‘bed SS eee eu ee eT goer ere - > mere mcm es esse wartrtatta/ sie Sas 89 none tage eone aceiene” O HARVARD UNIVERSITY cD LIBRARY OF THE Museum of Comparative Zoology Uliny Pilla YALE PEABODY MUSEUM or Naturat History Number 64 August 7, 1962 New Haven, Conn. TWO NEW PRIMATE SPECIES FROM THE AFRICAN OLIGOCENE Exwywn L. Simons One of the major objectives of the Yale 1961-1962 Paleon- tological Expedition to the Fayum region of Egypt was to re- cover a larger sample than previously known of the earliest mammalian microfauna from the continent of Africa, that of the Fluviomarine formation of the Fayum early Oligocene. During the course of our investigations the expedition staff succeeded in locating specimens assignable to two new species of Primates. In view of the considerable interest in, and rarity of, Old World Primates dating from this epoch it seems advis- able to publish a preliminary description of these two forms without delay, so that they will be available for consideration by other authors. It is intended that a fuller analysis of their morphology and relationships will be included in a study of the Fayum mammalian microfauna now being prepared by the writer. ACKNOWLEDGMENT I wish to express my sincere thanks to Dr. Osman Moharam, Director of the Department of Geological and Mineralogical! Research, Ministry of Industry, United Arab Republic; Dr. 2, Postilla Yale Peabody Museum No. 64 Riad Higasy, former Minister of Industry, and Dr. Y. Shawki Moustafa, Curator of Fossil Vertebrates, Cairo Museum of Geology, for generously supplying facilities and assistance which contributed greatly to the success of our expedition to the Fayum badlands. The field research of which this report 1s an outgrowth was made possible by a grant in Geology of the National Science Foundation. Illustrations were prepared by Margaret EK. Freeman of New Haven and their execution was partly financed by a grant from the Wenner-Gren Foundation of New York. The photograph (figure 3) was prepared by John Howard of the museum staff. ABBREVIATIONS A.M.N.H. _.. American Museum of Natural History, New York. YCC:P-E: Yale-Cairo, Paleontological Expedition (field numbers). Ys PAVic .. Yale Peabody Museum, New Haven. SYSTEMATICS Class MamMatia Order PRIMATES Suborder ANTHROPOIDEA Infraorder CATARRHINI OLIGOPITHECUS,' new genus Type: Qligopithecus savagei, new species Generic characters: Lower dental formula 2?. 1. 2. 3., size of mandi- ble approximately that of the living ceboid primate Leontocebus rosalius, slightly smaller than its contemporary Propliopithecus haeckeli. Differs from the latter in having a slightly shallower mandibular ramus, more ‘Named with reference to the Oligocene occurrence of this catarrhine. August 7, 1962 Two New Primate Species 5) anteroposteriorly elongated P., distinct paraconid on P,-M,, lower molar external cingula less distinct and hypoconulid shifted much more lingually than in Propliopithecus and not distinctly separated from entoconid. Differs from Parapithecus fraasi in its larger size and in possession of undoubted, large canine anterior to P,, in absence of metaconid cusp on P,,—present in Parapithecus, in possession of anteroposteriorly elongated P., not seen in the latter genus, and in having a much more lingually ex- tended paraconid ridge on M,-. c \ Figure 1. External view of left mandible, Oligopithecus savagei, Type, Y.C.P.E. 207. x 3 approx. Discussion: In depth of jaw and canine morphology Oligo- pithecus much more nearly resembles Propliopithecus than it does Parapithecus or Apidium, three other primates from the Fayum early Oligocene. On the other hand, the anteroposteri- orly elongate P., a feature which typically characterizes post- Oligocene cercopithecoids and pongids, is not to be seen in any specimens of these three Fayum genera. Such a lower P, does occur in a damaged mandible, A.M.N.H. 13389 from the Fayum, lacking tooth crowns other than on P,, described by Simons (1961), but in spite of this resemblance P., of A.M.N.H. 13389 is three rooted, a feature of great rarity among Pri- mates, while that of Oligopithecus is more normal in being two rooted. Moreover, A.M.N.H. 13389 is, in comparable measure- ments, over twice larger than the type of Oligopithecus. Pre- sumably the former specimen belongs to yet another Egyptian Oligocene primate species, diagnosis of which will have to await recovery of more satisfactory material; see Simons (1961:3). 4 Postilla Yale Peabody Museum No. 64 The relationship of Oligopithecus to Moeripithecus mark- grafi, also from the Egyptian early Oligocene, is more obscure, since comparison of M,. crown patterns in the two forms fails to reveal any significant similarity. The overall mor- phology of the tooth series preserved in Oligopithecus appears to be more like that of certain Kocene prosimians than are these patterns in other Fayum Primates, except that the P,.. are lost, the horizontal ramus is deep compared to tooth crown height, and P, is anteroposteriorly elongate—all similarities to Old World Anthropoidea. Among Eocene Primates perhaps the greatest resemblances of Oligopithecus are to Omomyidae, Anaptomorphidae and Necrolemurinae, which fact agrees well with the supposition that Anthropoidea are more closely allied to the so-called tarsioids of Kocene times than they are to other known Eocene, and earlier, families of the order. The lingual po- sition of the M,.. hypoconulid in Oligopithecus creates a par- tial posterior loph parallel to the anterior loph, between meta- conid and protoconid, which might represent an early stage im the transition to the bilophodont lower molar pattern of Cer- copithecoidea. This possibility will require further confirma- tion before definite assignment of this primate to the latter superfamily would be advisable. Placement of Oligopithecus among the Hominoidea is also conceivable but M,-. crown patterns are so primitive that such an association is equally dubious. Clearly, however, the creature is not a ceboid or a prosimian. OLIGOPITHECUS SAV AGEI Figures 1 and 4. Type: Left mandibular ramus, with C-M.,, inclusive, Y.C.P.E. 207. Horizon and locality: Yale Expedition quarry E, Fossil Wood Zone, Fluviomarine Formation, early Oligocene age, Fayum Province, Egypt, about 2.3 miles northeast of American Museum Quarry A (1907) and about 2 Named for Dr. D. E. Savage of Berkeley University who discovered the type and only specimen. * By agreement with the Egyptian authorities concerned, types collected on the Yale 1961-62 expedition will eventually be deposited in the paleontologi- cal collections of the Cairo Museum of Geology. Pending such assignment they are here identified by their field numbers. August 7, 1962 Two New Primate Species 5 eight miles west-northwest of Quasr el Sagha Temple, thirty feet below the top of the Fossil Wood Zone, see Beadnell (1905). Specific characters: Not distinguished from generic. Discussion: The species, O. savagei, is at present known only from the type individual so that little can be said of varia- bility in this primate. Although M, is missing in Y.C.P.E. 207 the fact that it had fully erupted is indicated by the preserva- tion of the anterior root of this tooth in the type. A large wear facet on the anterior crest descending from P., protoconid indi- cates that this species must have possessed a sizeable upper canine which sheared against this facet. Measurements in millimeters of Oligopithecus savagei are listed below in comparison with similar measurements on con- temporary Fayum Primates taken from Kalin (1961). Mandibular dimensions (Q. savagei). Anteroposterior Length of C through M, 19.4 Depth of mandible beneath M. 9.5 Depth of mandible beneath P,, a 10.2 MEASUREMENTS OF TEETH Oligopithecus Parapithecus Propliopithecus Moeripithecus savagei fraasi haeckeli markgrafi left side left side left side right side Canine lene thie. 4 3.7 3.2 (Cor P2) 4.0 (right) wali S68dac 3.3 2.5 (Cor P.) 4.3 (right) lee lenoth 22. =-- 4.2 3.3 3.5 Wwacthe es 2r 3.1 2.5 3.8 P, leno beens 3.3 3.3 3.5 WLC heer 3.0 2.5 4.0 M, lengthieeee 4.2 a 4.8 5.0 width). =. ..-- 3.4 4.5 4.8 M, lengthier: 4.2 4.3 4.8 5.4 Width "ee a.5 - 3.5 3.4 4.5 5.4 6 Postilla Yale Peabody Museum No. 64 Genus APIDIUM Osborn, 1908 Type: dA pidium phiomense Osborn, 1908. Discussion: Although Osborn (1908) was reluctant to state the ordinal position of this species, it subsequenlty came to be regarded as a primate, and Gregory (1922) held that it might be related to the basic cercopithecoid stock or to Oreopithecus; see Simons (1959: 14). More recently a few authors have again questioned the primate status of A. phiomense but its resem- blance to Oreopithecus seems more than convergent, as was discussed by the writer (1960). Moreover, the discovery of several isolated upper teeth of a species of this genus, (de- scribed below) in association in one quarry with several lower jaws of this form, shows us that similarities to Oreopithecus in the upper dentition are about equal to those seen in the lower teeth. Nevertheless, 4 pidiwm species are rather unlike other known Oligocene Anthropoidea in that the cheek-teeth are more polycuspidate, the mandibular ramus is shallow, and the canine may have been small, or alternatively there may have been three premolars—inadequate preservation of the new materials rendering this point equivocal. Clearly the stock of Apidium had differentiated from that of other Fayum Pri- mates at a considerable remove in time from their common ocur- rence in the Karly Oligocene of Egypt, but it is in many ways closest to Parapithecus insofar as the two forms can be com- pared. The question, therefore, as to the relation of species of these two genera to the earliest undoubted Old World Anthropoidea is interrelated. Provisionally the study of Api- dium suggests that it, together with Parapithecus, may even- tually be ranked with some certainty among the Anthropoidea. APIDIUM MOUSTAFAI,* new species Figures 2, 3. Type: Left mandibular ramus with P.,-M,, Y.C.P.E. No. 260. Hypodigm: Type and Y.P.M. numbers 18007, left mandibular rumus with damaged P,-M,; 18008, unassociated right P*-M*; 18009 right mandi- bular ramus with P,-M.; 18018 left mandibular ramus with C?-M, ; 18042 *Named in honor of Dr. Y. Shawki Moustafa, whose earnest assistance and advice were most helpful to our Egyptian expedition. ~ ‘ August 7, 1962 Two New Primate Species left mandibular fragment with P,-M,; 18081, unassociated left P'?, left M?, left M., left M,, all from Quarry G. Horizon and locality: Yale Expedition Quarry G, below upper fos- siliferous zone, Fluviomarine Formation, early Oligocene age, Fayum Pro- vince, Egypt. One mile due north of American Museum Quarry A and approximately 100 feet above top of lower Fossil Wood Zone. Specific diagnosis: About one-eighth smaller than 4. phiomense, with slighter development of lateral basal cingula and fewer accessory cuspules on P, heel and on M,—.,, and as a whole, comparatively smaller and more primitive than in 4. phiomense, with posterointernal angle of M, produced more lingually. Figure 2. A. Crown view and B. External view, 4 pidium moustafai Type, Y¥.C:P.E. 260. x 5 approx. Discussion: This species is both smaller and earlier occur- ring than the type of A. phiomense. That a significant lapse of time exists between the populations from which the two species are derived seems probable in view of studies of other faunal elements from the upper part of the Egyptian Fluviomarine Formation now in progress. Although the exact locality of the type of A. phiomense is not known, it was reported by Osborn (1908) as having been discovered upon the upper fossiliferous 8 Postilla Yale Peabody Museuni No. 64 level, which begins about 100 feet stratigraphically above the horizon of Quarry G. If the practice of some past students of fossil Primates were followed here, a case for generic separa- tion of these two species could probably be made, but this type of splitting should not be attempted until these forms are much better known. MeraAsvuREMENTS IN MILLIMETERS OF Apidium moustafat YPM. “YPM. | TYPE: 180158 18009 Y.C.P.E. 260 CDG or Es Lemip:bhy (35 vane. c cere couseeters 2.5 Pv Gin i ke cen se=usk oscars Si: 2.0 | Pe lengthy eeacee ssomm eat: 2.7 2.7 2.8 width ean thee, Seco htestis 2.0 1.9 2.0 P, lemothi pss cies yer 2.7 2.7 2.8 Width sg. oe. scouts 2.3 2.2 2.3 M, Leno; theese era 3.5 3.6 3.7 Wildithh is g7se scien ae 2.8 2.7 3.0 M. Lengthy. ee creo sees 3.6 3.6 width 284s ora ke fe ieee 3.2 3.1 M, NSM 2 egos pen Homn s 3.6 4.0 V0(6 U8 ler eaerene mesa Hoes 3.0 2.9 Depth of mandible beneath Mie Bye eice ck seas eeats aes 6.0 tel 6.8 Anteroposterior length ORIN Eg ie Se in AG ete at 15.3 16.8 | Ease Le ree eeema er coer S50 8.6 9.0 8.8 August 7, 1962 Two New Primate Species 9 CONCLUSIONS Recent collections of Primates from the Fayum early Olig- ocene deposits of Africa indicate the existence of at least two new forms of Primates. Study of the first of these, Oligo pithe- cus savagei, suggests that it is assignable to the Anthropoide: and that it may represent an early stage in the differentiation of Cercopithecoidea. Near agreement in expected size, and ap- proximation in horizon and locality suggests that a catarrhine frontal bone described by Simons (1959) may provisionally be referred to this species. Placement of this frontal with A pidium or Parapithecus, which are also of suitable size, seems less ap- propriate in view of non-anthropoidean features such as are to be seen in their anterior dentitions. Known Propliopithecus and Moeripithecus are probably too large to have had a frontal this size, but of course this skull fragment could well belong to a species otherwise unkown. A second species, A pidium mousta- fai, here described, appears to represent an earlier and ances- tral population to Apidium phiomense which was recovered from the upper part of the Fluviomarine Formation. Newly discovered upper teeth of Apidium strengthen the view that species of this genus, together with Oreopithecus bamboli, rep- resent a fourth major group of Old World Higher Primates distinct from cercopithecoids, pongids and hominids. 10 Postilla Yale Peabody Museum No. 64 Figure 3. Right mandibular ramus of Apidium moustafai, Y.P.M. 18009. x 6 approx. August 7, 1962 Two New Primate Species ila Figure 4. Crown view of dentition, Oligopithecus savagei, Type, Y.C.P.E. 207. x 6 approx. 12 Postilla Yale Peabody Museum No. 64 REFFRENCES Beadnell, H. J. L., 1905. The topology and geology of the Fayum provence of Egypt. Publ. Sury. Dept., Ministry Public Works, Egypt, p. 1-101, 10 fig., 24 pl. Gregory, W. K., 1922. The origin and evolution of the human dentition. Baltimore, Williams and Wilkins Co., p. 1-548. Kahlin, J., 1961. Sur les primates de VOligocéne inférieur d’ Egypte. Annales de Paléontoiogie, tome 47, p. 1-48, 18 fig., 7 pl. Osborn, H. F. 1908. New fossil mammals from the Fayum Oligocene, Egypt. Bull. Amer. Mus. Nat. Hist. v. 24, art. 16, p. 265-272 6 fig. Simons, EK. L., 1959. An anthropoid frontal bone from the Fayum Oligocene of Egypt: the oldest skull fragment of a higher primate. Amer. Mus. Nov., no. 1976, p. 1-16, 4 fig. ——_———.,, 1960. Apidium and Oreopithecus. Nature. v. 186, no. 4727, p. 824-826, 1 fig. ——————, 1961. An anthropoid mandible from the Oligocene Fayum beds of Egypt. Amer. Mus. Nov. no. 2051, p. 1-5, 1 fig. | Harvard MCZ Libra IWAUOITAQA TIN 3 2044 066 305 236 ee Due Sat 574 | AHS Re ne ee . ee Fir PSS mere ernie See ee ~~ Re pA ace natn! pin i oe “ : aut ae eee ae a par enetetae ren : ; rs . 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