Late Eocene Sivaladapid Primate from Guangxi Zhuang Autonomous
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Tao Qi Late Eocene sivaladapid primate from Institute of Vertebrate Guangxi Zhuang Autonomous Region, Paleontology & People’s Republic of China Paleoanthropology, Chinese Academy of Sciences, P.O. Box 643, Beijing A new genus and species of Sivaladapidae is described from the late 100044, People’s Republic of Eocene Gongkang Formation, Yongle Basin, western Guangxi China Zhuang Autonomous Region, southern China. Guangxilemur tongi, new genus and species, shows a combination of traits that occur separately in earlier and more primitive Asian adapiforms (Hoangho- K. Christopher nius and Rencunius) and in Miocene sivaladapines (Sivaladapis and Beard* Sinoadapis). Phylogenetic analysis of dental characters suggests that Section of Vertebrate Guangxilemur is closely related to the Miocene sivaladapine clade. Paleontology, Carnegie Miocene sivaladapines were the latest surviving members of a broad Museum of Natural History, radiation of Eocene adapiforms in Asia that included Hoanghonius, 4400 Forbes Avenue, Rencunius, and Wailekia in addition to Guangxilemur. European Pittsburgh, Pennsylvania Periconodon may also be specially related to this primarily Asian clade, 15213, U.S.A. E-mail: but current anatomical data are insufficient to test this possibility [email protected] adequately. Sivaladapine adapiforms and tarsiid tarsiiforms maintained relictual distributions in southern and/or southeastern Received 19 February 1998 Asia far beyond the extirpation of their closest relatives on other Revision received 24 April Holarctic continents near the Eocene–Oligocene boundary. This 1998 and accepted 27 April temporal persistence was mediated by Asian paleogeography, which 1998 allowed virtually continuous access to tropical refugia during a middle Cenozoic interval of climatic deterioration that coincided with the Keywords: Guangxilemur, extinction of adapiforms and tarsiiforms in Europe and North Sivaladapidae, Eocene, America. paleontology, phylogeny, 1998 Academic Press primates. Journal of Human Evolution (1998) 35, 211–220 Article No. hu980240 Introduction recognized by Gingerich & Sahni (1979, 1984) and subsequent authors, the Sivaladapid primates are distinctive ele- persistence of sivaladapids into the late ments of middle–late Miocene faunas from Miocene of southern Asia begs the question the Indian subcontinent and Yunnan of how this Neogene adapiform clade Province, China (Gingerich & Sahni, 1979, relates to the much broader Paleogene 1984; Wu & Pan, 1985; Pan, 1988). radiation of these animals. However, the Although the first fragmentary fossils of great stratigraphic disparity between these animals were misinterpreted as per- middle–late Miocene sivaladapids and taining to procyonid carnivores and lorisid all other known adapiforms, which are primates (Pilgrim, 1932; Lewis, 1933; virtually restricted to the Eocene (e.g. Tattersall, 1968), discovery of more nearly Szalay & Delson, 1979; Gheerbrant complete specimens allowed Gingerich & et al., 1993; Simons et al., 1995; Simons, Sahni (1979, 1984) to demonstrate the 1997), has hampered attempts to place adapiform affinities of these taxa. As sivaladapids within an explicit phylogenetic *To whom correspondence should be addressed. context. 0047–2484/98/090211+10 $30.00/0 1998 Academic Press 212 . . Figure 1. Map showing location of field area and type locality for Guangxilemur tongi. During a joint Institute of Vertebrate Systematic paleontology Paleontology & Paleoanthropology (IVPP)- Carnegie Museum of Natural History field Order Primates Linnaeus, 1758 expedition in February and March 1995, Suborder Strepsirhini Geoffroy, 1812 we prospected fossiliferous outcrops of the Infraorder Adapiformes Szalay & late Eocene Gongkang Formation along Delson, 1979 the shores of Chengbihu Reservoir in the Family Sivaladapidae Thomas & Yongle Basin, western Guangxi Zhuang Verma, 1979 Autonomous Region, China (Figure 1). Among the fossil vertebrates recovered during the course of that field work are Guangxilemur, new genus two teeth of a sivaladapid primate, probably pertaining to the same individual. Type species. Guangxilemur tongi, new These specimens are the first fossil species. primates to be reported from Guangxi Zhuang Autonomous Region. In addition, Diagnosis. Larger than Hoanghonius, Rencu- they form the basis for a new genus nius, and Wailekia.M2 differs from those of and species of Sivaladapidae, which is Hoanghonius and Rencunius in showing described below. In light of this new greater development of external shearing taxon, we also propose a new hypothesis crests, stronger parastyle and mesostyle, and regarding the content and phylogeny of weaker conules. M2 differs from those of Sivaladapidae. Sivaladapis and Sinoadapis in possessing 213 large pericone and hypocone on lingual Formation, Yuanqu Basin, Shanxi and cingulum. Henan Provinces, China, are decidedly more primitive than those of either the Krabi fauna of southern Thailand or the Guangxilemur tongi, new species Gongkang Formation. Indeed, all available Holotype. IVPP V11652, a left M2 [Figure biostratigraphic evidence suggests that the 2(a)]. IVPP V11653, a right C1 [Figure Heti Formation faunas are late middle 2(b)–(c)], probably pertains to the same Eocene, while those from Krabi and the individual (see below). Both specimens were Gongkang Formation are younger (Russell collected by Guo Jianwei on 1 March 1995. & Zhai, 1987; Holroyd & Ciochon, 1994; Ducrocq et al., 1997). As such, Guangxile- Type locality. Outcrop of Gongkang mur from the Gongkang Formation is sig- Formation on the north shore of Chengbihu nificantly younger than either Hoanghonius Reservoir, west of Wanjiang village (Figure or Rencunius from the Heti Formation, but is 1). Exact coordinates of the type locality are probably similar in age to Wailekia from as follows: 24)01.05*North, 106)35.94*East. southern Thailand. Known distribution. Late Eocene of Guangxi Diagnosis. As for the genus (currently Zhuang Autonomous Region, China. The monotypic). associated mammalian fauna from the Gongkang Formation was reviewed by Etymology. For our friend and colleague Russell & Zhai (1987), who were unable to Tong Yongsheng, in recognition of his many decide whether the fauna should be consid- contributions to knowledge of Paleogene ered late Eocene or early Oligocene in mammals in China and his field work in the age. However, since Russell & Zhai’s Baise and Yongle basins of Guangxi. (1987) survey, the position of the Eocene– Oligocene boundary with respect to Asian Description. The holotype is a left upper mammal faunas has shifted, such that many molar that we identify as M2 on the basis of faunas formerly considered as late Eocene comparisons with maxillary fragments of are now late middle Eocene, and those for- Sivaladapis and Sinoadapis that bear serially merly considered as early Oligocene are now associated upper molars. The crown late Eocene (e.g., Berggren & Prothero, measures 6·6 mm (mesiodistal length) by 1992; Ducrocq, 1993; Holroyd & Ciochon, 9·1 mm (buccolingual breadth). Buccal and 1994; Emry et al., 1998). lingual cingula are prominently developed Anthracotheriid artiodactyls are well rep- and virtually continuous around the entire resented in late middle Eocene and younger periphery of the crown. Lingually, the mammal faunas in Asia and are widely cingulum bears both a pericone and a regarded to be useful for purposes of bio- hypocone. In terms of volume, the latter stratigraphy (e.g., Holroyd & Ciochon, cusp is the larger, but both are highly con- 1994; Ducrocq, 1994, 1997; Ducrocq et al., spicuous. A weak crest connects the 1997). Anthracotheres from the Gongkang hypocone with a series of at least three small Formation, originally reported by Tang cuspules that lie buccal to the hypocone, on (1978), have recently been described as a cingular shelf below the level of the post- ‘‘almost identical’’ to a species of Anthraco- protocrista. The protocone is canted therium from the Krabi fauna of southern mesially, as is frequently the case in pri- Thailand, also of late Eocene age (Ducrocq mates, so that its apex lies nearer to the et al., 1997). Anthracotheres from the Heti paracone than the metacone. Pre- and 214 . . Figure 2. Guangxilemur tongi, n. gen. et sp.: (a) IVPP V11652, holotype left M2, occlusal view; (b–c) IVPP V11653, fragmentary right C1, probably associated with holotype, in lingual (b) and labial (c) views. All views are stereopairs. Scale=5 mm. postprotocristae emanate mesiobuccally and conules have been rendered indistinguish- distobuccally from the protocone to form able by wear, which is only moderate. the lingual margin of the trigon basin. If they Mesiobuccally, the preprotocrista joins the were ever present as discrete structures, the preparacrista just lingual to the parastyle. 215 Figure 3. Comparative schematic drawings of M2 in selected sivaladapid primates (not to scale): (a) Hoanghonius stehlini;(b)Guangxilemur tongi; (c) Sivaladapis nagrii. Note the presence of pericone and hypocone in Hoanghonius and Guangxilemur, and the increased development of external shearing crests with parastyle and mesostyle in Guangxilemur and Sivaladapis. See text for discussion of these and other characters and their possible phylogenetic significance. The postprotocrista merges with the post- crest bears a well-defined wear facet that cingulum distolingual to the metacone. must have formed by contact with the an- Buccal shearing crests are well-developed, teriormost lower premolar (presumably P2). connecting the parastyle with the