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 pri- mates 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 para- pithecid-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.