[=Kenyapithecus] at Fort Ternan, Kenya, in Deposits Which Have Been Dated by the K/A Method As About 14 Million Years Old

528 ANTHROPOLOGY: E. L. SIMONS PROC. N. A. S.

9 Garby, L., and M. Hjelm, Blut, 9, 284 (1963). 10 Borsook, H., J. B. Lingrel, J. L. Scaro, and R. L. Millette, Nature, 196, 346 (1962). These authors fractionated rabbit reticulocytes in BSA density gradients with methods developed in the early stages of this research. 11 Mateyko, G. M., and M. J. Kopac, Ann. N. Y. Acad. Sci., 105, 183 (1963). 12 Bock, R. M., and N. S. Ling, Anal. Chem., 26, 1543 (1954). 13 Lakshmanan, T. K., and S. Lieberman, Arch. Biochem. Biophys., 53, 258 (1954). 14 Bukantz, S. C., C. R. Rein, and J. F. Kent, J. Lab. Clin. Med., 31, 394 (1946). 15 de Duve, C., J. Berthet, and H. Beaufay, Progr. Biophys. Biophys. Chem., 9, 326 (1959). 16Holmquist, W. R., and J. R. Vinograd, Biochim. et Biophys. Acta, 69, 337 (1963). 17 Borsook, H., E. H. Fischer, and G. Keighley, J. Biol. Chem., 229, 1059 (1957). The composi- tion of NKM in gm/i is 7.61 NaCl, 0.389 KCl, 1.52 MgCl2-6H20. The osmolarity is 0.293. 18 Eagle, H., J. Exptl. Med., 102, 595 (1955). 19 Brecher, G., E. F. Jakobek, M. A. Schneiderman, G. Z. Williams, and P. J. Schmidt, Ann. N.Y. Acad. Sci., 99, 242 (1962). 20 Kabat, D., and G. Attardi, personal communication. 21 Ponder, E., Hemolysis and Related Phenomena (New York: Grune and Stratton, 1948). 22 Scatchard, G., A. C. Batchelder, A. Brown, and M. Zosa, J. Am. Chem. Soc., 68, 2610 (1946). 23 Hearst, J. E., and J. R. Vinograd, these PROCEEDINGS, 47, 999 (1961). 24 This equation may be expressed in terms of tonicity and liquid volume, V. (dp/dV)ef= dp/dV - dT/dV dp/dT. The method of evaluating dp/dV has been described. The value of dT/dV was obtained as the product AT/Ap-dp/dV, where AT and Ap are the difference in tonicity and density, respectively, in the two BSA solutions used in the preparation of the gradient. Buoyant density distributions may be obtained with the above procedures in centrifuge tubes of arbitrary shape. 26 Clark, P., Australian J. Sci., 22, 216 (1959). 26 Burwell, E. L., B. A. Brickley, and C. A. Finch, Am. J. Physiol., 172, 718 (1953). 27 Neuberger, A., and J. S. F. Niven, J. Physiol., 112, 292 (1951). 28 Farquhar, W. J., and E. H. Ahrens, Jr., J. Clin. Invest., 42, 675 (1963).

ON THE MANDIBLE OF RAMAPITHECUS* BY ELWYN L. SIMONS DEPARTMENT OF GEOLOGY, YALE UNIVERSITY Communicated by G. E. Hutchinson, December 26, 1963 During the past three years a number of findings have enlarged scientific under- standing of the initial differentiation of hominids from pongids. These advances are the outgrowth of significant developments in the study of man-like hominoids of Miocene and Pliocene age, recovered from deposits in Africa and Eurasia. In order of their occurrence these additions to knowledge are as follows: (1) The discovery and description by Dr. L. S. B. Leakey of an African member of Ram- apithecus [=Kenyapithecus] at Fort Ternan, Kenya, in deposits which have been dated by the K/A method as about 14 million years old. (2) The assignment to Ramapithecus by Simons' of a second maxilla (a referred specimen of Dryopithecus punjabicus originally figured by Pilgrim,2 from Haritalyangar in the Nagri zone, Siwalik Hills, North India). (3) The recent determination at Yale that several known mandibles from the latest Miocene and/or early Pliocene of the Siwaliks can plausibly be referred to Ramapithecus. This contribution is an attempt to relate the first two of these discoveries to previously published discussions of Downloaded by guest on September 24, 2021 VOL. 51, 1964 ANTHROPOLOGY: E. L. SIMONS 529

Ramapithecus (Leakey,3 Simons' 4) and to present new evidence showing that mandibles of the earliest hominid, Ramapithecus, are known from the latest Miocene and/or Pliocene of the Siwalik Hills, India. It seems advisable initially to point out that a revision of dryopithecine taxonomy will soon be forthcoming (Simons and Pilbeam,6 in press). Background and conclusions on dryopithecine taxonomy are presented in fuller detail in that study. Abbreviations.-A.M.N.H., American Museum of Natural History, New York; B.M.N.H., British Museum (Natural History); C.M.N., Coryndon Museum of Natural History, Nairobi; G.S.I., Geological Survey of India, Calcutta; M.C.Z., Museum of Comparative Zoology, Harvard; N.M.N.H.P., National Museum of Natural History, Paris; and Y.P.M., Yale Peabody Museum, New Haven. Material.-The type species of the genus Ramapithecus, R. brevirostris, is founded upon a right maxilla (Y.P.M. 13799) from the Siwalik Hills of North India containing the root of J2, alveolus of I', canine alveolus, and P3-M2. G. E. Lewis (written communication, 1964) states that it is probable that the specimen is not from the Tatrot as was originally indicated, but from the cuesta scarp at Hari- talyangar [Nagri(?), Simons 1961: 1; Lewis 1937: 142]. Recent intensive collect- ing in the Tatrot (?) north of Haritalyangar by K. N. Prasad (personal communication) failed to produce any fossil Primates, although he recovered without dif- ficulty a series of Primates at the cuesta scarp. Consequently, the occurrence of Primates in the Tatrot of the Siwaliks has not been proved. G. E. Lewis has kindly supplied the following statement regarding the collection of Y.P.M. 13799: Number 13799 was brought to me by a native, who took me to a spot some distance northward from Haritalyangar and insisted he found it there. To be sure it may have occurred there, but I eventually decided that he wanted to hide its true locality. After my 1937 publication, and after Krynine6 had had such good success in petrographic studies of large rock samples from many Siwalik localities, we examined the matrix on Y.P.M. 13799 and found it to be identical with that on the collections from Haritalyangar cuesta scarp-which as Pilgrim says2 "represents a decidedly older type than that of Dhok Pathan." I had originally intended to publish my later views, but other matters and the war intervened. Recently Simons referred a second right maxilla definitely from Haritalyangar (G.S.I. D-185) to Ramapithecus.1 This identification demonstrates the occurrence of this primate at Haritalyangar and reinforces the later view of Lewis that this site could be that of the type. An isolated upper third molar also from this site, recently recovered in India, is now under study by K. N. Prasad. It is of the proper, small size to belong to this same species. Thus, upper dental remains of three individuals have been recovered to date in India, all apparently from the Nagri zone cuesta at Haritalyangar. A fourth portion of an upper dentition representing this species is that of Ramapithecus [=Kenyapithecus] from Fort Ternan, Kenya. Since lower jaws with dentitions are quite commonly recovered in much greater numbers than are maxillae and upper teeth in dissociated fossil concentrations of continental deposition, there is a very high probability that if as many as four sepa- rate parts of upper dentitions of this taxon have been found, mandibular materials of this species exist unrecognized among known dryopithecine specimens. Approxi- mately four dozen binomials have been coined to date for fossils which may represent members of the Dryopithecinae. Not one of the types of any of these species Downloaded by guest on September 24, 2021 530 ANTHROPOLOGY: E. L. SIMONS PROC. N. A. S.

include definitely associated upper and lower dental materials. This suggests that some of the upper and lower dentitions from these deposits with different species names may represent the same species. The tentatively referred mandible of Ramapithecus, cf. R. brevirostris (Gregory, Hellman, and Lewis8) represented by a cast Y.P.M. 13870,7 does not, in my opinion,4 belong to the genus Ramapithecus, a conclusion which Lewis now believes to be justified, inasmuch as the mandible was referred (in Gregory, Hellman, and Lewis, 1938: 21) to Ramapithecus brevirostris only because of possible occlusal relation- ships with the holotype maxilla, whereas it was recognized that "the teeth are somewhat too narrow and the P- has its long axis too anteroposteriorly oriented to fulfill completely the requirements of a lower dentition of the genotype." Other reasons for doubting this reference are: (1) The specimen (G.S.I. D-618) is from a different horizon (Chinji) and locality (Salt Range) than are the maxillary materials of Ramapithecus. (2) Unlike most dryopithecines, this specimen has a distinct simian shelf. (3) It shows marked lower premolar heteromorphy which is not known to date in undoubted Hominidae. Should Ramapithecus have possessed a sectorial P3, like that seen in G.S.I. D-618, such a lower tooth would have produced a deep wear facet on the anterior face of P3 in Y.P.M. 13799, but no such facet exists in this specimen. In addition, the small upper canine of Ramapithecus figured by Leakey (not separated from P3 by a diastema in any of the three maxillae) will not allow for functional occlusion of a sectorial P3 such as that of G.S.I. D-618 (Y.P.M. 13870, cast). G.S.I. D-618 together with a second lower jaw (left side with dam- aged P4-M3) from the upper Chinji zone near Domeli, India (B.M.N.H. M-15243) both show small size and mandibular gracility differing from all other Siwalik hominoid mandibles. In these two mandibles there is fairly marked development of the simian shelf-otherwise both fall within the metric and morphologic range of variability of Proconsul africanus. When B.M.N.H. M-15243 and G.S.I. D-618, both of which have been separated in the mid-line of the symphysis, are set together along this line, they show a typical pongid U-shaped dental arcade, which could not occlude with that inferred for the Ramapithecus palate by Simons.4 These two specimens appear to represent in decreasing order of probability, (a) a new species, (b) a pigmy race of one of the described Eurasian pongids, or (c) an Indian variety of Proconsul africanus. Pilgrim's discussion2 of Dryopithecus punjabicus suggests a method of determining maxillary and mandibular associations in Ramapithecus. The material he assigned to his new species D. punjabicus included the type, consisting of portions of the right and left horizontal rami of the same mandible (G.S.I. D-118, and D-119), the right containing M3 and the left M2, and a referred maxilla (G.S.I. D-185). The subsequent taxonomic allocation of these two finds is interesting. Lewis9 assigned the mandibular fragments to the genus Bramapithecus, while (as is stated above) Simons1 referred the maxilla to Ramapithecus. If one, however, accepts Pilgrim's reason for placing these upper and lower dental materials in the same species, a view which I will attempt to confirm below, then the obvious implication of taxonomic work to date is that Bramapithecus mandibles are the lower jaws of Ramapithecus. Unless species distinctions can be supported, the prior binomial for this earliest known hominid must be Ramapithecus punjabicus (Pilgrim) 1910 and the following "species" become junior synonyms of this taxon; Downloaded by guest on September 24, 2021 VOL. 51, 1964 ANTHROPOLOGY: E. L. SIMONS 531

Ramapithecus brevirostris, Bramapithecus thorpei, Bramapithecus (?) sivalensis, and Kenyapithecus wickeri. Pilgrim's Contribution.-The type mandibular fragments of Ramapithecus punjabicus (Pilgrim) came from a high horizon in the Chinji Zone near the village of Chinji-the referred right maxilla, G.S.I. D-185, from the Haritalyangar cuesta scarp in the Nagri zone (Pilgrim, 1915: 9, 16). In 1915 Pilgrim2 dealt with the question of whether the two specimens could have been sufficiently separated temporally to lessen the probability of their belonging to the same species. He concluded that this was not the case. Problems regarding the rate of sedimentation, stratigraphic disconformities, and lithology in the Siwaliks are extensively discussed by Colbert."0 From this discussion, and others, it seems clear that the rate of sedimentation while the Upper Chinji-Nagri Zones were accumulating was rapid and that there are no abrupt faunal or lithologic changes between these two zones. There is some reason to think that typical Hipparion may not occur in the Chinji Zone, as was reported by some earlier workers, while other authors, such as Borisiak,11 present evidence that the Chinji has the oldest Hipparion fauna in Asia. These data would seem to favor a pre-Pliocene age for the Chinji. The primate materials here referred to Ramapithecus punjabicus come from both the Upper Chinji and the Nagri, and I would agree with Pilgrim that but one species, R. punjabicus, occurs in the two zones. Associated faunas of both Indian and African specimens suggest that known Ramapithecus lived at about the time of the Miocene-Pliocene temporal boundary. In the absence of a series of geo- chemically dated faunas or detailed studies of faunal correlation between mammals of the Fort Ternan and Siwalik localities, no more precise determination of the temporal position of Ramapithecus is possible at this time. It seems unlikely, however, that the temporal range of materials here referred to Ramapithecus punjabicus will ever prove to be sufficiently great to justify two or more time- successive species populations. At least l am not aware of any scientific evidence, now available, which would warrant such a conclusion. It is well known that the reference of unassociated upper and lower dentitions to the same species has its weaknesses. However, I believe that Pilgrim's decision to do so in this case can be justified on morphological grounds. Pilgrim2 sum- marized his conclusions on this point as follows: The dimensions of these teeth, the moderately low cusps, the complexity of the folding of the enamel, and, above all, the peculiar serrated outer edge of the molars incline one so strongly to the opinion that we have before us a maxilla and mandible which belong to the same species, that, unless fairly conclusive evidence were forthcoming of a close affinity to another genus, or of the existence of features in the maxilla, which told against an affinity with Dryopithecs [now, in this case, Ramapithecus], I should feel fairly certain that the similarity mentioned between the mandible and the maxilla indicated specific identity. The Yale Mandibles.-Three specimens referable to Bramapithecus are in the Yale collections-all are mandibular fragments with molars present (Y.P.M. Nos. 13806, 13814, and 13833). They are closely comparable to the type of Ram- apithecus punjabicus (G.S.I. D-118-119) although, as in all other hominoids, there is some variation in the crown pattern of the molars. The four specimens have in common a thick, robust mandible in the M2-3 region which, however, is quite shallow vertically. In this feature they differ significantly from Eurasian Downloaded by guest on September 24, 2021 532 ANTHROPOLOGY: E. L. SIMONS PROC. N. A. S.

DRYOPITHECUS SIVAPITHECUS SUGRIVAPITHECUS PROCONSUL

N.M.N.H.P? \ .M.N.H. J ( A.M.N.N 6.5.1. YPM r S N.H. C MN. 0-1 M-10970 19413 0-196 13611 M-1486 R (1942)

BREADTH/DEPTH 54.54.5. 53....53...... 55.2 ...... 57.3...... 6...... 5 54.1

RAMAPITHECUS

| *.M.N.H. 0.5.5. 9 0.6.5. 7YPM. / PM.. ) M-13264 0-119 D-116 13614 13606

DREP0TH 62.1..... 70.0...... 70.9..75.970 . 71.7. e FIG. 1.-Diagrammatic vertical sections of hominoid mandibles across mid-line of M3 showing distinctly lower breadth-depth ratios in the Dryopithecus group (A) than in Rama- pithecus (B). Diagrams are anterior views of left rami and reversed views of right rami. * Depth estimated either by adding height of M2, when M' missing, or by projecting the lower outline of mandible. and African dryopithecines, in which the mandible under M3 is considerably deeper relative to its thickness (Fig. 1). The molars tend to be quadrate and crowded rather than anteroposteriorly elongate and spaced out as in Dryopithecus and Proconsul. The one lower molar from Fort Ternan, figured by Leakey,3 and which he regards as belonging to the same individual as his East African Ramapithecus maxillae also has this shape. In Y.P.M. 13806 and 13814, M2 and M3 are essentially the same length. In the Dryopithecus group M3 is typically longer than M2. As in Ramapithecus upper molars, the individual cusps do not stand out in relief as in African and Indian pongids. Distinct, large, wear facets are present on the anterior and posterior faces of M2 in G.S.I. D-118, Y.P.M. 13806 and 13814, and on the anterior face of M3 in G.S.I. D-119 and Y.P.M. 13806, 13814, and 13833. Par- ticularly in Y.P.M. 13814, the type of "Bramapithecus thorpei," wear against adja- cent teeth has cut deep, arcuate "contact facets" into the anterior margins of M2 and M3. This, together with the quadrate form of the teeth, strongly suggests that the animal must have had a short face, and a crowded tooth row which lacked diastemata. Mr. D. R. Pilbeam has recently pointed out to me that the horizontal ramus begins to turn toward the symphysis at the level of M1 while in dryopithecines, such as Sugrivapithecus, the inward curvature toward the symphysis commonly does not begin until about the level of P3 (Fig. 2). This distinction is a direct reflection of the differences between tooth arrangement in a short-faced form and that seen in long-snouted hominoids. It is interesting to note that these same features, deep anterior contact facets, subequal size of quadrate M2 and Downloaded by guest on September 24, 2021 VOL. 51, 1964 ANTHROPOLOGY: E. L. SIMONS 533

M3, shallow but robust mandible and lingual curvature toward the sym- » -X' physis at the level of Ml, are present in the type of "Telanthropus capensis" and are about equally developed in several of Dart's Makapan mandibles of Australopithecus. Perhaps even more significant is the presence of / these same features, together with other similarities, in the Lantien jaw -1 discovered on July 19, 1963 in North- west China."2 This mandible may be Australopithecus (s.l.), or even a member ofmuch more recent Homo; which- ever it is, resemblances between "Bramapithecus" and such hominid jaws are striking. These strong similarities reinforce the probability that "Bramapithecus" mandibles belong A B to a taxon on or near the main line FIG. 2.-Comparison of Mio-Pliocene hominid mandible (A) Y.P.M. 13814, with of human descent. This conclusion contemporary Ramapithecus,dryopithecine (B) Sugrivapithecus, can be reached independently without Y.P.M. 13811. Unlike living or fossil apes, in the hominid the mandible begins to turn inward toward reference of these mandibles to the the symphysis at the level of Ml and the molars same species as the previously dis- are rounded with low relief. In the Sugrivapithecus cussed maxillae of Ramapithecus. mandible, Ms is restored from those of other dryo- Nevertheless, the two sets of data pithecines. Scale Xl. are interrelated and reinforce the association of upper and lower jaws. It seems most unlikely that hominid mandibles differentiated in one species while hominid maxillae first reached the hominid grade in a different taxon. Lewis clearly recognized the hominid ties of "Bramapithecus" discussed above, but until more be- came known about Ramapithecus maxillae, the phyletic and taxonomic implication of structure in "Bramapithecus" remained uncertain. Lewis13 commented percep- tively in his initial discussion of "Bramapithecus:" "...the sculpture of the crowns is highly suggestive of many human molars. The deep folds, persistent in spite of severe wear, are notable features. The author .., believes that the genus has affinities with Dryopithecus and was probably derived from a common stock. It may very well lie near to the stem which led to the Hominidae proper." In his revision of 1937, Lewis9 placed the four partial lower dentitions, discussed above, in the genus "Bramapithecus" but retained three different species, "B. thorpei," "B." punjabicus, and "(?) B. sivalensis." Lewis' recognition of the taxonomic affinity of these materials was another important step but, in view of known variability in living Hominoidea, these materials need not indicate more than one species. For this species "B." punjabicus (Pilgrim) 1910 has priority. An ad- ditional specimen (here assigned to this species), B M.N.H. M-13264, from the Attock district, Punjab is either from the uppermost Chinji or from the Nagri. Three of the finds, Y.P.M. 13833, 13834, and the'type G.S.I. D-118-119,!come from the Upper Chinji And one of them, "(?) B." sivalensis, YP.M, 13806, from the Downloaded by guest on September 24, 2021 534 ANTHROPOLOGY: E. L. SIMONS PROC. N. A. S.

Nagri age cuesta scarp at Haritalyangar. Thus, at Haritalyangar as well as at Ft. Ternan, Kenya, lower teeth of this sort have been found at a locality which has yielded Ramapithecus maxillae. It seems most unlikely that two species were independently differentiating toward Hominidae at this time (end of the Miocene and/or early Pliocene), one of which is known only from mandibles, the other only from maxillae, but both of which occur at the same sites. I can find no morphological, temporal, or distributional evidence for thinking that all the materials belonging to the "species" listed below should not be referred to one species, Ramapithecus punjabicus, which was originally diagnosed by Pilgrim"4 in 1910. In regard to the question of geographic distribution of this species, it is interesting that a right M2 from the Pontian of Melchingen, Wirttemberg, assigned by Koken'6 to Dryopithecus suevicus shows all the characteristic molar features outlined in the diagnosis below for Ramapithecus punjabicus. A left M2 from the same collection, markedly different, but also included in D. suevicus by Koken is a Dryopithe- cus-presumably D. fontani. Systematics Order PRIMATES Suborder Anthropoidea Superfamily Hominoidea Family HOMINIDAE Genus Ramapithecus Lewis, 1934 Dryopithecus Lartet, Pilgrim (partim) 1915:16 Bramapithecus Lewis 1934:173 Dryopithecus Lartet, Lewis (partim) 1934:171 Kenyapithecus Leakey, 1962:690 Type Species.-Ramapithecus brevirostris Lewis 1934'3 (considered here to be a subjective synonym of Dryopithecus punjabicus Pilgrim, 1910).14 Generic Diagnosis.-Differs from Australopithecus and members of the Dryo- pithecus group in the following general features: slightly smaller over-all size (ex- cept for Proconsul africanus), shallower mandible, less complex patterns of tooth crenulation, shorter face, little or no evidence of cingula or Carabelli's cusps. Incisors and canines reduced, in relation to cheek-tooth size when compared to Dryopithecus, but not as markedly as in Australopithecus; incisor procumbency intermediate. Differs from Dryopithecus and other apes in showing more widely spaced and much lower crowned molar cusps, so that central or occlusal fovea of molars covers more of the crown surface of the tooth, and the sides of the upper molars (particu- larly) are more vertical. Also differs from Dryopithecus in showing a larger and lower canine fossa, an arched palate, arcuate tooth row, and much shorter rostrum. Specific Diagnosis.-Same as for the genus, which is monotypic. Referred Species.-Ramapithecus brevirostris Lewis (1934:162), Bramapithecus thorpei Lewis (1934:173), Bramapithecus(?) sivalensis Lewis (1934:171), Kenya- pithecus wickeri Leaky (1962:690). Downloaded by guest on September 24, 2021 VOL. 51, 1964 ANTHROPOLOGY: E. L. SIMONS 535

Summary.--Earlier discussions of the mandible of Ramapithecus, the oldest probable forerunner of man, have been based on a jaw which actually belongs to a species of Dryopithecus. Materials previously assigned to species of the genus "Bramapithecus" are here ;__, referred to the genus and species Ramapithecus punjabicus. This species probably occurred widely iClH-2-- throughout Eurasia and Africa about 14 or 15 million years ago. Considered together, known ma- - terial of R. punjabicus indicates a Pan-sized primate with short FIG. 3.-Hypothetical appearance of the face ofRama- face, arcuate palate, and an pithecus. Maxilla Y.P.M. 13799; mandible, Y.P.M. AsrokAustralopithecus-likeofmandible 13814"Kenyapithecus(reversed). wickeri."Dotted outlineDashedof linescanineconjectural.from type (Fig. 3). Dental and facial Scale XO.72. characters are so close to Austra- lopithecus africanus as to make difficult the drawing of generic distinctions between the two species on the basis of present material. Provisionally the two genera, Ramapithecus and Australopithecus, are retained as distinct because of their consider- able time separation. Ramapithecus punjabicus is almost certainly man's forerunner of 15 million years ago. This determination increases tenfold the approximate time period during which human origins can now be traced with some confidence. The author wishes to acknowledge valuable discussions of the subject with W. E. LeGros Clark and D. R. Pilbeam. Figures were prepared at Yale by Miss Polly Porter and Miss Anna Held. * The research reported here was supported in part by a grant in geology from the National Science Foundation, no. GP-433, and by grants from the Wenner-Gren Foundation of New York. I Simons, E. L., Science, 141, 879 (1963). 2Pilgrim, G. E., Records Geol. Surv. Ind., 45, 1 (1915). 3Leakey, L. S. B., Ann. and Mag. Nat. Hist., 13, 689 (1962). 4Simons, E. L., PostiUa, 57, 1 (1961). 6 Simons, E. L., and D. R. Pilbeam, Folia Primatologica, in press. 6Krynine, P. D., Amer. J. Sci., 34, 422 (1937). 7The original of this mandible G.S.I. D-618 cannot now be located in the collections at Cal- cutta, therefore further comment as to the taxon which it represents will have to be based on Y.P.M. 13870 (cast). Simons4 inadvertently referred to this specimen as Y.P.M. 13807, now M.C.Z. 8386, the type of Ramapithecus hariensis Lewis (1934), which was ultimately assigned to Sivapithecus sivalensis Lewis8 and which is not a Ramapithecus. 8 Gregory, W. K., M. Hellman, and G. E. Lewis, Carnegie Inst. Wash. Publ. No. 495,1 (1938). 9 Lewis, G. E., Amer. J. Sci., 34, 139 (1937). "Colbert, E. H., Trans. Amer. Philos. Soc., 26, 1 (1935). 11 Borisiak, A. A., (translation) Internat. Geol. Rev., 4, 845 (1962). 12 Anon., IUust. Lond. News, 243 (6483), 742 (1963). 13Lewis, G. E., Amer. J. Sci., 27, 161 (1934). 14 Pilgrim, G. E., Records Geol. Surv. Ind., 40, 63 (1910). ' Koken, E., Fatrher durch die samlungen (Tubingen, Stuttgart: Geol.-Min. Inst., 1905), p. 1. Downloaded by guest on September 24, 2021