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Discovery of the Smallest Fayum Egyptian Primates (Anchomomyini, Adapidae)

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Discovery of the Smallest Fayum Egyptian Primates (Anchomomyini, Adapidae) Proc. Natl. Acad. Sci. USA Vol. 94, pp. 180–184, January 1997 Evolution Discovery of the smallest Fayum Egyptian primates (Anchomomyini, Adapidae) ELWYN L. SIMONS Department of Biological Anthropology and Anatomy and Primate Center, Duke University, Durham, NC 27705-5000 Contributed by Elwyn L. Simons, October 10, 1996 ABSTRACT Two new adapiform primate species from subfamily Ceracmoniinae of the Adapidae (as here) or it may locality 41, Jebel Qatrani Formation, Egypt, are described. prove to be a separate subfamily of its own. The first, here named Wadilemur elegans genus novum species Most of the newly described species from sites in the novum (holotype Cairo Geological Museum 42211), consists Afro-Arabian plate are discussed variously by those who of a right mandible with P3–M3. The second is assigned to named them as being either adapoid or anthropoidean or, in Anchomomys milleri species novum, with a holotype Cairo one case, basal omomyid. Nevertheless, with the exception of Geological Museum 42842, that includes the right mandible Djebelemur, which consists of a five-toothed, mandibular frag- with lower canine to M . Both species are allied closely with ment, the classification of these new Afro-Arabian primates is 3 based on isolated teeth, mostly from the lower dentition. The genera that are known to be from Eocene deposits either in newly described Paleogene African prosimians mentioned Europe, Tunisia, or the Sultanate of Oman (Arabia), thus above join an enigmatic, earlier Algerian find, Azibius trerki, enhancing earlier paleomagnetic evidence that locality 41 was reported by Sudre (16). In all cases, prosimians found outside deposited in Eocene times. the Fayum lack the critical parts of the cranial anatomy that would confirm definite ranking, whether with adapoids or with Collections made in the last 5 or 6 years have yielded more new anthropoideans (3, 17). information about early anthropoidean evolution than finds of The diversity of primates at L-41 rivals any site in the upper any previous decade of the century, and new descriptions have sequence of the Fayum, for example Quarry M, where six greatly added to knowledge of Paleogene prosimians in Africa. primate species have been found. The new genus and species Most of this information has come from the Fayum, Egypt (1–5). here described as well as the new species of Anchomomys There have been other finds further west in North Africa and in makes eight primate genera so far described from L-41. the Sultanate of Oman, Arabian peninsula—a biogeographic part Together, these constitute a distinct, much earlier, and differ- ent Fayum lower-sequence fauna in which both the prosimians of Africa (6). Godinot and Mahboubi (7–9) also described new and probably the anthropoideans represent more diversified primate genera and species from Algeria thought to be earliest primate groups than occur in the Fayum upper sequence. anthropoideans. Together with their descriptions, these authors The age of Fayum site recently was reviewed by Kappleman clearly disallowed any affinity between the new Eocene anthro- et al. (18) and Rasmussen et al. (19). In brief, the upper- poideans or ‘‘near anthropoideans’’ from Africa and the Chinese sequence quarries appear to be about 33 million years ago Eosimias (10)—a point also stressed by Simons (11). Djebelemur (Ma) and the L-41 level about 36 Ma, the latter being from Tunesia was figured and analyzed by Hartenberger and correlated as Priabonian late Eocene. Marandat (12). They considered this find to be an adapid, but Both of the new species described below are clearly related others thought it may be an anthropoidean (9). Lastly, Ducrocq to the Djebelemur–Anchomomys–Omanodon–Schizarodon et al. (13) described a new possible anthropoidean relative, group. It seems, on the other hand, that the idea that the latter Wailekia, from the Paleogene of Thailand. two of these genera are distinct from Anchomomys is not The transition between prosimians and anthropoideans is correct. However, it also seems that the described material of being narrowed by these new discoveries. There also have been Schizarodon is inadequate to sustain a valid generic diagnosis. striking advances in understanding other early Tertiary pros- As has been noticed by Gheerbrant et al. (13), Schizarodon, imian groups found in the African zoogeographic region. One Omanodon, and by inference, Anchomomys and Djebelemur group of these prosimians, the cercamoniine Adapidae, was bear a strong resemblance to the cheirogaleid lemur Microce- bus, as can be seen in Fig. 1 by comparison with a stereopho- first reported from Oman, Arabia, by Gheerbrant et al. (14). tograph of a Microcebus right mandible. Although the molar One year later, Simons and Rasmussen (15) described man- resemblances are striking, at least one of the species described dibles and a remarkable cranium of a rather large prosimian, below, A. milleri, which preserves the canine, shows that these Plesiopithecus teras, from the Fayum late Eocene locality 41 Paleogene prosimians did not have a tooth comb. The origin (L-41). Subsequently, Simons et al. (5) reported on an even of the tooth–comb prosimians is still a mystery. larger new genus and species of cercamoniine also from the late Eocene Fayum quarry L-41, Aframonius dieides. In this paper, another new genus and species of a diminutive adapoid SYSTEMATICS found in L-41 is reported, Wadilemur elegans, as well as a new Order Primates Linnaeus, 1758; Suborder Prosimii, Illiger, species of Anchomomys, Anchomomys milleri. All of these 1811; Family Adapidae, Trouessart, 1879; Family fossils represent an important addition to understanding the Cercamoniinae, Gingerich 1977; Tribe Anchomomyini radiations that may have produced the Anthropoidea. The Szalay and Delson, 1979 (sensu Godinot 1988). Anchomomys group may be considered either a tribe under the Wadilemur, New Genus The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked ‘‘advertisement’’ in Generic diagnosis. Distinctly larger than Omanodon and accordance with 18 U.S.C. §1734 solely to indicate this fact. slightly larger than Shizarodon, Waldilemur shows typical Copyright q 1997 by THE NATIONAL ACADEMY OF SCIENCES OF THE USA 0027-8424y97y94180-5$2.00y0 Abbreviations: L-41, locality 41; Ma, million years ago; CGM, Cairo PNAS is available online at http:yywww.pnas.org. Geological Museum; DPC, Duke University Primate Center. 180 Downloaded by guest on October 3, 2021 Evolution: Simons Proc. Natl. Acad. Sci. USA 94 (1997) 181 Djebelemur in having a more mesiodistally compressed trigo- nid, a comparatively larger M2 relative to M1, a relatively smaller M3 talonid basin, and a more distinctly set off hypo- conulid (Fig. 2). In addition, the lower molars of Wadilemur are more ‘‘notched’’ externally between the talonid and trigonid, especially in M2-3, whereas in Djebelemur, the lateral or buccal outline of these teeth is smoothly rounded. Type species. W. elegans. Distribution. The Fayum, Egypt, quarry L-41. Etymology. From Arabic Wadi, meaning a valley or wash, plus Lemur/Lemures L. W. elegans, New Species Holotype. CGM specimen 42211, right mandible with P3–M3. Hypodigm. Type and DPC 13439 left dentary with P4–M3 and partial ascending ramus. Species diagnosis. Same as generic diagnosis. Etymology. From Latin elegans, meaning neat, elegant. Type species. W. elegans. Anchomomys, Stehlin 1916 Type species. Anchomomys gaillardi. Distribution. Middle Eocene site at Lissieu, France. Etymology. From Greek anchi, meaning near, plus Omomys, name of a North American Eocene primate. This name is from Greek omos, meaning shoulder (many other meanings) com- bined with Greek mys, -mouse. FIG. 1. Stereopair of right dentary of a mouse lemur (Microcebus murinus). Duke University Primate Center (DPC) osteo.035, crown view. (311.) Note caniniform P2 molars with broad talonids and mesially extended entoconids. adapoid lower molars with crescentic ridges connecting the cusps. Lower first and second molars are subequal in size and length, with M3 longer and showing central hypoconulid—no hypoconulid on M1-2. Lower molars differ from those of O. minor in that the cristid obliqua does not run up to join the metaconid cusp crown as in both Omanodon and Shizarodon; also, the entoconid of O. minor is shifted further lingually than in Wadilemur, and the high point of the paraconid crest (presumably the actual paraconid) in Omanodon is shifted more laterally than in Wadilemur. Also, in Wadilemur,a continuous crest or ridge runs from the tip of the protoconid around to the metaconid encircling the anterior fovea without a break; in Omanodon, there is a deep, central notch in this crest. Wadilemur differs from Shizarodon in being slightly larger, with lower molar metaconid shifted further anteriorly and with a paraconid cusp located more posteriorly in relation to the protoconid so that the trigonid is less open anterolat- erally. In both D. martinezi and Algeripithecus minutus, the cristid obliqua of M1 runs up into the apex of the metaconid whereas in Wadilemur, this cristid runs into the middle of the wall between the protoconid and metaconid. Algeripithecus also shows a more crenulate talonid with possible traces of a twinned entoconid—characteristics not seen in Wadilemur. Close to Djebelemur but much larger and apparently consid- erably younger, Wadilemur has lower molars of subequal length that narrow posteriorly whereas, in contrast, Djebelemur has a larger M1 and a smaller M3 compared with M2.IntheM1and M2 of Djebelemur, the crest running posteriorly from the hypocone is relatively higher above the basin and has a slightly FIG. 2. Stereopair of W. elegans type CGM Cairo Geological developed cuspule, which may be the analog of the M3 Museum (CGM) 42211. (311.) Right mandibular fragment with hypoconulid, whereas in Wadilemur, this crest is lower and less P3–M3. Note that the large and lingually directed entoconids are well defined and has no cuspule. The latter also differs from broken off on both M1 and M2.
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