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AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3124, 10 pp., 1 table April 5, 1995

Crania of Apidium: Primitive Anthropoidean (Primates, Parapithecidae) from the Egyptian Oligocene

ELWYN L. SIMONS'

ABSTRACT Cranial remains ofparapithecid primates found anthropoidean cranial characteristics had been de- in Oligocene deposits of the Fayum badlands veloped in the African parapithecids by Oligocene southwest ofCairo, Egypt, are assigned to two spe- times, 34 million years ago. Parapithecoidea may cies of the genus Apidium. A new partial skull be a sister group to Catarrhini. One partial cra- confirms reference of an earlier described frontal nium discussed here belongs to a new species de- bone to the genus and species Apidiumphiomense. scribed below. Together these two finds demonstrate that several

INTRODUCTION In 1959, I published on a primate frontal covered by Markgraf in 1908 in an area 1 or with anthropoidean characteristics that had 2 km to the northwest ofthe quarries named been collected in the Fayum badlands in 1908 A and B and worked in 1906 and 1907 by by Richard Markgraf and sent to the Amer- an AMNH expedition. This would presum- ican Museum of Natural History (AMNH), ably put it in or near quarries I, P, or J. Be- New York, in early 1909 (figs. 1, 2). The exact cause there were no other fragments associ- location site of this fossil frontal, numbered ated with this nearly complete frontal, it seems AMNH 14456, is not indicated on the field that it must have been deposited in river sands label. The attached matrix appeared to be as a separate bone. identical with that on other small fossils re- Although this specimen was early identi-

' Research Associate, Department of Anthropology, American Museum of Natural History; James B. Duke Pro- fessor, Department of Biological Anthropology and Anatomy, Duke University, Durham, NC 27705.

Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $1.60 2 AMERICAN MUSEUM NOVITATES NO. 3124 fied as a "primate," it was not announced to be primate. The specimens of the then scientifically other than in a passing reference apparent association were described by Gin- by Gregory (1922). His sole observation was gerich (1973). Much later Cartmill et al. (1981) that the frontal closely resembles the corre- reviewed additional material that included sponding bone in some of the smaller Old four isolated primate petrosals from quarry World monkeys. When I first noticed the I. These are YPM 25972, YPM 25973, YPM frontal in 1956, its importance seemed clear. 25974, and YPM 23968. The first of these is It gave the first clue that postorbital closure, about 25% larger than the remaining three, a fundamental characteristic of higher pri- which Cartmill et al. (1981) considered to mates, had been achieved by Oligocene times. belong to Apidium phiomense. In reviewing Prior to this frontal's description (Simons, the anatomy ofthe petrosal in the three spec- 1959), there was no evidence, other than imens of Apidium, the latter authors noted through relationships suggested by dental fea- that Gingerich (1973) was correct in con- tures, that any Oligocene primate had reached cluding that the carotid canal lacks a stapedial the anthropoidean grade of organization. branch, resembles Anthropoidea in this re- Reference of this isolated frontal to a par- gard, and differs from early tarsioids as well ticular Fayum primate species was not pos- as most other prosimians, although they re- sible because it is about the size expected for marked that (p. 9): "certain grooves running the type species of two different genera: Api- from the carotid canal across the promon- dium and Parapithecus, Simons (1959). Since tory's ventral surface may have contained then, a series of expeditions to the Fayum caroticotympanic arteries ... one of which badlands originating first from Yale (between may represent a vestige ofthe stapedial stem." 1961 and 1968) and then from Duke (from The detailed discussion of Cartmill et al. 1977 to date) has resulted in the discovery (1981) emphasizes that the anatomy of sev- of hundreds of mandibles, maxillae, and po- eral isolated petrosals is anthropoid-like and stcranials of small primates, the commonest does not reveal any particular resemblance of which is Apidium phiomense, Osbom to the tarsiero-momyid group. They also show (1908). The type specimen ofA. phiomense, that the association ofApidium material cata- a mandible with a fourth premolar and three loged under the number YPM 23968 con- molars was, like the frontal, also found by tained a squamosal that must belong to a Markgraf but somewhat earlier. The original nonprimate, that had erroneously been la- field label of this type said: "NW of Quarry beled as part of Apidium. Because of this

[A] ... new genus ?primate, Feb. 17, 1907, squamosal not being primate, the conclusion collr. R. Markgraf." of Gingerich (1973) that Apidium had an ec- Apidium phiomense occurs (principally) in totympanic that was "free and intrabullar" the upper sequence ofthe Fayum deposits at now lacks proof. If the isolated petrosals in- quarries I and M, and quarry I lies in a NW deed are of Apidium then the ectotympanic direction from quarry A. Over the years, sev- is a simple annulus like that of ceboids and eral other isolated small primate frontals re- of Aegyptopithecus, fused to the squamosal sembling that described in 1959 have been at both ends. discovered in quarry I. The first ofthese was Although Apidium phiomense is the com- reported, Simons (1967), as probably being monest small mammal in the upper sequence referable to A. phiomense. Simons subse- ofthe Jebel Qatrani Formation, Parapithecus quently (1971) described another frontal fraasi, another similarly sized primate spe- fragment found associated with separate teeth cies described from the Fayum early in this of A. phiomense. This find appeared to in- century by Schlosser (1911), has continued dicate, with some certainty, that the AMNH to be known only from the type specimen. frontal and other small frontals subsequently Meanwhile, Simons (1974) proposed a sec- recovered at quarry I belonged to A. phio- ond species of the genus, Parapithecus gran- mense. Even so, a temporal fragment found geri, and this species occurs with A. phio- at the same spot with the upper teeth and the mense at quarries I and M. Although their piece of frontal bone from the region of the molar structure is quite different, certain sim- interorbital septum, subsequently proved not ilarities in the dentition, such as, among an- 1995 SIMONS: CRANIA OF APIDIUM thropoideans, a unique central cusp on the upper premolars, show that Apidium and Parapithecus belong in the same family. In comparable parts, Parapithecus grangeri is about 20% larger than A. phiomense and, consequently, would have a frontal distinctly larger than that described in 1959 (AMNH 14456). No specimen ofParapithecusfraasi has ever been found in the upper sequence localities of the Fayum and therefore the type surely came from a different, presumably lower, part of the section. Because it is both larger than would be expected for P. fraasi and appar- ently younger, the frontal described in 1959 could not belong to the type species of Para- pithecus. The only other common larger primates at the quarry I-M level are Parapithe- 0 1 2 cus grangeri, Aegyptopithecus zeuxis, and L,.1ll.LLLLLLLL llJ Propliopithecus chirobates, all of which have frontals larger than that ofAMNH 14456. At quarry M there is an additional small parapithecid-known from a single mandible- Fig. 1. Cranial remains ofApidiumphiomense, Qatraniafleaglei, but this animal is too small dorsal view. Left, Duke specimen, DPC 9867; right, to relate to this frontal. Eliminating all these American Museum specimen, AMNH 14556. other species that differed in size and/or level ofoccurrence strengthens, but does not prove, the conclusion that AMNH 14456 belongs to of attached partial parietals that on grounds A. phiomense. of size are assignable to P. grangeri. These In 1989, after 27 years ofcollecting at quar- new specimens are described below. ry I, a partial cranium ofApidium phiomense, DUPC 9867, with five attached upper molars ACKNOWLEDGMENTS was discovered (figs. 1, 2). This specimen has I thank the frontal largely intact and it definitely es- Friderun Ankel-Simons and Tho- tablishes that the frontal discovered by Mark- mas M. Bown for helpful criticism during AMNH preparation of the manuscript together with graf, 14456, indeed belongs to Api- H. H. Covert and P. D. Gingerich for re- dium phiomense (see fig. 1). Regrettably, viewer's comments. This research was sup- DUPC 9867 does not preserve the petrosal ported by several NSF grants in Anthropol- on either side and, consequently, the earlier the found petrosals ofApidium must still be re- ogy, most recent of which are BNS-85- ferred only provisionally. 46024, BNS-88-09776, and BNS-91-08445. In 1984 at about 78 m Abbreviations: YPM = Yale Peabody Mu- quarry V, below the seum; AMNH = American Museum ofNat- I/M level, Dr. Thomas Bown of the U.S. = Geological Survey, Denver, quarried out an ural History; CGM Cairo Geological Mu- associated frontal, partial face, and upper seum; DUPC = Duke University Primate dentition of a new small species of Apidium Center. Photographic illustrations have been (DPC 5264). During the 1980s, cranial parts prepared by R. L. Usery, Duke Audiovisual of Parapithecus grangeri were also being Services, and by the author. This is Duke found. At quarry I, in 1986, an associated left Primate Center Publication No. 550. maxilla and frontal of Parapithecus grangeri (DPC 6641) were collected. Earlier at I in DESCRIPTION 1978, we had also recovered another perti- The partial cranium, DPC 9867, is slight- nent specimen (DPC 1098), which consists ly larger than that ofthe common marmoset, 4 AMERICAN MUSEUM NOVITATES NO. 3124

Callithrixjacchus, and is somewhat more than the eye socket is preserved. The bones have 1 /2 times the size typical ofthe mouse lemur, been crushed together in such a way that the Microcebus murinus. Anteroposteriorly, the exact size and position ofthe orbitotemporal new frontal is no longer than it usually is in opening cannot be determined. However, on the owl monkey, Aotus trivirgatus, but the the obital floor, side, and back, portions of parietals are much less broad than in the lat- the maxillary, zygomatic, and frontal plates ter. Consequently, the brain volume of Api- can be seen enclosing the eye socket. The dium phiomense is considerably smaller than medial wall ofthe orbit, with a large lacrymal that of A. trivirgatus. In the palate of DPC foramen, is the least distorted part of the or- 9867 upper teeth are preserved that confirm bit. On the dorsal surface of the skull, the the assignment ofthis skull to Apidium phio- temporal crests, or lines, run backward from mense. The teeth that remain in this skull the external angular processes and converge include the left M'-3 and the somewhat dam- in the midline at the fronto-parietal suture. aged right Ml-2. The absolute size of these Back of this, the interparietal suture can be molars is considerably larger than in Aotus followed posteriorly to the inion. The original or Saimiri, extant primates that clearly have frontal, AMNH 14556, exhibits a ledge of larger brain volumes. The molar size in this bone, particularly on the left side, that would fossil is closer to those of Callicebus torqua- have extended back under the parietal. In this tus. As with Aotus and Saimiri, the brain vol- region of the skull in platyrrhines, there is a ume of Callicebus is much greater than that frontal/alisphenoid contact, whereas in cata- estimated for Apidium phiomense. rrhines the parietal has a sutural contact here In its present condition, DPC 9867 is 5.3 with the jugal. Fleagle and Kay (1988) sug- cm long from the inion to the anteriormost gested that this posterior ledge, or extension remaining part ofthe rostrum. The complete ofthe frontal, may have achieved such a plat- cranium would not have projected farther yrrhine-like sutural contact with the ali- posteriorly and, because the face is somewhat sphenoid. In DPC 9867, the right parietal is distorted and flattened dorsoventrally, the shifted forward onto the zygomatic area in rostrum also was probably not much longer such a way as to leave this possibility equiv- in life, with a total anteroposterior length es- ocal. The same situation obtains for AMNH timated to be from 5.5 to 5.8 cm. Relying 14556; consequently at present this issue can- particularly on AMNH 14556, which is more not be resolved. complete in the orbital region, an estimate of Examination of the basicranium of DPC 2.75 cm for the maximum breadth across the 9867 (fig. 2) indicates that the posterior mar- orbits is plausible. It is clear from these de- gin of the hard palate probably extended terminations that Apidium phiomense had a somewhat backward beyond a transverse line brain volume that was only about 60 to 75% across the posterior sides of the third molars the volume of that in a modern platyrrhine and graded off into the pterygoid wings. The such as Callicebus torquatus with teeth of basicranium is considerably damaged, but on similar size. This partial cranium shows, as the right side the glenoid fossa and the part does the frontal, AMNH 14556, that the dor- ofthe squamosal at the back ofthe zygomatic sal margin of the orbit is relatively flat with arch are still preserved. Unfortunately, there the rather angular dorsomesial corner ofeach is almost no detail left in what fragments re- orbital opening adjacent to the interorbital main of the right auditory region. This cra- septum and is structured in outline somewhat nium (in overall view) is rather elongate com- like the squared-off orbits of modern Calli- pared to its breadth and, in this regard, is cebus. DPC 9867 is important as well in con- reminiscent of crania of Saimiri and Calli- firming that parapithecids had full postor- thrix, in contrast to skull shape in most other bital closure, a fact also indicated by the living monkeys. This difference may only arise postorbital plate ofthe frontal, AMNH 1456, because all Miocene-Recent anthropoideans although this was first reported when the lat- have distinctly larger brains, compared to size ter's familial affinities were not known. Even of teeth and overall body size. though part ofthe zygomatic rim ofthe right From quarry I in the Fayum upper se- orbit is broken away, much of the interior of quence there is a partial right maxilla and 1995 SIMONS: CRANIA OF APIDIUM 5 face, DPC 6641, of Parapithecus grangeri, a somewhat larger relative ofApidium. As stat- ed above, this specimen was directly associated with a frontal of comparable size. As mentioned from quarry I, there is also a pair of parietals that appear to belong to Parapi- thecus grangeri. Although not from the same individual as the frontal DPC 6641, these parietals, DPC 1098, articulate well with it. The internal surface of the braincase in the skull DPC 9867 is almost featureless. How- ever, these latter two specimens show enough of the modeling of the dorsal surface of the brain to indicate some resemblance to that in Aegyptopithecus zeuxis. Placing the parietals together with a relatively complete man- ;qqhff 0 1 2 dible of Parapithecus grangeri makes it pos- sible to prepare a composite reconstruction of the skull of this strangely adapted parapithecid, which had a greatly foreshortened face (fig. 3). The original frontal of A. phiomense (AMNH 14556) can now be combined (similarly) with a right maxilla (CGM 26929) Fig. 2. Ventral view ofpartial skull ofApidium and mandible (YPM 21018) in order to par- phiomense. Left, DPC 9867; right, frontal ofsame tially reconstruct the splanchnocranium of species, AMNH 14556. Apidium phiomense (fig. 3). From these re- constructions it is clear that in both contem- poraneous species from quarries I, M, and J there existed foreshortened faces, rather rect- contact with the frontal and, on its internal angular orbital openings, and dorsally con- surface a faint imprint of the dorsal surface verging temporal lines. The left maxilla of of the brain is preserved. DPC 6641, P. grangeri, has a conjoined zy- Under the orbital opening ofanother Para- gomatic bone, which contains a noticeably pithecus grangeri, DPC 6641, there is a dis- large zygomatic foramen. The large size of tinct infraorbital foramen. The lachrymal this foramen is a primitive platyrrhine or pro- bone and lachrymal foramen are both situ- simianlike feature and this foramen in P. ated inside the orbit, a feature characteristic grangeri is comparatively much larger than of higher primates. In this specimen the zy- in various Fayum specimens of Aegyptopi- gomatic arch rises relatively far forward, just thecus (family Propliopithecidae). The pari- above a line between first and second molars. etals here assigned to Parapithecus grangeri, The ascending external face of the maxilla is DPC 1098, are too small to be from the two deep and rises medially to the point ofcontact propliopithecids, Aegyptopithecus zeuxis and with the nasals. There is no distinct canine Propliopithecus chirobates, that are its con- fossa. Viewed from below, the palate extends temporaries at quarries I and M. Hence it medially from the root sockets of the upper seems reasonable to associate them with P. p2-3 to a well-developed midline suture. As grangeri. As arranged in figure 3, parietal ar- far as presentation of the specimen allows, chitecture suggests a relatively long, low cra- this shows that anteriorly the upper tooth nial vault similar to that preserved in Api- rows converge markedly and are separated dium phiomense (DPC 9867, cf. figs. 1, 3). by a reduced transverse space, other evidence The parietals of P. grangeri (DPC 1098) are of a very foreshortened face. On the associ- solidly fused along the midline but are both ated frontal fragment, the metopic suture is broken off irregularly at their lateral, ventral fully fused but the temporal lines have not margins. Situated anteriorly is the suture for yet quitejoined where they pass offthe frontal 6 AMERICAN MUSEUM NOVITATES NO. 3124

Fig. 3. Left, three-quarter view of Apidium phiomense. Composite reconstruction of skull: frontal, AMNH 14556; right maxilla, CGM 26929; mandible, YPM 21018; canine and upper incisors isolated finds. Right, lateral view ofskull ofParapithecus grangeri. Composite reconstruction ofskull: left anterior maxilla with canine, P34, DPC 2385; posterior maxilla and frontal, DPC 6641; parietals, DPC 1098; and mandible, DPC 2807. posteriorly. Parapithecus grangeri has a 21018, figure 3, these larger lateral incisors strange adaptation, not seen in Apidium, in have almost always fallen out. This is because that all lower incisors have been lost, and the the lateral incisors have straight, conical roots lower canines are the anteriormost pair of which do not hold the teeth in their sockets. teeth. The canines are closely appressed with Isolated teeth of Apidium phiomense are so wear facets on their contacting mesial faces, common at quarry I that many upper incisors proving unequivocally that the lower incisors of this species have been identified. These are gone. This adaptation is unique among teeth demonstrate that, as is typical of an- primates (see Simons, 1986), and it is not thropoideans, the lateral incisor pair is dis- certain whether or not upper incisors existed tinctly smaller than the central. All these fea- in P. grangeri, as no premaxillae are known. tures combine to produce a rostrum in A. However, occluding the upper and lower den- phiomense that is less abbreviated than in P. titions indicates that there was rather restrict- grangeri, and one that therefore looks similar ed space for upper incisors between the large, to the snout of a typical marmoset. blunt upper canines, a condition also sug- Finally, figures 4 and 5 show the partial gested by the very narrow anterior palate. splanchnocranium of Apidium from quarry Because oflower incisor loss (and, at the very V (DPC 5264). This find belongs to a new least, incisor reduction above) the front of species ofApidium described below. This dis- the face in Parapithecus grangeri is foreshort- covery provides little additional anatomical ened, blunt and, seen from the sides, has an knowledge ofthe cranium ofApidium. It does almost parrotlike, rounded aspect in profile. show that the posterior margins of the two This conformation is quite unlike the con- nasal bones form an M like sutural outline, dition seen in the related species Apidium a condition that closely resembles the struc- phiomense where incisor development is like ture of this region in some specimens ofAe- that typical of all other Anthropoidea. gyptopithecus zeuxis. DPC 5264 also recon- Apidium also shows another basal anthro- firms the central fusion ofthe metopic suture poidean character: the central lower incisor originally reported by Simons (1959) for pair is smaller than the lateral. As in YPM AMNH 14556. 1995 SIMONS: CRANIA OF APIDIUM 7

Fig. 5. View showing partial palate with upper dentition ofApidium bowni, DPC 5264. Specimen preserves: left canine C, M3, right P2-4, and M2-3.

Fig. 4. Frontal view of Apidium bowni; pre- open lingually without a premetacristid and serving partial maxillae, nasals, and frontal bone, where buccal cingulum and cetroconid are DPC 5264. invariably lacking. Differs from Serapia which has P2 that is larger than P3-4 in having P24 series that increases in size posteriorly. SYSTEMATICS Apidium bowni, new species ORDER PRIMATES LINNAEUS, 1758 Figure 4 SUBORDER ANTHROPOIDEA MIVART, 1864 TYPE SPECIMEN: CGM 42199 (DPC 8921), SUPERFAMILY PARAPITHECOIDEA, KALIN, 1961 a right mandibular corpus with root sockets FAMILY PARAPITHECIDAE, SCHLOSSER, 1911 of I112, C1, and P2, and P3-M3 complete. Genus Apidium Osborn, 1908 HYPODIGM: Type and DPC 2958, 3884, 5406, 5411, and 6282; mandibular fragments REVISED GENERIC DIAGNOSIS: Apidium dif- with teeth; DPC 6295, right maxilla with C- fers from other parapithecids in having pre- M3 and DPC 5264 splanchnocranium with molars and molars that are more cuspidate upper dentition lacking incisors, right canine, than in other genera in the family-Serapia, and right M1. For comparative measure- Qatrania, and Parapithecus. Cheek teeth with ments see Appendix 1. inflated cusps; large hypocone, displacing the protocone somewhat labially toward the LOCALITY: Fossil vertebrate quarry V, at paracone and metacone; labial cingulum cus- the 165 m level, Jebel Qatrani Formation pidate. Lower molars typically increasing in (lower Oligocene), Fayum Province, Egypt. size posteriorly or with the M2 and M3 sub- Considerably older than the basalt overlying equal. Unworn lower molars with distinct this formation 31 ± 1 Ma [Fleagle et al., centroconid in middle of the central basin. 1986]. Recent dates for the basalt overlying Hypoconulid of M3 is normally flanked by the Jebel Qatrani Formation reported in many small cuspules. Parapithecus differs Kappelman et al., 1992, that are younger than from all other parapithecids in lacking cus- 31 ± 1 are not based on samples from the pidation on tooth crowns and in having mo- lowest Jebel Qatrani basalt (contra Kappel- lars subequal in size, particularly M2.3. Dif- man et al., 1992: 653) and hence do not in- fers from Qatrania in lacking large trigonid validate it. 8 AMERICAN MUSEUM NOVITATES NO. 3124

DIAGNOsIs: Apidium bowni resembles Ap- known parapithecid species, all of which are idium phiomense and A. moustafai and dif- mandibles. The upper dentition ofDPC 5264 fers from species of Parapithecus and Qat- is more dentally complete and its choice as rania in having distinct, well-developed a type was considered but rejected because centroconids on the lower molars. Differs distinctions between upper dentitions have from Qatrania in having relatively larger M2 never been drawn for parapithecids and, in and M3 compared to M1. Differs from A. fact, upper dentitions are either not known, phiomense and more nearly resembles A. or incompletely known in five of the eight moustafai in that M2 is similar in length to parapithecid species, namely in Parapithecus M3, rather than M3 being distinctly longer fraasi, Qatrania wingi, Qatraniafleaglei, Ser- than M2 as is typical ofA. phiomense. Differs apia eocaena, and Apidium moustaji. from the other two species ofApidium in its The type specimen shows detailed anato- much smaller size, being a third to a quarter my of the five posterior teeth as well as the smaller than average A. moustafai from quar- root sockets ofthe right side. The right man- ry G, as well as from several specimens of dible seems to have separated, in this case, Apidium (cf. A. moustafai) also found at quar- at or about the midline suture. Numerous ry V. other finds of Apidium show that the man- ETYMOLOGY: Named for Thomas M. Bown dibular suture is fused even in subadult in- in dividuals. CGM 26919 shows clearly the who found one of the specimens, recog- comparatively large size of M2-3 relative to nition of his significant additions to our Ml that typifies this species. The molar cen- knowledge of the geology, paleontology, pa- troconids, although distinctly discernible, are leoecology, and biostratigraphy of the Fa- not as clearly defined as in DPC 3884, DPC yum. 5406, DPC 541 1, and DPC 6282, probably DEscRWrIoN: The type specimen, CGM because the type has suffered some abrasion 26919 (= DPC 8921), a right mandible, is to the teeth, and may also be of a somewhat the most complete known lower dentition and older individual with more advanced tooth is comparable with the types of all other wear.

DISCUSSION AND CONCLUSIONS The parapithecids discussed here are a mens of Apidium or any other parapithecid group of anthropoideans that in Africa had yet known definitely establish the construc- reached the "monkey" grade of organization tion and placement of the ectotympanic. Si- with postorbital closure and metopic and mons and Kay (1988) and Fleagle and Kay symphyseal fusion. The arrangement ofbones (1988) have discussed in detail the status of at the back of the jugal may be closer to that knowledge of the relationships of parapithe- of Platyrrhini than of Catarrhini. No speci- cids to other primates.

REFERENCES Cartmill, M., R. D. E. MacPhee, and E. L. Simons dentition. Baltimore: Williams and Wil- 1981. Anatomy of the temporal bone in early kins, 548 pp. anthropoids, with remarks on the prob- Fleagle, J. G., T. M. Bown, J. D. Obardovich, and lem ofanthropoid origins. Am. J. Phys. E. L. Simons Anthropol. 56: 3-21. 1986. Age ofthe earliest African anthropoids. Gingerich, P. D. Science 234: 1247-1249. 1973. Anatomy of the temporal bone in the Fleagle, J. G., and R. F. Kay Oligocene anthropoid Apidium and the 1987. The phyletic position of the Parapithe- origin ofAnthropoidea. Folia Primatol. cidae (Primates, Anthropoidea). J. Hum. 19: 329-337. Evol. 16: 483-531. Gregory, W. K. Kappelman, J., E. L. Simons, and C. C. Swisher, 1922. The origin and evolution of the human III 1995 SIMONS: CRANIA OF APIDIUM 9

1992. New age determinations for the Eocene- 1971. A current review of the interrelation- Oligocene boundary sediments in the ships of Oligocene and Miocene Cata- Fayum Depression, northern Egypt. J. rrhini. In A. A. Dahlberg (ed.), Dental Geol. 100: 647-668. morphology and evolution, pp. 193-208. Osborn, H. F. Chicaco: Univ. Chicago Press. 1908. New fossil mammals from the Fayum 1974. Parapithecus grangeri (Parapithecidae, Oligocene, Egypt. Bull. Am. Mus. Nat. Old World higher Primates): new spe- Hist. 24: 265-272. cies from the Oligocene of Egypt and Schlosser, M. the initial differentiation ofCercopithe- 1911. Beitriige zur Kenntnis der Oligoziinen coidea. Postilla 166: 1-12. Landsiiugetiere aus dem Fayum. Beitr. 1986. Parapithecus grangeri of the African Palaeontol. Geol. Oesterr.-Ung. 24: 51- Oligocene: an archaic catarrhine with- 167. out lower incisors. J. Hum. Evol. 15: Simons, E. L. 205-213. 1959. An anthropoid frontal bone from the Simons, E. L., and R. F. Kay Fayum Oligocene of Egypt: oldest skull 1988. New material of Quatrania from Egypt fragment ofa higher Primate. Am. Mus. with comments on the phylogenetic po- Novitates 1976: 16 pp. sition of the Parapithecidae (Primates, 1967. Review of the phyletic interrelation- Anthropoidea). Am. J. Primatol. 15: ships of Oligocene and Miocene Old 337-347. World Anthropoidea. Coll. Int. Cent. Nat. Rech. Sci. 63: 597-602.

APPENDIX 1 Comparison of Representative Apidium phiomense, and A. moustafai with A. bowni Measurements (mm) Length Breadth Trigonid Talonid DPC A. moustafai 10705 C 3.22 P2 2.85 2.18 P3 3.03 2.15 P4 2.79 2.51 MI 3.39 2.83 2.97 M2 3.42 3.12 3.10 M3 3.4e 3.00 A. moustafai 8707 P3 2.4 1.78 P4 2.86 2.09 M2 3.68 2.71 2.66 M3 3.61 2.52 2.07 A. moustafai 3830 P4 2.69 2.40ea MI 3.18 2.50e 2.50e M2 3.57 2.80 2.83 M3 4.10 2.90 2.69 A. moustafai 5670 P4 2.44 2.37 MI 2.54 M2 3.78 2.97 3.08 A. bowni 3884 P4 2.07 1.78 MI 2.82 2.18 2.36 M2 2.87 2.47 2.34 M3 2.87 2.28 1.75 A. bowni 5406 M2 3.07 2.33 2.12 M3 2.96 2.12 1.95 A. bowni 2958 C 2.00 1.80 P2 1.75e 1.35 10 AMERICAN MUSEUM NOVITATES NO. 3124

APPENDIX 1-(Continued) Measurements (mm) Length Breadth Trigonid Talonid P3 1.50e 1.55e P4 2.10e 1.70e A. bowni 5411 M2 2.97 2.30e 2.26 M3 2.70 2.25 1.85 A. bowni 8921 P3 1.95 1.62 P4 2.20 1.94 MI 2.68 2.10 2.24 M2 2.82 2.46 2.43 M3 3.02 2.36 2.1 A. bowni 6282 P3 2.2 1.45 P4 2.12 1.73 MI 2.81 1.85 2.20 M2 3.12 2.65 2.50 CGM A. bowni 26919b P3 1.97 1.58 P4 2.07 1.92 M I 2.66 2.12 2.23 M2 2.69 2.44 2.35 M3 3.03 2.35 1.85 YPM A. phiomense 20905 P3 2.64 2.06 P4 2.65 2.20 M I 3.51 2.60 2.64 M2 3.45 3.0 2.85 M3 3.45 2.89 2.54 A. phiomense 20914 P3 2.48 2.0 M I 3.26 2.52 2.42 M2 3.25 2.71 2.46 M3 3.13 2.52 2.11 A. phiomense 20911 P2 2.32 2.48 P3 2.58 2.32 P4 2.81 2.52 M I 3.52 2.8 2.75 M2 3.54 3.09 3.15 A. phiomense 20920 P3 2.59 2.05 P4 2.74 2.37 M I 3.73 2.85 2.89 M2 3.8 3.24 3.0 M3 3.75 3.03 2.51 20911 P2 2.32 2.48 P3 2.58 2.32 P4 2.81 2.52 M I 3.52 2.8 2.75 M2 3.54 3.09 3.15 a e = estimate. b Type was DPC 8921.

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