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A New Genus of Pliopithecoid from the Late Early Miocene of China and Its Implications for Understanding the Paleozoogeography of the Pliopithecoidea

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A New Genus of Pliopithecoid from the Late Early Miocene of China and Its Implications for Understanding the Paleozoogeography of the Pliopithecoidea Journal of Human Evolution 145 (2020) 102838 Contents lists available at ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol A new genus of pliopithecoid from the late Early Miocene of China and its implications for understanding the paleozoogeography of the Pliopithecoidea * Terry Harrison a, Yingqi Zhang b, c, , Guangbiao Wei d, Chengkai Sun e, Yuan Wang b, c, Jinyi Liu b, c, Haowen Tong b, c, Baiting Huang f,FanXuf a Center for the Study of Human Origins, Department of Anthropology, New York University, New York, NY, 10003, USA b Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 100044, People's Republic of China c CAS Center for Excellence in Life and Paleoenvironment, Beijing, 100044, People's Republic of China d Chongqing Institute of Geological Survey, Chongqing, 401122, People's Republic of China e Division of Natural History, Shandong Museum, Jinan, 250014, People's Republic of China f Cultural Heritage Administration of Fanchang County, Wuhu City, Anhui Province, Wuhu, 241200, People's Republic of China article info abstract Article history: A diversity of pliopithecoids is known from Miocene localities in Europe, but until recently, this group Received 20 March 2020 was relatively poorly represented in China. However, new discoveries have shown that Chinese pliopi- Accepted 26 May 2020 thecoids were taxonomically diverse and geographically widespread. The earliest pliopithecoids in China Available online 10 July 2020 (and Eurasia) are Dionysopithecus and Platodontopithecus from the Early Miocene of Sihong, Jiangsu (~19À18 Ma). During the Middle Miocene (~15À12 Ma), several species of pliopithecoids are recorded at Keywords: localities in Gansu Province (Laogou), Inner Mongolia (Damiao), Xinjiang Uygur Autonomous Region Pliopithecid (Tieersihabahe), and Ningxia Hui Autonomous Region (Tongxin). Finally, a late-surviving anapithecine Crouzeliid Dental morphology crouzeliid, Laccopithecus robustus, is known from the Late Miocene (~7 Ma) of Shihuiba in Yunnan, which Phylogeny postdates the extinction of pliopithecoids in Europe (during MN 10). Paleontological investigations at a Zoogeography late Early Miocene locality near Fanchang in Anhui Province have yielded a large sample of isolated teeth (more than one hundred) of a previously unknown species of pliopithecoid. The associated micro- mammals indicate an age contemporaneous with the Shanwang Formation in Shandong Province (MN 3 e4, ~18À17 Ma). All of the permanent teeth are represented except for I2. With its unique suite of dental features, the Fanchang pliopithecoid can be attributed to a new species and genus. Shared derived features of the lower molars confirm that the Fanchang pliopithecoid has its closest affinities with Eu- ropean crouzeliids, but a number of primitive traits indicate that it is a stem member of the clade. The evidence points to China as an important center for the early diversification of pliopithecoids. Contrary to previous zoogeographic scenarios, the occurrence of an early crouzeliid in China implies that the Plio- pithecidae and Crouzeliidae may have diverged from a stem pliopithecoid in Asia during the Early Miocene before their arrival in Europe. © 2020 Elsevier Ltd. All rights reserved. 1. Introduction Eurasia at ~21 Ma (Andrews et al., 1996; Harrison, 2005, 2013). The pliopithecoids were a relatively diverse and successful group, with Pliopithecoids are an extinct clade of stem catarrhines from the 10 genera and 19 species currently recognized, spanning more than Miocene of Eurasia. They were the first catarrhines to migrate out of 10 million years, with a geographical range that extended from the Africa, soon after the collision of the Afro-Arabian plate with Iberian Peninsula to eastern China (Andrews et al., 1996; Begun, 2002; Harrison, 2013; Marigo et al., 2014). They became extinct in Eurasia during the Late Miocene, presumably as a consequence of progressive cooling and increased seasonality at higher latitudes * Corresponding author. associated with a corresponding shift from subtropical evergreen E-mail address: [email protected] (Y. Zhang). https://doi.org/10.1016/j.jhevol.2020.102838 0047-2484/© 2020 Elsevier Ltd. All rights reserved. 2 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 woodlands to habitats dominated by deciduous broad-leaved the Early Miocene of Sihong, Jiangsu (~19À18 Ma), are the earliest woodlands and C4 grasslands (Fortelius et al., 2014). pliopithecoids in China (and Eurasia) (Harrison and Gu, 1999). The alpha taxonomy of the clade is well established (Andrews During the Middle Miocene (~15À12 Ma), several species of plio- et al., 1996; Begun, 2002, 2017; Harrison, 2005, 2013; Alba and pithecoids are recorded at localities in Gansu Province (Laogou), Moya-Sol a, 2012), with differences of opinion mainly pertaining Inner Mongolia Autonomous Region (Damiao), Xinjiang Uygur to the taxonomic ranks applied to the various subclades. The clas- Autonomous Region (Tieersihabahe), and Ningxia Hui Autonomous sification adopted here is presented in Table 1. The Pliopithecoidea Region (Tongxin) (Harrison et al., 1991; Wu et al., 2003; Deng, is differentiated into four families: Dionysopithecidae, Pliopitheci- 2004; Deng et al., 2004; Zhang and Harrison, 2008; Kaakinen dae, Crouzeliidae, and Krishnapithecidae. The dionysopithecids et al., 2015). Finally, a late-surviving crouzeliid, Laccopithecus (including Dionysopithecus and Platodontopithecus) represent early robustus (Wu and Pan, 1984, 1985; Pan, 1988), is known from the stem pliopithecoids from China (Harrison and Gu, 1999). The plio- Late Miocene (~7 Ma) of Shihuiba in Yunnan, which postdates the pithecids (including Pliopithecus) and crouzeliids (including Ple- extinction of pliopithecoids in Europe (during MN 10; Fig. 1). The siopliopithecus, Anapithecus, Laccopithecus, Egarapithecus, and present study provides further evidence to document the remark- Barberapithecus), from the Middle and Late Miocene of Eurasia are able diversity of pliopithecoids known from the Miocene of China. more derived than the dionysopithecids (Andrews et al., 1996; In 1984, fossil vertebrates of Miocene age were discovered in a Begun, 2002; Harrison, 2005, 2013; Alba and Moya-Sol a, 2012). limestone quarry at Laili Mountain near the village of Suncun in Krishnapithecidae includes a single highly derived species, Krish- Fanchang County, Anhui Province. The fossils were forwarded to napithecus krishnaii, from the Late Miocene of India (Sankhyan the Anhui Museum in Hefei and the Institute of Vertebrate Pale- et al., 2017). ontology and Paleoanthropology (IVPP) in Beijing (Zheng, 1993; Jin Pliopithecoids are best known from Europe (with at least 12 and Liu, 2009). Full-scale excavations at the site, directed by species), and until recently, they were relatively poorly represented Changzhu Jin in 1999 and 2000, led to the recovery of a diverse in Asia. However, new discoveries in Thailand, India, and especially assemblage of fossil mammals, including more than one hundred China have shown that Asian pliopithecoids were taxonomically isolated teeth of a large species of pliopithecoid (Jin and Wei, 1999). diverse and geographically widespread (Wu and Pan, 1984, 1985; The associated micromammals (Qiu and Jin, 2016, 2017) correspond Qiu and Guan, 1986; Pan, 1988; Suteethorn et al., 1990; Harrison most closely to those from the Shanwang and Xiacaowan Forma- et al., 1991; Wu et al., 2003; Harrison, 2005, 2013; Zhang and tions (Qiu et al., 1999; Qiu and Qiu, 2013) and indicate an estimated Harrison, 2008; Chaimanee et al., 2015; Kaakinen et al., 2015; age of ~18À17 Ma (late Early Miocene, equivalent to middle Sankhyan et al., 2017). Dionysopithecus and Platodontopithecus from Table 1 Classification of the Pliopithecoidea (after Andrews et al., 1996; Alba and Moya-Sol a, 2012; Harrison, 2013; Alba and Berning, 2013; Sankhyan et al., 2017). Order: Primates Linnaeus, 1758 Suborder: Anthropoidea Mivart, 1864 Infraorder: Catarrhini E. Geoffroy Saint-Hilaire, 1812 Superfamily: Pliopithecoidea Zapfe, 1961a Family: Dionysopithecidae Harrison and Gu, 1999 Dionysopithecus Li, 1978 D. shuangouensis Li, 1978 D. orientalis (Suteethorn et al., 1990) Platodontopithecus Gu and Lin, 1983 Plat. jianghuaiensis Gu and Lin, 1983 Family: Krishnapithecidae Sankhyan et al., 2017 Krishnapithecus Ginsburg and Mein, 1980 K. krishnaii (Chopra and Kaul, 1979) Family: Pliopithecidae Zapfe, 1961a Pliopithecus Gervais, 1849 Plio. antiquus (Blainville, 1839) Plio. bii Wu et al., 2003 Plio. canmatensis Alba et al., 2010 Plio. piveteaui Hürzeler, 1954 Plio. platyodon Biedermann, 1863 Plio. vindobonensis Zapfe and Hürzeler, 1957 Plio. zhanxiangi Harrison et al., 1991 Family: Crouzeliidae Ginsburg and Mein, 1980 Subfamily: Crouzeliinae Ginsburg and Mein, 1980 Crouzelia Ginsburg, 1975 C. auscitanensis Ginsburg, 1975 C. rhodanica Ginsburg and Mein, 1980 Plesiopliopithecus Zapfe, 1960 Plesio. lockeri (Zapfe, 1960) Barberapithecus Alba and Moya-Sol a, 2012 B. huerzeleri Alba and Moya-Sol a, 2012 Subfamily: Anapithecinae Alba and Moya-Sol a, 2012 (new rank) Anapithecus Kretzoi, 1975 A. hernyaki (Kretzoi, 1975) cf. A. priensis (Welcomme et al., 1991) Laccopithecus Wu and Pan, 1984 L. robustus Wu and Pan, 1984 Egarapithecus Moya-Sol a et al., 2001 E. narcisoi Moya-Sol a et al., 2001 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 3 Figure 1. Map showing
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