of the fore limb. (5) The pelvis is relatively short and, conversely, the femur very long. (6) The tibia and third metatarsal are relatively shorter than the radius and the third metacarpal, respectively. (7) Baluchitherium by this showing belongs among neither the cursorial nor the graviportal types of Osborn, except in certain features (e.g., graviportal ratio of tibia to femur). It is, on the other hand, a gigantic, long-limbed rhinoceros, the ratios of its limb segments being most easily derivable from those of primitive subcursorial ancestors allied with Allacerops and Trigonias. RELATIONSHIPS As to generic relationships, our material indicates that both Baluchi- therium and Indricotherium are close to or even synonymous with Para-

B Fig. 33. Left maxilla and premaxilla with teeth. A. Epiaceratherium turgaicum. After Borissiak. X1/6. B. Baluchitherium grangeri. After Osborn. X I/12. 1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 55 ceratherium, although possibly representing slightly different species. In the first place, the type upper molars of Paraceratherium bugtiense Pilgrim exhibit no conspicuous differences from those of Baluchitherium osborni; secondly, the lower jaw referred to Paraceratherium bugtiense by Forster Cooper seems to us to be indistinguishable in generic characters from one of our jaws (No. 26166) that is associated with humerus, radius, ulna and metacarpal III of the general size and characters of Borissiak's Indricotherium; thirdly, the cast of the skull referred by

/1 Fig. 34. Right upper cheek teeth. A. 107.8 Epiaceratherium turgaicum. After Borissiak. X 200- B. Indricotherium asiaticum. After 4 Borissiak. X 200

Forster Cooper to Paraceratherium reveals essential similarities at all points to our large skulls of Baluchitherium grangeri Osborn. Unfor- tunately, however, this substantial identity is obscured by the fact that the Paraceratherium skull is crushed vertically and its rostrum is broken off. Fourthly, the peculiar lower front teeth of Paraceratherium are matched precisely in Baluchitherium osborni and in Borissiak's Indrico- therium. Fifthly, we have numerous fully adult limb bones, astragali 56 Bulletin American Museum of Natural History [Vol. LXXII and metapodials that collectively comprise a closely graded series (Figs. 44, 45) from the small Paraceratherium through Baluchitherium osborni to B. grangeri and finally to a super-Indricotherium. On the other hand, Borissiak has pointed out that in Forster Cooper's Paraceratherium the protoloph of the fourth upper premolar is higher than in Indricotherium, the whole crown is slightly more hypsodont and the cingulum better developed; also the incipient " crochets" of the upper molars are a little

D Fig. 35. Atlas and axis. X 1/6. A, C. Epiaceratherium. After Borissiak. B, D. Baluchitherium. C U j ,1~ {r

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Fig. 36. Dorsal vertebrae. X i/. A. First (?) dorsal of Epiaceratherium. After Borissiak. B. First dorsal of Baluchitherium. C. Fourth (?) dorsal of Epiaceratherium. After Borissiak. D. Fourth dorsal of Baluchitherium. E. Fourteenth (?) dorsal of Epiaceratherium. After Borissiak. F. Thirteenth dorsal of Baluchitherium. 57 58 Bulletin American Museum of Natural History [Vol. LXXII more pronounced; assuredly, however, the evidence assembled in Fig. 2 above is not favorable to the idea that Paraceratherium, Baluchitherium and Indricotherium are distinct genera, although there are minor and perhaps specific differences, especially in the second upper premolars. Moreover, our experience with the remarkable variability of Oligocene

Fig. 37. Left scapula and left humerus. X 1/5. A. Scapula of Indricotherium. After Borissiak but outline restored and enlarged to size Grade I. B. Scapula of Epiaceratherium. After Borissiak. C. Left humerus of Baluchitherium. Anterior view. D. Left humerus of Epiaceratherium. Anterior view. After Borissiak. "N~ ~ /

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rhinoceros premolars would make us hesitate to allow a superspecific value to such differences.1 We accordingly continue to use the name Baluchitherium grangeri Osborn for the Mongolian material. As noted by Matthew, the type of Baluchitherium mongoliense Os- born (1924) from the Loh formation, ? Lower Miocene, represents a far more advanced type of rhinocerotid, with submolariform p4 and com- ' Cf. Gregory and Cook, 'New Material for the Study of Evolution: A Series of Primitive Rhi- noceros Skulls (Trigonias) from the Lower Oligocene of Colorado.' Proc. Colorado Mus. Nat. Hist., VIII, No. 1, February, 1928. B

Fig. 42. Right pes. X1/. A. Baluchitherium grangeri (size Grade IV). B. Epiaceratherium. After Borissiak. 63 64 Bulletin Anmerican Museum of Natural History [Vol. LXXII

plex hyposodont molars. It appears to belong to a very different sub- family of rhinoceroses from Baluchitherium. As to the derivation of Baluchitherium from more primitive hornless rhinoceroses, Borissiak in his memoir on "Epiaceratherium turgaicum" (a small light-limbed rhinoceros contemporary with Baluchitherium) has remarked (1918, p. 82) that Epiaceratherium is morphologically related to the primitive Aceratherium and that all the differences between it and Indricotherium, although so marked, even apart from size, in the form of

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Fig. 43. Right astragalus and left calcaneum. X 1/'. A. Baluchitherium grangeri. B. Epiaceratherium. After Borissiak. C. Baluchitherium grangeri. D. Epiaceratherium. After Borissiak, 1936] Cranger and Oregory, B3aluchitherium from the Oligocene of Mongolia 65 the bones and of their articular surfaces, ought to be attributed to the unique specialization of Indricotherium. After comparing Borissiak's excellent figures of the teeth and bones of "Epiaceratherium" (= Alla- cerops') with those of Baluchitherium and Indricotherium, we would go even farther than he does and say that in regard to its vertebrae and limb bones this small subcursorial rhinoceros (Figs. 33-43) makes an ideal structural ancestor for its huge contemporary. The accompanying table (Table IX) of limb measurements and limb ratios suggests that "Epiaceratherium" (= Allacerops) affords an intermediate structural stage connecting Baluchitherium with more primitive ancestors typified by Eotrigonias Wood. RESTORATION The principal parts of the skeleton, except the sternebrae, are represented in the collection. As noted above, there is an enormous range in the size of the adults, the smallest middle metacarpal of the manus (Figs. 44, 45) measuring 405 mm., the longest, 635 mm. in length. We have grouped the material used in the restoration under four descending grades of size. The middle metacarpal of Grade I is about 1.4 times as long as that of Grade IV, 1.3 times that of Grade III and 1.2 times that of Grade II. Consequently these factors, along with others, have been used (Fig. 46) in enlarging bones of the smaller grades to the probable size of Grade I, which is represented by several gigantic cervical vertebrae and by the third metacarpal. Grade II includes the huge skull, a lower jaw associated with a humerus, radius and middle metacarpal, and several ribs (Amer. Mus. No. 26166). Grade III is represented by the smaller occiput, atlas, axis. Grade IV includes associated manus and pes and various associated vertebrae, ribs, femur, tibia and middle metatarsal. Those who are familiar with the difficulties in securing consistent consecutive measurements from large fossil bones that are more or less imperfect or distorted will not expect our work to be free from errors. After repeated revisions our restoration (Fig. 47) represents an animal of the largest grade, seventeen feet, three inches in height at the shoulder (top of spine at first dorsal vertebra). The height at the shoulder as thus estimated exceeds that of the tallest hitherto known land mammal.2 Estimated length (Grade I) in standing pose, from tip of 1 Dal Piaz (1930) has shown that "Epiaceratherium turgaicum" Borissiak is not congeneric with E. bolcense, the genotype, and H. E. Wood, 2d. (1932), has made Borissiak's species the genotype of Allacerops. 2 With the possible exception of the Upper Miocene Dinotherium giganteum, to which M. Boule (1935, p. 610) assigns a possible maximum height of 5 meters. B C U D

Fig. 44. Right third metacarpals. X 1/6. A. ? Paraceratherium sp., No. 26190 (extremely small). B. Baluchitherium grangeri, No. 26389. C. Baluchitherium grangeri. No. 21618 (size Grade IV). D. Baluchitherium grangeri, No. 26166 (size Grade II). 66 E F Fig. 45. Right third metacarpals (continued). X 1/5. E. Indricotherium asiaticum, No. 26973. Cast; original in Moscow. (Size between Grades I and II.) F. Baluchitherium grangeri, No. 26175. (Size Grade I.) 67 b0

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1936] Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 60 premaxilla to ischial tuberosity, about twenty-seven feet. Size Grade II animals would be about fourteen feet at the shoulder, size Grade IV about twelve feet. The skull is relatively small; the axis is compara- tively long and low but cervicals 4-7 are relatively very broad and low as compared with those of recent rhinoceroses. On the whole, our restoration makes Baluchitherium not unlike one of the primitive hornless Oligocene rhinoceroses, except for its titanic size and relatively long radius, long femur, small head, elongate axis and wider mid-cervicals. This is also to be inferred from the detailed characters of the individual bones. We can find no evidence for the okapi- like restorations by earlier authors. LIST OF SPECIMENS OF BALUCHITHERIUM AND RELATED FORMS FROM MONGOLIA, 1922-1930 BARON SOG AND HOULDJIN BEDS 26165 Posterior half of skull-no teeth. Urtyn Obo, 1928. 26167 Posterior half of skull-no teeth. Urtyn Obo, 1928. 26172 Lower jaws, P2-M3 right and left. Nom Kong Obo (Holy Mesa), 1928. 26171 Lower jaw fragment with four cheek teeth. Urtyn Obo, 1928. 26166 Humerus, left; radius, right; metacarpal III, left; two ribs (D6, D7 ?); lower jaw, right ramus complete and P2-4 left. Size Grade II. Urtyn Obo, 1928. 26169 Distal portion of humerus, left; femur, right; distal half of left femur; tibia and fibula, left; proximal ends of right ulna and radius and left radius and distal ends of both ulnae. Metatarsal III, left; 3d dorsal vertebra and two ribs (D12, 14, right). Size Grade IV. Nom Kong Obo (Holy Mesa), 1928. 26387 Sacrum, three lumbar and two dorsal vertebrae; left astragalus; astragalus, calcaneum, cuboid, ectocunei- form and metatarsal III of right pes; two proximal and two ungual phalanges of median digit; two ribs (D8 left, D10 left); fibula; patella. Also measured in situ but not col- lected, a femur, scapula, radius and pelvis. Size Grade IV. 25 miles S. W. of Iren Dabasu, 1930. 26168 Two cervical vertebrae-C3 and C6 (association uncertain). Nom Kong Obo (Holy Mesa), 1928. 26173 Anterior dorsal vertebra. Urtyn Obo, 1928. 26390 Axis vertebra. 25 miles S. W. of Iren Dabasu, 1930. 26392 Seventh cervical vertebra. Size Grade III. 25 miles S. W. of Iren Dabasu, 1930. 26393 Femur, left. 25 miles S. W. of Iren Dabasu, 1930. 26175 Metacarpal III, right (maximum size). Size Grade I. Urtyn Obo, 1928. 70 19361 Granger and Gregory, Baluchitherium from the Oligocene of Mongolia it

26388 Metacarpal III, left. 25 miles S. W. of Iren Dabasu, 1930. 26389 Metatarsal III, left. 25 miles S. W. of Iren Dabasu, 1930. 26179 Two calcanea, right and left (unasso- ciated). Urtyn Obo, 1928. 26174 Calcaneum, left. Urtyn Obo, 1928. 26189 Patella, astragalus and carpal (no association). Nom Kong Obo (Holy Mesa), 1928. 18651 Calcaneum and a few fragments of other bones. Iren Dabasu, 1922. 26190 Atlas vertebra; two calcanea; one metatarsal III, one metacarpal III; three phalanges? (Small size-possibly not Baluchitherium.) Jhama Obo, 1928. HSANDA GOL BEDS (Tsagan Nor Region) 18650 Skull and fragmentary lower jaws. Type of Baluchitherium grangeri Osborn. Size Grade II. 1922. 21618 Fore and hind feet complete except left hind foot which lacks tarsus. Size Grade IV. 1925. 18652 Distal end of humerus, left (found near skull No. 18650). 1922. 20446 Proximal ends of ulna and radius. 1922. 21619 Femur, left; dorsal vertebra (no association). Size Grade IV. 1925. 21749 Proximal end of left metatarsus, and one indet. foot bone. 1925. LITERATURE CITED 3ELIAJEVA, E. 1927. 'Catalogue of the Geological Museum. Vertebrata. Mam- malia. Rhinocerotidae. Genus Indricotherium Boriss.' Travaux du Mus6e Geol. pres l'Acad6mie des Sciences de 1'URSS, IV, pp. 241-272. 1929 (?1930). 'Catalogue of the Geological Museum. Vertebrata. Mam- malia. Rhinocerotidae. Genus Epiaceratherium Abel.' Tra- vaux du Mus6e G6ol. pr6s l'Academie des Sciences de l'URSS, VI, pp. 179-193. BORISSIAK, A. 1915a. 'Epiaceratherium turgaicum, n. sp.' Bull. de l'Acad. Imp. des Sciences, pp. 781-787. 1915b. 'Indricotherium, n. gen.' Gcological Messenger, No.3, pp. 131-134. 1916. 'The dental apparatus of Indricotherium.' Bull. de l'Acad. Imp. des Sciences, X (6), pp. 343-348. 1918. 'Sur l'ost6ologie de 1'Epiaceratherium turgaicum, nov. sp.' Soc.. Pal6ont. d. Russie. M6m. I, 82 pp., 3 pls. 1921a. 'On the lower jaw of a small rhinoceros from the Indricotherium beds of Turgai region.' Monogr. Russian Paleont. Soc., 1918, pp. 39-44. 1921b. 'On the lower jaw of a small rhinoceros from the Indricotherium beds of Turgai region.' Bull. d. l'Acad. d. Sci. d. Russie, pp. 397- 402. 1923a. 'Sur un nouveau representant des Rhinoc6ros gigantesques de l'Oligoc6ne d'Asie, Indricotherium asiaticum, n.g., n.sp.' MWm. d. 1. Soc. d. France: Pal6ontologie. M6m. No. 59, XXV, pp. 1-15, PIs. ix-xIII. 1923b. 'Indricotherium n.g. (cem. Rhinocerotidae).' M6m. d. l'Acad. d. Sci. d. Russie, (8) XXXV, No. 6, 128 pp., 11 pls. 1923c. 'A Reconstruction of Indricotherium.' Bull. d. l'Acad. d. Sci. d. Russie, pp. 111-114. 1924a. 'New additional material. Indricotheriinae Boriss. (Baluchi- theriinae Osb.).' Bull. d. l'Acad. d. Sci. d. Russie, (6) XVIII, pp. 127-150. 1924b. 'Indricotherium and Baluchitherium.' Comptes Rendus de l'Acad. d. Russie, pp. 148-149. 1924c. 'Uber die Unterfamilie Indricotheriinae Boriss. = Baluchitheriinae Osb.' Centralblatt f. Min., etc., No. 18, pp. 571-575. 1927a. 'On the Paraceratherium.' Comptes Rendus d. l'Acad. d. Sci. d. l'URSS, pp. 1, 2. 1927b. 'On the Brachypotherium from the Jilancik-beds of Turgai.' Comptes Rendus d. l'Acad. d. Sci. d. l'URSS, pp. 93, 94. COOPER, C. FORSTER. 1911. 'Paraceratherium bugtiense, a new genus of Rhino- cerotidae from the Bugti Hills of Baluchistan.-Preliminary No- tice.' Ann. and Mag. Nat. Hist., (8) VIII, pp. 711-716, PI. x. 1913. 'Thaumastotherium osborni, a new genus of Perissodactyles from the Upper Oligocene deposits of the Bugti Hills of Baluchistan.-Pre- liminary Notice.' Ann. and Mag. Nat. Hist., (8) XII, pp. 376- 381. 72 19361 Granger and Gregory, Baluchitherium from the Oligocene of Mongolia 73

1923. 'Baluchitherium osborni (? syn. Indricotherium turgaicum, Borris- siak).' Philos. Trans. Roy. Soc. London, (B), "L," pp. 35-66. 1924. 'On the skull and dentition of Paraceratherium bugtiense: a genus of aberrant rhinoceroses from the Lower Miocene deposits of Dera Bugti.' Philos. Trans. Roy. Soc. London, (B), pp. 212, 369-394, 19 figs. DIETRICH, W. 0. 1927-1929. 'Uber Rekonstruktionen fossiler Saugetiere.' Zeitschr. f. Saugetierkunde, II Band, Heft 3, 1929, pp. 177-186. [Estimated ratios of limb segments in Baluchitherium but not based on associated specimens. ] GRANGER, WALTER, AND GREGORY, WILLIAM K. 1935. 'A revised restoration of the skeleton of Baluchitherium, gigantic fossil rhinoceros of Central Asia.' Amer. Mus. Novitates, No. 787, 3 pp., 2 figs. GREGORY, WILLIAM K. 1928. (With Harold J. Cook.) 'New material for the study of evolution: a series of primitive rhinoceros skulls (Trigo- nias) from the Lower Oligocene of Colorado.' Proc. Colorado Mus. Nat. Hist., VIII, No. 1, 32 pp., 6 pls., 8 tables, 5 graphs. 1935. 'Building a super-giant rhinoceros.' Natural History, XXXV, No. 4, pp. 340-343, 3 figs. KROKOS, W. I. 1917. 'Aceratherium Schlosseri Web. du village de Grebeniki du gouvernement de Kherson.' Mem. Soc. Rural Economy of New Russia, LXXXVII, part 2, 96 pp., 3 pls. MAYET, LUCIEN. 1924. 'Un Rhinoc6ros g6ant: Le Baluchithere.' La Nature, No. 2596, pp. 1-3. OSBORN, HENRY FAIRFIELD. 1898. 'The extinct rhinoceroses.' Mem. Amer. Mus. Nat. Hist., I, part 3, pp. 75-164, Pls. XII A-XX. 1923. 'Baluchitherium grangeri, agiant hornless rhinoceros from Mongolia.' Amer. Mus. Novitates, No. 78, pp. 1-15. PAVLOW, MARIE. 1922. 'Indricotherium transouralicum, n.sp.' Bull. d. 1. Soc. d. Naturalistes d. Moscou, Nouvelle S6rie, XXXI, pp. 95-116, 2 pls. PETERSON, 0. A. 1911. 'A mounted skeleton of Diceratherium cooki Peterson.' Ann. Carnegie Mus., VII, No. 2, pp. 274-279, 1 pl. PILGRIM, GUY E. 1910. 'Notices of new mammalian genera and species from the Tertiaries of India.' Rec. Geol. Sur. India, XL, part 1, pp. 63-71. ["Aceratherium bugtiense, n.sp.-From the Upper Nari of the Bugti Hills; allied to A. perimense, but larger and more primitive in character, possessing no crochet and a hardly appreciable post- fossette." P. fi5.] 1912. 'The vertebrate fauna of the Gaj Series in the Bugti Hills and the Punjab.' Mem. Geol. Surv. India, Palaeontologica Indica, (N. S.) IV, Mem. No. 2, pp. 1-83, Pls. I-xxx. [Aceratherium bugtiense Pilgrim, pp. 26-30, Pls. vIII, IX, X. ] SCHMALTZ, REINHOLD. 1909. 'Atlas der Anatomie des Pferdes.' Zweiter Teil: Topographische Myologie. Taf. 24-62. Berlin. 4to. WOOD, HORACE ELMER, 2D. 1927. 'Some early tertiary rhinoceroses and hyraco- donts.' Bull. Amer. Paleontology, XIII, No. 50, pp. 1-89, 7 pls. 1932. 'Status of Epiaceratherium (Rhinocerotidae).' Jour. Mammalogy, XIII, No. 2, pp. 169-171. PLATE I Endocranial cast of Baluchitherium grangeri. Type skull. BULLETIN A. M. N. H. VOL. LXXII, PLATE I

occip.pole tran verse f7ssure

forlac. ant. + for: r (7T1-YiP ,-P,z2,P,e) PLATE II Interior of braincase of Baluchitherium grangeri. Type skull. A. Oblique view. B. Inner side view, partly restored. BULLETIN A. M. N. H. VOL. LXXII, PLATE II

(mnt. carotid etc.) PLATE III Sixth cervical vertebra of Baluchitherium grangeri compared with that of white rhinoceros (Ceratotherium sirnum). About 1/3 natural size. BULLETIN A. M. N. H. VOL. LXXII, PLATE III

i-X.. PLATE IV Model of Baluchitherium grangeri to scale with six-foot man. By John W. Hope. 0~~~~~~~~~~~~~~~~~~~~~~~~~~~~......

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