od he ‘ AM ay »y Le Le ; a SOM TEM FCN Ss Ayan dove a tr) Shah tA rt tA ees if MAD L het, 4 is ay Shey ae pts : : : eto =< t tircigs . f ee ' it ty ef “ ul @ ‘ : iv t: ' At ! i : ae beet beet ea hy ee I, f Hone ’ SE kae Te Te OM Pe o8 a aly ay ay ey Phy aa i$ . Lae (ite (yt ae Hygena 0 Mt At 7 Dt iY SAGES 5-0 ite. Tot 4 pa oe os * dba) he POSTILLA PEABODY MUSEUM YALE UNIVERSITY NUMBER 166 20 DEC. 1974 PARAPITHECUS GRANGERI (PARAPITHECI- DAE, OLD WORLD HIGHER PRIMATES): NEW SPECIES FROM THE OLIGOCENE OF EGYPT AND THE INITIAL DIFFERENTIATION OF CEROPITHECOIDEA ELWYN L. SIMONS POSTILLA Published by the Peabody Museum of Natural History, Yale University Postilla includes results of original research on systematic, evolution- ary, morphological, and ecological biology, including paleontology. Syntheses and other theoretical papers based on research are also welcomed. Postilla is intended primarily for papers by the staff of the Peabody Museum or on research using material in this Museum. Editor: Zelda Edelson Postilla is published at frequent but irregular intervals. Manuscripts, orders for publications, and all correspondence concerning publications should be directed to: Publications Office Peabody Museum of Natural History New Haven, Conn. 06520, USA Lists of the publications of the Museum are available from the above office. These include Postilla, Bulletin, Discovery, and special publica- tions. Postilla and the Bulletin are available in exchange for relevant publications of other scientific institutions anywhere in the world. Inquiries regarding back numbers of the discontinued journal, Bulletin of the Bingham Oceanographic Collection, should be directed to: Kraus Reprint Co. Route 100 Millwood, New York 10546 PARAPITHECUS GRANGERI (PARAPITHECIDAE, OLD WORLD HIGHER PRIMATES) NEW SPECIES FROM THE OLIGOCENE OF EGYPT AND THE INITIAL DIFFERENTIATION OF CERCOPITHECOIDEA ELWYN L. SIMONS Peabody Museum of Natural History and Department of Geology and Geophysics, Yale University, New Haven, Connecticut 06520 (Received May 16, 1974) ABSTRACT Among many primate fossils from the badlands of Oligocene age in the Fayum Province, Egypt, are specimens of a new species of the genus Para- pithecus. The new materials for the first time provide evidence of the up- per dentition and mandibular materials show that all the early determi- nations as to the dental formula of the type species of Parapithecus, P. fraasi, were incorrect. The new species, P. grangeri, is here described. It is suggested that the family Parapithecidae is best ranked in Cerecopith- ecoidea and that, in fact, Parapithecus and Apidium are the earliest known cercopithecoids. POSTILLA 166: 12 p. 20 DECEMBER 1974 2 POSTILLA 166 INTRODUCTION Several score fossil primates have been recovered in the course of seven expeditions to the Fayum badlands of Egypt, UAR, between 1961 and 1968. These specimens include some four dozen finds (mostly isolated teeth) of a new species of Parapithecus, larger than the type species and presumed to be somewhat younger than it. All known specimens of the new species of Parapithecus (described below) were recovered from Yale Quarry I which is located in the Upper Fossil Wood Zone of the Je- bel el Qatrani Formation about 250 feet below the top of that formation. The level of Quarry I is the highest Fayum horizon that is richly fossilifer- ous. The Jebel el Qatrani Formation is capped by a basalt that has been dated by the potassium/argon method at 24.7 + 2 million years B.P. by Evernden and Curtis at Berkeley and at 27.0 + 3 by Armstrong at Yale (see Simons and Wood, 1968). Geological evidence suggests that the basalt was implaced on the underlying Jebel el Qatrani Formation a considerable time after deposition of those beds. In places the entire Formation (110-270 meters thick) had been eroded away before the basalt flow occurred. Thus, a tentative age of 28 to 30 million years seems probable for the Upper Fos- sil Wood Zone from which the fossils described here were recovered. Such a dating supports the evidence derived from faunal correlation that all the Fayum mammalian fossils from the Jebel el Qatrani are of Oligocene age and that they are all older by around ten million years than are any other African deposits that yield fossil cercopithecoids. The new species of Parapithecus is of special interest as its dental an- atomy appears to provide plausible evidence of relationship to the ances- try of the Old World Monkeys, Ceropithecoidea. The new material also provides adequate evidence to make a definite settlement of the taxonomic position of not only Parapithecus but Apidium as well. The latter is rep- resented in our new collection by an even greater number of specimens. These two genera are by far the most common African Oligocene primates. They are known not only from jaws, teeth and cranial fragments but also probably are represented in the nearly 100 isolated postcranial bones from Yale Quarry I which are definitely primate. On grounds of their proper size, anatomy, and frequency of correlation with finds of jaws and teeth most of these can be provisionally referred to the Parapithecidae, to which both Parapithecus and Apidium belong. ABBREVIATIONS Abbreviations used in this paper are as follows: € canine (C' = upper canine, C, = lower canine) dP deciduous premolar (dP* = third upper deciduous premolar) M molar (M! = first upper molar, M, = second lower molar) P premolar (P* = third upper premolar, P, = fourth lower premolar) PARAPITHECUS GRANGERI 3 CGM _ Cairo Geological Museum, Cairo, Egypt SNM _ Naturhistorisches Museum, Stuttgart, Germany YPM Peabody Museum of Natural History, Yale University, New Haven, Connecticut SYSTEMATICS ORDER PRIMATES SUPERFAMILY CERCOPITHECOIDEA FAMILY PARAPITHECIDAE SUBFAMILY PARAPITHECINAE GENUS Parapithecus Schlosser 1910, 1911 TYPE Parapithecus fraasi (Fig. 2) GENERIC DESCRIPTION. Dental formula g-L33., as in only Apidium and probably Amphipithecus among catarrhines. Differs from the contempo- LG: rary parapithecine genus Apidium in showing comparatively larger € and markedly smaller M,, centroconid typical of Apidium absent and hypo- conulids of M,_, relatively reduced, principal upper cusps at corners of a square, not with hypocone much more lingually situated as in Apidium. Parapithecus lacks the large pericone cusp developed from the anterior part of the lingual cingulum of the protocone in Apidium. Differs from later Cercopithecidae and from all Old World Higher Primates, but agrees with Apidium in uniformly showing small central cusp in upper P?-* be- tween main inner and outer cusps and apparently homologous with the para- conule of M!-%. Differs from Apidium in showing no trace of the wrinkling and polycuspidation of teeth characteristic of the latter. Parapithecus grangeri, new species’ (Fig. 1) TYPE. CGM 26912, left mandibular ramus with P,-M,, collected from the eastern edge of Yale Quarry I, by E. L. Simons in February, 1966. 1This species is named in honor of the late Walter Granger of the American Mu- seum of Natural History, whose untiring collecting efforts in the Fayum in 1906 led indirectly to the discovery of earliest Higher Primates there. In an earlier paper (Simons, 1969) I used the name Parapithecus grangeri and presented drawings and photographs of its dentition and that of the type of P. fraasi. However, this was not intended to be the publication establishing the name of the new species, and a careful review of that paper shows that the technical phrasing of Article 13 (a) of the International Code of Zoological Nomenclature (Stoll, 1964) is not. satis- fied: ‘‘. . . a name published after 1930 must be . . . accompanied by a statement that purports to give characters differentiating the taxon.’’ Therefore the 1969 pa- per can be ignored for purposes of nomenclature. In my book on primate evolution 4 POSTILLA 166 HYPODIGM. Type and CGM 26918, right jaw fragment with P,-M,; YPM 21017, right mandibular fragment with M,,; 21019a, right mandibular frag- ment with dP, ,, M,_., M, in crypt; 23954, right jaw fragment with P,-M, and part of ascending ramus; 23973, left jaw fragment with M,_, and about 40 isolated upper and lower teeth at Yale. (This is a tentative count. Pos- itive identification is not possible for every one of the 40 teeth.) FIG. 1. Stereo pair of the occlusal view of the teeth, type specimen of Parapithecus grangeri, CGM 26912. Scale x 2. HORIZON AND LOCALITY. All known specimens from Yale Expedition Quarry I, Upper Fossil Wood Zone, Jebel el Qatrani Formation, Oligocene Epoch, Fayum Province, Egypt. SPECIFIC CHARACTERS. Comparable measurements on teeth and mandible ranging from about 10 to 25% larger than in type species, P. fraasi, which is presumably older and from lower in the section (see Table 1). P. gran- geri showing a tendency toward more marked reduction of M, relative to M, and with much larger and more robust mandible relative to absolute size of teeth in full adults (with M, erupted) than in type species. Mandib- (Simons, 1972: p. 191) the species P. grangeri was mentioned a second time as a species then in press, although again it was not my intention to make that brief ref- erence to the work that established the name. Even though no type specimens were designated, the passage did make a partially comparative statement: ‘‘Most of the new Parapithecus finds are 15 to 20 percent larger than the type of Parapithecus fraasi, which was evidently found at a lower level than Quarry I, where the new species occurs. This new parapithecine has been named Parapithecus grangeri (Simons, 1972).’’ The present contribution is the actual paper that was then in press at a date prior to publication of my book, but because of a difference about its editing, that paper was not published in the journal to which it had been sub- mitted, and it is here published for the first time as the initial description of this species. ‘JUIUIIAINSVIU PIIBUINISI = yu ‘(LIGI) Jassopyog Jo asoyy Burjoa1109 ([ 96) UNRY wor sjUIUIAINSPATW, = OTL 0°6 66 Oo! o'6 Lol GL *W Jopun yidap srejnqipueur == Oe) 06 8 a G8 06 GL *d Jopun yidap aepngipuew = 9°61 —_— — — — —_— Og! ssao0id aepnonae jo yoeq 0} *Fw jo yuo. ~~ 966 a = = = = GSS "WO D9 Woy YsuUIT 1ori1asodo19jue = CFI aa rae 20°ST OF GPF irl siejour jo ysuay Ior1a}sodo19jue = 816 a 0°06 = 20°06 0°66 “8ST "W Od Sq YISUDT Io11a}sodo19} ue s]UdWaAINseaUI AZIS [[RIIAO a 8's ae og O'F ae 68 g§ €§ Yi pesiq - oF = 66 0G €F oF VP GF yisua, "IW GP SP 8 2 8g GP P= «x9GP Gis a {peaq SF 6F 6F ae GG LY 0g GF PV yisuzy “WW IF oF OF UF <a a VF g§ og Yi peaiq 0g GP oP oF “as LY LY GF re yisuzy "WW = G¢ = 0's = 9'¢ 8'¢ CSG LG yipeaiq a Sh = OF = UF oF gg gg yisugt az og y L6 = = Ca Go Gs Yipesiq a oF i: 8°¢ = — Sh 0g gg yisuay *d €L6ES $5666 96LE3 96LES LIOIS 81696 61696 WYBIy YT UnUr WdA WdA WdA& WdA WdA W990 W909 244 L P6E93L WNS 2d4.L ur yq293 Jo suaumoads apex masups3 snoayiidvivg 1asuvs3 snaayndvivg sisppaf snaaynidvivg SUSUR OSEIN PARAPITHECUS GRANGERI ‘snoayjidving Ul sJudWAINseaU AvpNGIpueW pue [eJUap aANeIedwoDg *[ ATAVL 6 POSTILLA 166 ular depth, anteroposterior breadth of ascending ramus and length of tooth row from 20 to 30% larger than in type species. Horizontal ramus deepens posteriorly in P. grangeri from P, to M, (see Table 1), while that in type of P. fraasi does not. GENERAL DISCUSSION The type and only specimen of Parapithecus fraasi has long figured in textbooks of anthropology and paleontology as an important basal form with various postulated relationships to later primates. The history of study of this animal, or rather history of misinterpretation of it, is instructive since it clearly demonstrates the problems that arise when there is only one individual fossil specimen (representing a group) and when few of those who wrote of it had bothered to see the actual’specimen itself. Confusion began with the initial description, for at the start Schlosser (1911) drew three wrong conclusions about it. These were: (1) that the type specimen was a juvenile; (2) that the symphysis was unfused; and (3) that the dental formula for it is lis (the same as in modern Tarsius). Although Schlosser recognized that the morphology of the molars and pre- molars of Parapithecus justified placement of the group it represented among the Higher Primates, he concluded that the Parapithecidae must have been an extinct side branch in primate evolution. This was because he was un- able to reconcile the apparent reduction of the lower incisors to one pair with an ancestral relationship to descendant forms that possess two incisor pairs. As I have discussed at length (Simons, 1972) the numerous new jaws of Parapithecus and of the closely related genus Apidium show that the symphysis was fused in members of both genera at a subadult stage of growth; therefore the asymmetrical crack in the symphyseal region of the type of Parapithecus fraasi is an artifact. It is not indicative that the ani- mal possessed an unfused symphsis as Schlosser (1911) initially and Kalin (1961) later believed. Both were misled because of damage there to the type and only specimen. The alveolae and surrounding bone of the central incisor pair, as well as that of the larger lateral incisors, were entirely broken away before Schlosser studied the find, and the two mandibular bodies and the central incisors were then simply glued together. This missing bone also led to an unnatural distortion of the relationships of the horizontal rami of the mandible, for when glued together with this wedge missing they diverge at a much higher angle than would have been the actual case in life. All known Oligocene and Miocene monkeys and apes do have posteriorly divergent horizontal rami, but in this case the divergence is exaggerated. It thus (incorrectly) resembles the high angle of posterior divergence seen in Tarsius. It was this mistaken resemblance in mandibular construction to that of Tarsius, together with the incorrect determination of dental form- ula that impeded understanding of the phyletic relationship of Parapithecus. DE CE 4 TAT RA FS. PARAPITHECUS GRANGERI 4 FIG. 2. Comparison of Parapithecus fraasi, type (A) with adult Parapithecus gran- geri (C) and a juvenile specimen of the latter (B). Scale approximately x 4.9 for both. (A) SMN 12639a, type of P. fraasi, lower dentition lacking lateral incisors and right P,. Scale approximately x 4.9. A is taken from Kalin, 1961. (B) YPM 23796, juvenile P. grangeri right lower dP, ,, and M,_,. Scale approximately x 4.9. (C) YPM 23954, P. grangeri right lower P,-M,. Scale approximately x 4.9. 8 POSTILLA 166 The animal, for these two mistaken reasons, seemed to have affinities with the tarsioids. After description of Parapithecus in 1911, stereophotographs of the oc- clusal views of teeth in the type and only specimen of P. fraasi were dis- tributed. From these crown views it was apparently not possible to see the great disparity in height between the tall tooth Schlosser took to be the canine and that immediately following it. In 1915 Schwalbe incorrectly con- cluded, after examining only a cast, that the dental formula was 2.1.2.3. be- low, as in Old World monkeys and apes. Later Gregory (1922) who was also working from photographs and casts announced in no uncertain terms that Schwalbe’s interpretation of the dental formula had to be correct. But Gregory was wrong, and he even went further to state the incorrect con- clusion that Parapithecus ‘*. . . . may well be regarded as standing in or quite near to the line of ascent leading to the anthropoid apes and eventu- ally to man.’’ He also considered Parapithecus more tarsier-like than Pro- pliopithecus, and concluded that the latter stood ‘‘in or near the base of the gibbon line.’’ Although Gregory thus implied strongly that Parapithecus should be considered a stage typifying the earliest Hominoidea, he noted the overall similarity in the premolars and molars that exists between Para- pithecus and Apidium. He further discussed the similarity between Apid- ium and Oreopithecus which was subsequently dealt with in detail by Si- mons (1960), and concluded that Apidium conformed well with what should be expected for an early Oligocene stage in the evolution of the cercopith- ecoid monkeys. This was apparently the only early recognition of Apidium as related to cercopithecoid monkeys. Remane (1921), writing about the same time as Gregory, suggested that because of the nearly complete re- duction of the paraconid crest in Parapithecus that it might be a primitive representative of the Hylobatidae, but reasoned that because the paraconid crest (still present in the dryopithecines) had already been eliminated, Para- pithecus should be excluded from the ancestry of Pongidae. Much more re- cently Kalin (1961) published a full study of Parapithecus fraasi which appeared only a scant two years before the flow of new material from the Fayum represented by the many Yale expedition primate finds. Kalin also objected to Schlosser’s early interpretation of the dental formula, which had been correct, save for the incisor count. Moreover, he too concluded that the crack in the symphyseal region of the type specimen constituted evi- dence that the animal possessed an open symphysis. Therefore Kalin was just as puzzled about the relationships of Parapithecus as most authors who had written of it previously had been. He reasoned that one could not derive Propliopithecus from a Parapithecus stage as Gregory had implied. In any case such a derivation would be highly suspect because species of the two genera are contemporaries. Kalin decided that Parapithecus was a primate transitional between Higher and Lower Primates: in this, he echoed Schlosser who had based his conclusion on a mistake. In addition Kalin concluded that the form represented such a distinctive group that it should be made the basis of a new superfamily Parapithecoidea. PARAPITHECUS GRANGERI 9 With the discovery of dozens of new specimens of both Parapithecus and Apidium in the Fayum it became clear that the correct dental formula for both genera is phe that both these parapithecids had fused sym- physes and comparatively small canines, but, unlike the similar-sized mar- mosets, had the articular condyle of the mandible situated relatively higher above the level of the tooth row. In a series of papers (Simons, 1967, 1969, 1971) I have pointed out the extraordinary likeness to be found between the molar morphology of the modern African swamp monkey. Miopithecus talapoin, and that of Parapithecus. In Parapithecus the lower molar cusps are already arranged in a quadrate pattern as in Old World Monkeys. More- over, in Parapithecus the paraconid is missing and the hypoconulid is too . reduced to be significant functionally. The much-reduced hypoconulid of Parapithecus is situated in exactly the position where the talapoin pos- sesses a flattened area or shelf, presumably representative of the formerly present hypoconulid. Like monkeys, the unworn molars of Parapithecus are more high-crowned than is the case for the contemporary Fayum dry- opithecines, and the upper molars are much more quadrate in arrangement of the principal cusps than is the case in Apidium or the Fayum apes. It would not be difficult to convert the upper molar of Parapithecus into that seen in the modern cercopithecoids. In this connection I should point out my disagreement with the argument of Von Koenigswald (1969) that the crown morphology of the teeth of Apidium is not relevant to consideration of the origin of the cercopithecoid dentition. The modern cercopithecoid monkeys are very frequently cited as hav- ing (among Primates) remarkably uniform tooth structure, and they pos- sess a standard dental formula as well: abs Any student of mammalian paleontology will be aware that many families of Mammalia include much greater diversity of dental shapes and dental formulae than do the modern Old World monkeys, particularly when a group is known with “‘time-depth”’ as is the case here. This point was well-discussed long ago by Gregory (1920). Moreover, most mammalian families that have an adequate fossil record can be traced back to Eocene times when they include species much more primitive than are any extant members of such families. There- fore, neither the generalized features of some of the parapithecine post- cranial bones, nor the possession of PS seem adequate to me to justify re- tention any longer of K4lin’s superfamily, Parapithecoidea, for these Afri- can Oligocene primates. Most important is the recent study of Conroy (1974) on parapithecid postcranial bones. This shows through morpholog- ical and multivariate analysis that such postcranials as can be confidently assigned to the Parapithecidae are all (in their morphometrics) close to those of various monkeys. In the same fashion Conroy, Schwartz and Sim- ons (1974) have shown that the dental eruption sequence in Apidium, which appears to be the same in Parapithecus, is like monkeys and apes, not like prosimians. For all these reasons there can no longer be any doubt that 10 POSTILLA 166 Parapithecidae are monkeys, not prosimians: zoogeographic considera- tions ally them with cercopithecoids—not ceboids. What one looks for in determining the relatedness of ancestors and de- scendants among fossil mammals is the first emergence of the distinctive or “‘specialized’’ features that later become more exaggerated, or some- times, uniformly typical of the descendant group. Paleontologists will be familiar with a whole series of papers in which the earliest emergent char- acters of a higher category (superfamily, suborder or order) are discerned, for instance, Schaeffer (1947) and Radinsky (1969). In making such a place- ment of Parapithecus close to Old World Monkeys as is advocated here it should not be forgotten that, even if it should prove to be near the an- cestry of modern African monkeys, species of this genus, Parapithecus, are dated at around 28 to 30 million years old. Having existed so long ago, Parapithecus could reasonably be expected to have possessed primitive features that are no longer found in Old World Monkeys. These should not disqualify it from superfamilial association with them, any more than do such features in the basal members of any other group of mammals that evolved during the last two-thirds of the Tertiary. Thus parapithecids can correctly be termed: primitive monkeys. In sum, the loss of the paraconid crest in lower molars of Parapithecus can be taken as a resemblance to monkeys rather than gibbons, and rein- forces the other evidence of marked similarity in molar morphology be- tween Parapithecus, Cercopithecus, and Miopithecus. Research that I have reported elsewhere (Simons, 1967) shows that Apidium is generically dis- tinct from Parapithecus but both have almost identical morphology of the anterior teeth and should therefore be placed in the same subfamily. The recent Yale expeditions to the Fayum badlands of Egypt under the direc- tion of the author and of G. E. Meyer have provided more information as to the craniology of Apidium than is the case for Parapithecus, but it seems highly probable that the two resemble each other in major structural de- tails. Both had fused mandibular symphyses, and in Apidium the metopic suture is fused in early life and postorbital plates develop. Therefore, there seems to be no doubt that these animals had reached at least the grade of organization of the New World Monkeys if not higher. Both had much more foreshortened faces than did their dryopithecine contemporary Aegypto- pithecus, and probably both differed from it slightly in the shape and po- sition of the tympanic. Both should be placed in the same family, Parapith- ecidae. Their ranking among the Cercopithecoidea need in no way imply that the parapithecids would or could have been directly ancestral to any surviving Old World Monkey group but does leave open the possibility that Parapithecus may well prove to have been such an ancestor. This arrange- ment, rather than classifying these genera in a separate superfamily, makes the best sense in the state of present evidence. Working on the assumption that Parapithecus is close to the ancestry of the cercopithecoids, two heretofore little recognized facts emerge. These PARAPITHECUS GRANGER! 11 are firstly, that loss of the second premolars among the ancestors of the cercopithecoid monkeys occurred independently from and later than the similar reduction already found in the earliest apes (Oligopithecus, Pro- pliopithecus, Aegyptopithecus, and Aeolopithecus). Secondly, the small heels of M, in Parapithecus resembling as they do those of Miopithecus and Cercopithecus, strongly indicate the probability that, among Cercopith- ecoidea, enlarged M, heels are a later, or more specialized, development. The cases of independent development of large M, heels among various separate lines of herbivorous mammals are too numerous to require de- tailed tabulation, but they certainly show that various cercopithecoid lin- eages could have separately developed such talonid enlargement at some time subsequent to the Oligocene. SUMMARY A new species of Parapithecus, P. grangeri, is described. The parapith- ecines are the most common mammals of the Fayum Oligocene. They prob- ably did not become extinct thereafter. Their cheek tooth morphology and indeed the morphology of the whole mandible is extraordinarily like that of the smallest African monkey, the swamp monkey. Not many known or strikingly different features separate parapithecids and cercopithecids. To some, Parapithecus, as described here, may still seem so clearly set off from modern monkeys as to require placement outside Cercopithecoidea. The degree of separateness, however, is exaggerated by a lack of interme- diate forms resulting from the very poor paleontological knowledge that we have of Miocene, Pliocene, and early Pleistocene monkeys in Eurasia and Africa. Were intermediate forms better known it would be possible to be much more definite than anyone can now be as to the times and nature of the development of the narrowly limited dental and locomotor systems of the Old World Monkeys. [See Schultz (1970)]. ACKNOWLEDGMENTS The research results reported here were supported in part by grants from the Earth Sciences Section, National Science Foundation (G-18102, GP- 433, GP-3547, GA-723, and GA-11145) as well as by the Smithsonian For- eign Currency Program (Grants 5, 23, and 1841). I also wish to thank David Pilbeam, Glenn Conroy. Friderun Ankel-Simons and Philip Gingerich for discussion and assistance in the preparation of this paper. The field re- search was facilitated by the generous contribution of personnel and equip- ment by the Cairo Geological Museum and the Geological Survey of Egypt. 12 POSTILLA 166 LITERATURE CITED Conroy, G. C. 1974. Primate postcranial remains from the Fayum Province, Egypt, UAR. Ph.D. dissertation unpublished. Yale Univ. Conroy, G. C., J. A. Schwartz and E. L. Simons. 1974. Dental eruption patterns in the earliest Cercopithecoidea. Folia Primatologia. In press. Gregory, W. K. 1920. On the structure and relations of Notharctus, an American Eocene primate. Mem. Amer. Mus. Nat. Hist., N.S. 3(2): 1-193. 1922. The origin and evolution of the human dentition., Williams and Wil- kins, Baltimore, 548 p. ; Kalin, J. N. 1961. Sur les primates de |’Oligocéne inférieur d’Egypte. Ann. Paléont., Paris, 47: 1-48. Koenigswald, G. H. R. von. 1969. Miocene Cercopithecoidea and Oreopithecoidea from the Miocene of East Africa, p. 39-52. In L. S. B. Leakey [ed.] Fossil ver- tebrates of Africa, vol. 1. Radinsky, L. B. 1969. The early evolution of the Perissodactyla. Evolution 23(2): 308-328. Remane, A. 1921. Beitrdge zur Morphologie des Anthropoidengebisses. Arch. Na- turgesch. Abt. A, 87:1-89. Schaeffer, B. 1947. Notes on the origin and function of the artiodactyl tarsus. Amer. Mus. Nat. Hist. Novitates 1356: 1-24. Schlosser, M. 1911. Beitrage zur Kenntnis der oligozanen Landsaugetiere aus dem Fayum, Agypten. Beitr. Palaont. Oesterreich-Ungarns und Orients 24: 51-167. Schultz, A. H. 1970. The comparative uniformity of the Cercopithecoidea, p. 39-51. In J. R. and P. H. Napier [ed.] Old World Monkeys, Academic Press, New York. Schwalbe, G. 1915. Uber den fossilen Affen Oreopithecus bambolii: Zugleich ein Beitrag zur Morphologie der Zahne der Primaten. Ztschr. Morphol. Anthropol. 19: 149-254, Simons, E. L. 1960. Apidium and Oreopithecus. Nature 186: 824-826. 1967. The significance of primate paleontology for anthropological studies. Amer. J. Phys. Anthrop. 27: 307-332. 1969. The origin and adaptive radiation of the Primates. Ann. New York Acad. Sci. 167: 319-331. 1971. A current review of the interrelationships of Oligocene and Miocene Catarrhini, In A. A. Dahlberg [ed.] Dental Morphology and Evolution, Univ. Chicago Press. 1972. Primate evolution: an introduction to Man’s place in nature. Macmil- lan, New York. 322 p. Simons, E. L. and A. E. Wood. 1968. Early Cenozoic mammalian faunas[,] Fayum Province, Egypt. Bull. Peabody Mus., Yale Univ. 28: 1-105. Stoll, N. R., chairman, Editorial Committee, International Commission on Zoolog- ical Nomenclature. 1964. International code of zoological nomenclature. Interna- tional Trust for Zoological Nomenclature, London, England. 176 p. REVIEW STYLE FORM TITLE ABSTRACT NOMENCLATURE ILLUSTRATIONS FOOTNOTES TABLES REFERENCES -AUTHOR’S COPIES PROOF COPYRIGHT INFORMATION FOR AUTHORS The Publications Committee of the Peabody Museum of Natural History reviews and approves manuscripts for publication. Papers will be published in approximately the order in which they are accepted; delays may result if manuscript or illustrations are not in proper form. To facilitate review, the original and one carbon or xerox copy of the typescript and figures should be submitted. The author should keep a copy. Authors of biological papers should follow the CBE Style Manual Third Edition (Amer. Inst. Biol. Sci.). 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