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Geographical and Body Size Distributions of the Pondaung Primates with a Comment on the Taxonomic Assignment of NMMP 20, Postcra

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Geographical and Body Size Distributions of the Pondaung Primates with a Comment on the Taxonomic Assignment of NMMP 20, Postcra ANTHROPOLOGICAL SCIENCE Vol. 112, 67-74, 2004 Geographical and body size distributions of the Pondaung primates with a comment on the taxonomic assignment of NMMP 20, postcranium of an amphipithecid NAOKO EGI1*, SOE THURA TUN2, MASANARU TAKAI3, NOBUO SHIGEHARA3, TAKEHISA TSUBAMOTO3 Laboratory of Physical Anthropology, Department of Zoology, Kyoto University, Kitashirakawa Oiwakecho, Sakyo-ku, Kyoto, 606–8502 Japan,1 Department of Geology, University of Yangon, Yangon, Myanmar,2 Primate Research Institute, Kyoto University, Inuyama, Japan3 Received 23 May 2003; accepted 1 October 2003 Abstract Relative abundance and body size were compared among six primate genera found from the Pondaung Formation (latest middle Eocene, Myanmar). Pondaungia, consisting of two species, is the most common primate in the Pondaung fauna. A high abundance of Pondaungia savagei, the larger species, is recorded at the northern fossil localities (Mogaung area). At the southwestern localities (Bahin area), the primates are taxonomically more diverse than in the other areas, and there are no sig- nificant differences in abundance of the amphipithecid genera, Myanmarpithecus, Amphipithecus, and Pondaungia. The body sizes of the three eosimiid taxa (Eosimias-like eosimiid, undescribed eosimiid, and Bahinia) and Myanmarpithecus are distinct from one another, but those of the Amphipithecus and Pondaungia species are inseparable. The postcranial bones of a large-sized primate (NMMP 20) from the Pondaung Formation, consisting of humeral, calcaneal, ulnar, and vertebral fragments, most likely belong to Pondaungia cotteri or Amphipithecus mogaungensis, based on comparisons of body mass estimates of NMMP 20 with those of the taxonomically allocated dentognathic specimens. Although NMMP 20 was previously cited as Pondaungia savagei, we consider the current evidence to be insuf- ficient to resolve its species or generic-level taxonomic assignment. Key words: Pondaung primates, abundance, body size range, taxonomic assignment Introduction quently the material was described by Ciochon et al. (2001). Unfortunately, the postcranial material was found without The Pondaung Formation in central Myanmar, dating to any accompanying dental remains that would help settle its the latest middle Eocene, has yielded a diverse assemblage taxonomic assignment. No other primate specimens have of mammals, including primates (Cotter, 1914; Colbert, been recovered from the locality (Pk1). Ciochon et al. 1938; Aye Ko Aung, 1999; Tsubamoto et al., 2000, 2002; (2001) assigned NMMP 20 to Pondaungia savagei, the Tsubamoto, 2001). Two of the primates, Pondaungia and larger of the two species of Pondaungia, based on estimated Amphipithecus, have been known since the early twentieth body size and relative abundance of Pondaungia in the century (Pilgrim, 1927; Colbert, 1937). Other primate gen- Pondaung fauna, although Gunnell et al. (2000) did note that era, such as Myanmarpithecus (Takai et al., 2001), Bahinia assignment of postcranial material to Pondaungia or (Jaeger et al., 1999), an Eosimias-like eosimiid (Gebo et al., Amphipithecus was not possible due to the small size differ- 2002), and an undescribed eosimiid genus (Takai, in prep.) ence between the two genera. were discovered within the past five years. Until quite Given that the NMMP 20 specimen can, for the first time, recently, the Pondaung primates were known only from den- elucidate the postcranial morphology of the larger Pondaung tognathic remains, so their systematic position has predomi- primate, its assignment to any of the known Pondaung pri- nantly been discussed in terms of their dental morphology. mate taxa evokes two procedural questions: 1) whether or In 1997, postcranial specimens of a large primate (NMMP not the Pondaung primates are separable in terms of their 20; NMMPNational Museum of Myanmar, Primates), con- body size, and 2) whether or not there are significant differ- sisting of humeral fragments, two ulnar trochleae, a calca- ences in relative abundance of the primates of the Pondaung neal fragment, limb bone shafts, and vertebral fragments, Formation fauna, either in the composite fauna or in that of were collected from the Pondaung Formation, and subse- each area. Several new specimens of Pondaung primates have been collected since NMMP 20 was discovered. In this * Corresponding author. e-mail: [email protected] study, we clarify the relative body size range and geographic phone: 81-75-753-4094; fax: 81-75-753-4083 distributions of the Pondaung primates, using an enlarged Published online 16 April 2004 sample that includes the new material. We then reevaluate in J-STAGE (www.jstage.jst.go.jp) DOI: 10.1537/ase.00076 whether or not the current evidence can conclusively be used © 2004 The Anthropological Society of Nippon 67 68 N. EGI ET AL. ANTHROPOLOGICAL SCIENCE to assign NMMP 20 to a particular Pondaung primate taxon. ness of the strata, from the fossil bearing horizon to the base of the overlying Yaw Formation, is much greater in the Relative Abundance of the Pondaung Primates Mogaung area than in the other two areas. Nannoplanktonic fossils in rock samples from the Mogaung area are repre- The Pondaung Formation is lithologically divided into sented by index taxa of NP 15 to 18, while rock samples two parts, “the Lower Member” and “the Upper Member.” from the Bahin and Pangan areas predominantly contain All of the primate fossil-bearing horizons occur in the lower index nannoplanktonic fossils of NP 17 to 20. However, half of “the Upper Member” (Aye Ko Aung, 1999, in press). those from the Pangan area also contain nannoplanktonic This level is exposed at numerous places in the Pondaung fossils of NP 14 to 16 and NP 18 to 22 (Hla Mon, 1999). area, and there are many vertebrate fossil localities. The fos- Although resolution of relative age estimations based on sil localities are geographically divided into three areas, these index fossils is not high, and some of the samples Mogaung, Bahin, and Pangan (Figure 1; Pondaung Fossil apparently combine rocks of the Pondaung and other Paleo- Expedition Team, 1997; Aye Ko Aung, 1999; Tsubamoto, gene formations thus obscuring biostratigraphic relation- 2001). Although we have not yet reached a full understand- ships especially in the case of the Pangan area, the ing of the geological structure of the Pondaung Formation, nannoplanktonic fossils indicate that sediments of the some data suggest that the fossil horizons exposed in the Pondaung Formation of the Mogaung area are generally Mogaung area are lower in the sequence than those exposed older than those of the other two areas. in the Bahin and Pangan areas. Aye Ko Aung (1999, in press; The Pk1 locality (Sabapontaung kyitchaung in the local Aung Naing Soe et al., 2002) demonstrated that the thick- language), where NMMP 20 was found, is located in the Figure 1. Map of Pondaung area. The rectangles with thick gray border indicate the three main fossil locality assemblages (Mogaung, Bahin, and Pangan areas). Ellipses are fossil localities, and the locality numbers catalogued by Kyoto University are indicated in italics. Localities with black colored ellipses yielded primate fossils (listed in Table 1). Squares are townships and villages. Vol. 112, 2004 RELATIVE OCCURRENCES OF THE PONDAUNG PRIMATES 69 Bahin area. Ciochon et al. (2001) mentioned the greater more common than Amphipithecus mogaungensis. abundance of Pondaungia specimens compared with the In the Bahin area, where NMMP 20 was collected, all other Pondaung primates, and added a comment that a man- Pondaung genera are represented. Amphipithecus, Myan- dible of Pondaungia (P. savagei) had been found at a nearby marpithecus, and Pondaungia are represented by four, three, locality (Pk2) in the Bahin area. Although this evidence was and two individuals, respectively. As mentioned above, sin- used as the basis upon which they made their taxonomic gle individuals of each eosimiid taxon has been collected. assignment of NMMP 20 to Pondaungia, no data on the rel- All of these primate fossils, except one specimen, have been ative abundance of primate fossils from the Pondaung area collected from localities a few kilometers west of Pauk- were presented. kaung and Bahin villages (Figure 2). Among these primate- Our current sample of the Pondaung primates includes a bearing localities, Pk2, Pk3, and Pk5 are similar in strati- total of 33 specimens. Table 1 shows the abundance of each graphic level to Pk1 (Maung Maung, 2003). Of the two indi- primate species based on the number of individuals instead viduals of Pondaungia, one (NMMP 24) was found at Pk2, of the number of specimens. Some individuals have several as noted by Ciochon et al. (2001). An additional Pondaungia specimen numbers because they were collected by different specimen (NMMP 41) has been collected from Pk5. How- researchers or on different dates. Thus, counting the number of specimens will bias the relative abundance of the pri- mates. The sample of 33 specimens consists of 24 individu- als from six genera. Pondaungia has been divided into two species, a smaller P. cotteri and a larger P. savagei, by Gun- nell et al. (2002). We follow their classification for the spec- imens included in their study. Concerning the specimen collected after their study (NMMP 41), the species-level tax- onomy is left as indeterminate. When all three areas are combined, Pondaungia savagei and Amphipithecus mogaungensis are the most abundant species in the fauna, represented by six and seven individu- als, respectively. The other species are represented by smaller numbers of individuals, three each for Myanmarp- ithecus yarshensis and for Pondaungia cotteri and one each for the eosimiid taxa (Bahinia, Eosimias, and the unde- scribed new genus). Pondaungia is found to be the most Figure 2. Detailed map of the Bahin and Paukkaung localities in abundant primate in the Pondaung Formation, as suggested the Bahin area. Closed circles are the fossil localities, and the black by Ciochon et al. (2001). Pondaungia savagei is the most circles are the primate-bearing localities. Locality names are indicated abundant species of the current sample, but it is only slightly in italics.
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