Journal of Human Evolution 145 (2020) 102838

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Journal of Human Evolution

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A new genus of pliopithecoid from the late Early of China and its implications for understanding the paleozoogeography of the

* 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 (~1918 Ma). During the Middle Miocene (~1512 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- indicate an age contemporaneous with the Shanwang Formation in Shandong Province (MN 3 e4, ~1817 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 (~1918 Ma), are the earliest woodlands and C4 grasslands (Fortelius et al., 2014). pliopithecoids in China (and Eurasia) (Harrison and Gu, 1999). The alpha of the clade is well established (Andrews During the Middle Miocene (~1512 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 ) and crouzeliids (including Ple- extinction of pliopithecoids in Europe (during MN 10; Fig. 1). The siopliopithecus, , 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 ~1817 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: Linnaeus, 1758 Suborder: Anthropoidea Mivart, 1864 Infraorder: 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: 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-Sola, 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 the location of paleontological sites in China that have yielded pliopithecoids. Damiao (Inner Mongolia Autonomous Region), Pliopithecoidea indet.; Fanchang (Anhui Province), gen. et sp. nov.; Laogou (Gansu Province), Pliopithecus sp.; Shihuiba (Yunnan Province), Laccopithecus robustus; Sihong (Jiangsu Province), Dio- nysopithecus shuangouensis and Platodontopithecus jianghuaiensis; Tieersihabahe (Xinjiang Uygur Autonomous Region), Pliopithecus bii; Tongxin (Ningxia Hui Autonomous Region), Pliopithecus zhanxiangi.

Orleanian, European Land Zone MN 3e4) (Mein, 1999; located 20 km south of the Yangtze (Chang Jiang) River. Laili Hilgen et al., 2012; Qiu and Qiu, 2013). Mountain is composed of limestone associated with the Biandan The range of metrical and morphological variation in the sample Formation of age (Yuan et al., 2009). During the later of teeth from Fanchang does not exceed that observed within Neogene, a number of fossiliferous karstic fissures and caverns species of extant anthropoids, so we recognize all of the specimens were produced by meteoric dissolution of the limestone (Jin and as belonging to a single species. Comparisons demonstrate that the Liu, 2009). The oldest fissure, the pliopithecoid-bearing Tangkou Fanchang species exhibits a distinctive suite of dental features that fissure, occurs near the top of the southern slope of Laili Mountain distinguishes it from all other pliopithecoids. On the basis of these at an elevation of about 143 m above sea level (Qiu and Jin, 2016; differences, a new genus and species is recognized. The Fanchang Fig. 3). Vertebrate remains on the paleolandscape surface were pliopithecid is inferred to be younger than the dionysopithecids washed into the fissure along with colluvial and fluvial sediments. from the Xiacaowan Formation at Sihong (Harrison and Gu, 1999; Younger fossiliferous fissures and caverns are located near the base Qiu and Qiu, 2013; Qiu et al., 2013) but still represents one of the of the mountain, and these include the Early Pleistocene Paleolithic earliest known pliopithecoids. The discovery adds to the growing site of Renzidong and the Middle and Late Pleistocene localities of diversity of pliopithecoids from China and provides crucial insights Xidong and Ludong, respectively (Qiu and Jin, 2016; Fig. 3). The into their early evolutionary history and biogeography. We present Tangkou fissure was exposed as a result of commercial quarrying by here a detailed description of the new taxon, along with a discus- the Suncun Cement Factory. Limestone extraction ceased after the sion of its phylogenetic and zoogeographic relationships. discovery of important paleontological and archaeological finds at Renzidong, and today, the fossil localities at Laili Mountain are protected. 1.1. Geology and chronology The sedimentary sequence in the Tangkou fissure consists of an alternating series of well-cemented sandstones and conglomerates, The fossils were recovered from a fissure filling at Laili Mountain more than 20-m thick (Qiu and Jin, 2016; Fig. 4). Four lithological (Lailishan), located 2 km northwest of Suncun Village and 10 km units are recognized (Qiu and Jin, 2016). The lowermost unit, layer southwest of the county town of Fanchang in Anhui Province, 1, exceeding 9.6 m in thickness, consists of yellow-gray sandstones eastern China (Jin and Liu, 2009; Qiu and Jin, 2016; Fig. 2). The site is 4 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838

Figure 2. Map showing the location of Laili Mountain (Lailishan), near the county town of Fanchang in Anhui Province, eastern China, where the pliopithecoid-bearing Tangkou Fissure is located.

Figure 3. Aerial view of the southern slope of Laili Mountain showing the location of the Tangkou Fissure (center right). The Early Pleistocene locality of Renzidong, the Middle Pleistocene locality of Xidong, and the Late Pleistocene locality of Ludong are also highlighted. The museum complex is in the foreground. Note the extensive exposure of limestone as a result of past commercial quarrying of Laili Mountain. and gravels that grade upward from fine-grained sandstones at the have produced fossil vertebrates, but most, including all of the base to coarser sandstones and gravels at the top. Lenticular pliopithecoids, are derived from layer 3. The faunas from the sandstones and gravels occur locally. Layer 2 consists of a thin band different layers at Tangkou are of varied ages, ranging from Early (80-cm thick) of brown-gray sandstones and gravels interbedded Miocene to Pleistocene (Qiu and Jin, 2016, 2017). The Pleistocene with siliceous bands. Layer 3, with a thickness of 4.8 m, consists of a fossils are yellowish red in color and show no evidence of having well-cemented conglomerate consisting of sandstones and well- been water transported. The pliopithecoids and the other fossils rounded and poorly sorted gravels and cobbles. The fissure is derived from layer 3 are grayish black or brown, and many show capped by layer 4, which consists of a collapsed breccia composed evidence of having been rolled as a result of fluvial transportation. of blocks of limestone (up to 50 cm in diameter) in a matrix of red- The Tangkou large mammal fauna has not yet been systemati- brown clay, yellow shales, and poorly sorted gravels. All four units cally studied. However, the rodents provide a reasonably good basis T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 5

wherein the serial position is known. Canines have been sexed based on their size and morphology. The preservation of the fossils is quite distinctive. Only isolated teeth are represented. Many of the teeth exhibit various degrees of polishing and rolling as a result of transportation in a high-energy fluvial sedimentary environment. Most of the teeth lack roots, and some consist only of enamel caps. Owing to the effects of polishing, few molars preserve the interproximal contact facets, making identification of their serial positions somewhat challenging. The teeth were presumably washed into the fissure from the surface and then tumbled around with the sand and gravel infilling before consolidation and cementation of the sediments. There is no evi- dence of digestive etching on the teeth or other indications that carnivores or other vertebrates were accumulating agents. A few specimens show evidence of fine plant root etching on the enamel surface, which indicates that vegetation grew in the fissure after burial of the teeth. The majority of the permanent cheek teeth (at least in which the original wear pattern can be discerned) are either germs or unworn to very lightly worn (n ¼ 47/79, ~60%), suggesting that the composition of individuals may have been skewed toward juveniles and young adults. However, preservation and sampling issues preclude the possibility of constructing a detailed age profile. This published work and the nomenclatural acts it contains have been registered with ZooBank: LSID, urn:lsid:zoo- bank.org:pub:3951290E-0266-43A9-987F-171C8BFA6423.

2.2. Data collection

Dimensions of the teeth were measured to the nearest 0.1 mm using Mitutoyo digital calipers (Table 2). Mesiodistal (MD) and buccolingual (BL) dimensions were taken on the incisors, lower second premolar, upper premolars, molars, and deciduous incisors and premolars. Crowns were oriented using the interproximal Figure 4. Schematic stratigraphic section of the Tangkou fissure at Laili Mountain facets (when preserved). For upper and lower canines and P3, the (after Qiu and Jin, 2016). Layer 1: well-cemented grayish yellow sandstones; layer 2: maximum length (maxL) and perpendicular breadth (perpB) were coarse, well-indurated brownish gray sandstones with siliceous bands; layer 3: brownish gray sandstones and conglomerates; layer 4: reddish brown sandy clays with measured in place of MD and BL dimensions. Height of the crown of limestone blocks and occasional yellowish shales. Most mammal fossils, including all the incisors and canines was measured at the buccal face (BHT). The of the pliopithecoids, were recovered from layer 3. (For interpretation of the references vertical height of the crown on the buccal face (BHT) and the length to color in this figure legend, the reader is referred to the Web version of this article.) of the mesiobuccal honing face (HHT) were taken on P3. The height of the lingual and buccal cusps of the upper premolars (LHT and BHT, respectively) was measured from the cementum-enamel for biochronological correlation for level 3. Diatomys shantungensis junction to the cusp apex. Lower molar flare (MF) is calculated and Neocometes sinensis are species shared with the Xiacaowan using the formula developed by Singleton (2003):MF¼ 1 mesial Formation at Sihong (Qiu and Jin, 2017), and D. shantungensis and intercuspal breadth/maximum mesial breadth. The lingual notch Plesiosciurus zhengi are shared with the Shanwang Formation at height (LNH) index of the lower molars is calculated as follows: Shandong (Qiu and Qiu, 2013; Qiu and Jin, 2016). P. zhengi from LNH ¼ height from crown cervix to the base of the lingual notch/ Tangkou has close affinities with Plesiosciurus sinensis from Sihong, height from the crown cervix to the apex of the metaconid. but the larger size and dental features of the former indicate that it Measurements of comparative specimens were taken from is a more derived species (Qiu and Jin, 2016; Qiu, 2017). Overall, the either the original specimens or published sources. Molar rodents indicate that the Tangkou fauna correlates best with that morphology terminology follows Harrison and Gu (1999; Fig. 5). from the Shanwang Formation (~1817 Ma) and is slightly younger Body mass estimation is based on the M and M to body mass than that from the Xiacaowan Formation (~1918 Ma; Qiu et al., 1 2 regression formulas for extant primates (Gingerich et al., 1982). 2013).

2.3. Data analysis 2. Materials and methods The classification of the pliopithecoids follows that of Harrison 2.1. Samples and Gu (1999) and Harrison (2005, 2013) with the following up- dates and amendments: (1) the recently described crouzeliid, The sample of pliopithecoids from Fanchang consists of 108 Barberapithecus huerzeleri (Alba and Moya-Sol a, 2012), is included; isolated teeth (Supplementary Online Material [SOM] Table S1). All (2) Krishnapithecus krishnaii is recognized as a pliopithecoid within of the permanent teeth, except I2, and several deciduous teeth are the family Krishnapithecidae (following Sankhyan et al., 2017); (3) represented in the collection. The specimens are housed in the IVPP, Pliopithecus piveteaui, previously subsumed into Pliopithecus anti- Chinese Academy of Sciences. Upper and lower first and second quus (Andrews et al., 1996; Harrison, 2013), is provisionally molars are assigned to their appropriate serial position based on retained as a separate species (following Hürzeler, 1954; Begun, morphological criteria derived from other species of pliopithecoids 2002; Gagnaison et al., 2006, 2009; Alba et al., 2010); (4) some 6 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838

Table 2 Table 2 (continued ) Dental dimensions (mm) of Fanchangia jini.a Tooth Specimen MD BL BHT LHT Tooth Specimen MD BL BHT LHT V 18115.41 5.6 6.5

I1 V 18115.26 3.2 3.9 5.8 V 18115.101 5.3 6.3 C (male) V 18115.31 9.3 6.4 e 1 MD ¼ mesiodistal length, maximum length for canines and P ;BL¼ buccolingual C (female) V 18115.30 6.4 4.6 e 3 1 breadth, perpendicular breadth for canines and P ; BHT ¼ buccal height of incisors P V 18115.32 5.9 4.0 3 3 and canines, height of buccal cusp of upper premolars; LHT ¼ height of lingual cusp V 18115.95 5.2 3.9 of upper premolars. P V 18115.27 6.8 5.5 4 a Includes only those teeth in which accurate measurements could be taken (see V 18115.28 6.3 5.7 SOM Table S1 for a comprehensive list of specimens and their dimensions). All V 18115.96 5.4 4.9 specimens are housed in the Vertebrate Paleontology collections (V), Institute of V 18115.97 5.7 5.0 Vertebrate Paleontology and Paleoanthropology, Beijing, China. V 18115.98 5.5 5.3

M1 V 18115.1 7.8 6.0 V 18115.7 8.8 6.9 authors (Begun, 2002, 2017; Alba et al., 2015) include the pliopi- e V 18115.14 8.4 thecid from Devínska Nova Ves in the genus , but we V 18115.17 8.4 6.5 fi V 18115.19 7.8 5.9 do not consider the differences suf cient to merit its exclusion from V 18115.22 8.1 6.1 the genus Pliopithecus (following Zapfe and Hürzeler, 1957; Zapfe, V 18115.24 7.5 6.2 1958); (5) the isolated M1 of a pliopithecoid from Priay in France V 18115.87 7.4 6.1 is referred to cf. Anapithecus priensis (following Alba and Moya- V 18115.90 7.8 5.8 Sola, 2012); (6) Crouzelia, previously synonymized with Plesioplio- M2 V 18115.2 9.8 7.8 V 18115.3 9.3 7.4 pithecus (see Andrews et al., 1996 for rationale), is recognized here V 18115.4 9.4 7.6 as a separate genus, distinguished mainly by the absence of crests V 18115.5 e 7.6 on the lower molars linking the entoconid and hypoconulid so that V 18115.8 9.5 7.7 the distal fovea communicates directly with the talonid basin; and V 18114 9.5 7.4 V 18115.23 9.4 7.6 (7) Crouzeliinae is elevated to family rank (following Begun, 2002) V 18115.91 e 7.6 in recognition of the morphological distinctiveness and taxonomic

M3 V 18115.9 10.2 7.4 diversity of the clade (see Table 1). V 18115.10 9.5 6.3 To assess the relationships of the pliopithecoid from Fanchang, V 18115.11 9.1 6.4 we carried out a phylogenetic analysis using PAUP* (version 4.0a, V 18115.71 8.4 6.2 V 18115.88 10.6 7.6 build 166; Swofford, 2002). All known pliopithecoid genera from

dP4 V 18115.15 7.5 5.8 Eurasia were included (except the poorly known Krishnapithecus; V 18115.20 7.9 6.0 Sankhyan et al., 2017). Characters included in the analysis are based V 18115.21 8.3 5.8 entirely on the dentition because the cranial and postcranial I1 V 18115.64 e 5.6 8.0 V 18115.68 6.1 5.4 8.1 morphology is unknown for most genera of pliopithecoids. In V 18115.75 6.2 5.9 8.5 addition, three genera (i.e., Plesiopliopithecus, Crouzelia, and Egar- I2 V 18115.58 3.4 3.5 4.3 apithecus) are known principally or exclusively by the lower post- V 18115.63 4.5 4.6 5.1 canine dentition. The 25 characters (SOM Tables S2 and S3) selected V 18115.69 4.5 4.5 5.2 for the phylogenetic analysis have previously been found to be C1 (female) V 18115.76 6.0 4.9 6.0 V 18115.78 6.5 5.7 e helpful in discriminating the different genera of pliopithecoids P3 V 18115.57 5.6 8.1 ee(Andrews et al., 1996; Harrison and Gu, 1999; Moya-Sol a et al., V 18115.80 5.5 7.6 6.1 3.6 2001; Begun, 2002; Harrison, 2013). V 18115.93 5.4 8.2 7.0 4.2 Sixteen characters are coded as binary (states 0 and 1), and 9 of P4 V 18115.55 5.2 8.5 ee V 18115.62 5.3 8.3 eethem are coded as ternary (states 0, 1, and 2). Character states are V 18115.67 5.2 eeecoded as '?' when data are missing. Among the characters used, 7 V 18115.70 5.8 8.6 eeare continuous but coded as discrete based on empirical catego- V 18115.79 5.3 9.4 eerizing (i.e., characters #2, 8, 17, 18, 20, 22, and 25 in SOM Table S2) ee V 18115.94 5.2 7.1 (following Moya-Sol a et al., 2001). Characters #11 and #19 are M1 V 18115.33 7.6 9.3 V 18115.34 6.7 8.2 coded as multistate because of variable traits among Pliopithecus V 18115.39 7.1 8.1 species (i.e., vestigial occurrence of the pliopithecine triangle in V 18115.50 6.8 8.5 Plio. vindobonensis and M3 smaller than M2 in Plio. piveteaui) All the V 18115.53 7.3 8.5 characters are set to Wagner type (ord in PAUP*; Farris, 1970; V 18115.54 7.6 e V 18115.104 e 7.8 Felsenstein, 1983), which means that characters are allowed to M2 V 18115.35 8.2 10.2 transform both forward and backward. Two stem catarrhines, V 18115.36 9.0 10.3 and , are included in the analysis as V 18115.37 8.3 10.4 successive outgroups to the pliopithecoids to polarize the charac- V 18115.48 8.4 10.3 ters and root the trees. The two outgroups are set as paraphyletic to V 18115.49 8.5 9.9 V 18115.51 7.9 10.0 the in-group. Because the data matrix only contains 12 taxa and 25 V 18115.74 8.1 9.5 characters, an exhaustive search method is applied to search for the M3 V 18115.42 6.7 7.7 most parsimonious trees. A strict consensus tree is computed for V 18115.73 7.7 9.7 the most parsimonious Wagner trees. V 18115.84 8.4 e V 18115.107 7.4 9.6 1 dI V 18115.16 5.4 4.4 4.7 3. Systematic paleontology dC1 V 18115.83 5.6 4.5 e dP4 V 18115.52 6.6 7.7 V 18115.40 6.2 7.6 Order Primates Linnaeus, 1758 Infraorder Catarrhini E. Geoffroy Saint-Hilaire, 1812 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 7

mesial fovea hypoparacrista preprotocrista preparacrista paraconule trigon basin

paracone protocone

postparacrista lingual cingulum

buccal cingulum

premetacrista prehypocrista

metacone

hypocone postmetacrista distal fovea posthypocrista crista obliqua mesial fovea hypoprotocristid protostylid hypometacristid preprotocristid premetacristid protoconid metaconid

postmetacristid mesial arm of pliopithecine triangle mesostylid postprotocristid talonid basin pliopithecine triangle prehypocristid pre-entocristid distal arm of pliopithecine triangle entoconid hypoconid buccal cingulum

postentocristid posthypocristid

hypoentocristid prehypoconulid cristid postcristid hypoconulid distal fovea posthypoconulid cristid

Figure 5. Molar terminology used in this study (following Harrison and Gu, 1999). Right upper molar (top) and right lower molar (bottom). Mesial at the top. Not to scale.

Superfamily Pliopithecoidea Zapfe, 1961a size; upper premolars with a buccolingually compressed paracone; Family Crouzeliidae Ginsburg and Mein, 1980 P4 with a poorly developed lingual cingulum; M1 and M2 with Subfamily Crouzeliinae Ginsburg and Mein, 1980 relatively narrow crowns that taper slightly lingually, a narrow Genus Fanchangia gen. nov. trigon, a lingual cingulum weakly to moderately developed, and a distal basin with little or no wrinkling; marked size differential Etymology d Named after the county town of Fanchang in between M1 and M2;M3 relatively small; I relatively tall with a Anhui Province. 1 weakly developed lingual cingulum and lacking a lingual pillar; P3 Diagnosis d A genus of crouzeliid pliopithecoid of large dental with a small metaconid; P3 much smaller than P4;P4 moderately size, with an estimated mean body mass of ~12e14 kg. Lower narrow with a long and robust preprotocristid ending in a distinct molars slightly smaller in area than those of K. krishnaii and cf. protostylid and forming a mesial protuberance bordered by a rib- A. priensis and similar in size to those of Platodontopithecus jian- like buccal cingulum, the talonid much longer than the trigonid, ghuaiensis, but larger than those of all other pliopithecoids. Upper and relatively small distal tubercles; lower molars relatively long incisors relatively small compared with the length of the upper with rounded, conical cusps and elevated crests, a well-developed molars with a well-developed lingual cingulum and buccolingually pliopithecine triangle, and a weakly to moderately developed expanded crown base; upper canines with a moderate degree of buccal cingulum; M1 narrows mesially; M1 and M2 with a short bilateral compression and marked degree of sexual dimorphism in mesial fovea, an oblique distal wall to the mesial fovea, an elevated 8 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 trigonid in relation to the talonid basin, a moderately long and buccal cusps are rounded (sharp and flange-like in Krishnapithecus), slightly obliquely oriented cristid obliqua no subsidiary tubercles on the postmetacristid and preentocristid, a (postprotocristid þ prehypocristid), a relatively small hypoconulid shallower and less expansive talonid basin, a better developed placed slightly to the buccal side of the midline of the crown, and a pliopithecine triangle (only incipiently developed in Krishnapithe- moderately large distal fovea; marked size differential between M1 cus), the presence of crests separating the mesial and distal foveae 3 and M2; and M3 smaller than M2 (Table 3). from the talonid basin, and a shallower lingual notch; and M with the paracone and metacone less widely spaced, more rounded cusps, a lower and more rounded postparacrista and premetacrista 3.1. Differential diagnosis (sharp and flange-like in Krishnapithecus), a better developed crista obliqua, and a less expansive trigon basin. Fanchangia differs from Pliopithecus, Platodontopithecus, and Fanchangia differs from Laccopithecus in the following features: Dionysopithecus in the following features: I1 and I2 with a well- I1 relatively higher crowned with a lingual cingulum less apically developed lingual cingulum and buccolingually expanded crown positioned; I2 smaller in relation to I1; upper incisors larger in base (unknown for Platodontopithecus); upper incisors relatively comparison with the length of the upper molar series; upper and small compared with the length of the upper molar series (un- lower canines relatively lower crowned; upper molars narrower, known for Platodontopithecus); upper premolars with a buccolin- with a less pronounced buccal flare and tapering of the crown gually compressed paracone (more conical in pliopithecids and lingually, a less protuberant parastyle, a stronger lingual cingulum, dionysopithecids); P4 relatively narrower with a less well- a larger hypocone and distal basin, and greater size differential developed lingual cingulum (and greater cusp height differential between M1 and M2 (Table 3); M3 smaller than M2 (larger than M2 in dionysopithecids); M1 and M2 with relatively narrower crowns in Laccopithecus); lower incisors with more pronounced MD (except Dionysopithecus), narrower trigon, less prominent lingual waisting; P3 relatively narrower, with a narrower distal basin, cingulum, and distal basin with little or no wrinkling; I1 lacking a better developed metaconid (weak to absent in Laccopithecus), and lingual pillar with a weakly developed lingual cingulum (unknown stronger lingual cingulum; P3 much smaller than P4 (the average for dionysopithecids); P3 with a less well-developed lingual occlusal area of P4 is 144% of that of P3 in Fanchangia, whereas the cingulum; P4 relatively narrower in dionysopithecids and relatively lower premolars are subequal in area [99.7%] in Laccopithecus); P4 broader in Pliopithecus;P4 with a long and robust preprotocristid more ovoid, with a less pronounced mesial protuberance, and ending in a distinct protostylid and forming a prominent mesial weaker distal tubercles; lower molars with more conical cusps, a protuberance bordered by a rib-like buccal cingulum; lower molars shorter and broader mesial fovea, a less oblique distal wall of the longer relative to breadth; M1 narrows mesially; M1 and M2 with a trigonid, shorter and less oblique cristid obliqua, less pronounced more oblique distal wall to the mesial fovea, a more elevated trig- mesial protuberance and mesiobuccal cingular rib, less BL waisting, onid basin relative to the talonid basin, a relatively smaller hypo- a well-developed pliopithecine triangle (vestigial to absent in Lac- conulid and distal fovea, and a relatively weak buccal cingulum; copithecus), less marked differential in elevation between the and M3 with a more oblique distal wall to the mesial fovea. In trigonid and talonid; and M smaller than M (larger on average in addition, Fanchangia differs from Pliopithecus in having a relatively 3 2 3 Laccopithecus; Table 3). smaller M and M3 (except for Plio. piveteaui, which is similar to Fanchangia differs from Anapithecus in the following features: I1 Fanchangia in having a relatively small M3). relatively narrower; upper canine in presumed females less buc- Fanchangia differs from Krishnapithecus in the following fea- colingually compressed with a less prominent lingual cingulum; tures: M1 and M2 lower crowned with lower and more rounded upper premolars lack a well-developed parastyle and post- cusps (tall and mammilate in Krishnapithecus), crests linking the paracrista style; upper molars relatively slightly broader with less marked lingual tapering, lower and more rounded cusps and crests, Table 3 a larger protocone, a less well-developed and less protuberant Size differential of the upper and lower molar series in pliopithecoids.a parastyle; a shorter and less obliquely oriented mesial fovea, Taxon Lowers Uppers broader trigon, better developed lingual cingulum, less pronounced 3 1 2 3 buccal cingulum, and relatively smaller distal basin; M relatively M1:M2:M3 M :M :M 2 smaller in comparison with M (Table 3); I1 taller and narrower, Fanchangia jini 69:100:92 72:100:78 with a less prominent lingual cingulum; P3 much narrower, with a Dionysopithecidae more distally directed distolingual crest, weaker lingual and buccal Dionysopithecus shuangouensis 77:100:91 97:100:81 Platodontopithecus jianghuaiensis 68:100:90 84:100:96 cingula, lacking a distinct protostylid, and narrower distal basin; P4 Pliopithecidae narrower and more ovoid, with a shorter mesial fovea, a shallower Pliopithecus antiquus 78:100:108 distal basin, and less prominent distal tubercles; P3 much smaller Pliopithecus bii —:100:90 than P4 (the average occlusal area of P3 is only 70% of that of P4 in Pliopithecus canmatensis 82:100:126 81:100:— Pliopithecus piveteaui 76:100:96 Fanchangia, whereas the lower premolars are subequal in area Pliopithecus platyodon 86:100:116 91:100:90 [98%] in Anapithecus); M1 and M2 with lower and more rounded Pliopithecus vindobonensis 82:100:114 80:100:102 cusps, a shorter mesial fovea, a less pronounced mesial protuber- Pliopithecus zhanxiangi 76:100:115 85:100:95 ance and mesiobuccal cingular rib, a less elevated trigonid in Crouzeliidae relation to the talonid basin; shallower occlusal basins, a shorter Anapithecus hernyaki 92:100:115 93:100:91 Barberapithecus huerzeleri 80:100:109 88:100:97 and less oblique cristid obliqua, a larger hypoconulid that is slightly Crouzelia auscitanensis 84:100:— more buccally placed, and a larger distal fovea; greater size differ- Egarapithecus narcisoi 93:100:108 ential between M1 and M2; and M3 smaller than M2 (much larger Laccopithecus robustus 79:100:104 90:100:104 on average in Anapithecus; Table 3). a fi Size differential is calculated as the mean areas of the rst and third molars Fanchangia differs from Crouzelia in the following features: P4 expressed as a percentage of the mean area of the second molar. The values use longer and narrower (Table 4), with a longer talonid basin relative combined data from isolated teeth and associated molars. Sources: Bergounioux and to the mesial fovea; M and M with more rounded and voluminous Crouzel (1965); Ginsburg and Mein (1980); Harrison et al. (1991); Harrison and Gu 1 2 (1999); Moya-Sol a et al. (2001); Kordos and Begun (2001); Wu et al. (2003); Alba cusps and crests, a shorter and more restricted mesial fovea; a et al. (2010); Alba and Moya-Sol a (2012); authors' unpublished data). shorter and less obliquely oriented cristid obliqua, a shallower T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 9

Table 4 Fanchangia differs from Barberapithecus in the following fea- a,b Relative breadth of P4,M1, and M2 in pliopithecoids. tures: I1 relatively lower crowned with a buccolingually expanded

Taxon B/L index, P4 B/L index, M1 B/L index, M2 crown base; upper and lower canines in females lower crowned Egarapithecus narcisoi 73.0 78.9 73.7 and less buccolingually compressed with a less prominent Crouzelia rhodanica 75.0 cingulum; upper molars relatively narrower with more rounded Crouzelia auscitanensis 95.5 78.0 79.7 and voluminous cusps; greater size differential between M1 and M2 Barberapithecus huerzeleri 82.0c 80.7 79.7 3 2 and M much smaller than M (Table 3); P3 much smaller than P4 Plesiopliopithecus lockeri 84.0 78.3 (the average occlusal area of P is only 70% of that of P in Fan- Fanchangia jini 88.9 77.9 79.8 3 4 Laccopithecus robustus 88.9 83.8 80.0 changia, whereas the lower premolars are subequal in area [96%] in Anapithecus hernyaki 93.8 80.2 81.0 Barberapithecus); P4 more ovoid (subrectangular in Barberapithe- Pliopithecus vindobonensis 94.3 85.8 81.7 cus) with a narrower talonid basin and relatively smaller distal Krishnapithecus krishnaii 83.2 tubercles; lower molars with rounded and more voluminous cusps Dionysopithecus shuangouensis 84.8 86.0 83.3 Dionysopithecus orientalis 83.1 (not buccolingually compressed), a less protuberant mesial margin, Pliopithecus zhanxiangi 93.2 83.7 a shorter mesial fovea, and a less well-developed buccal cingulum; Platodontopithecus jianghuaiensis 85.9 84.8 84.3 greater size differential between M1 and M2; and M3 smaller than Pliopithecus canmatensis 91.7 85.4 87.3 M2 (M3 larger than M2 in Barberapithecus; Table 3). Pliopithecus antiquus 117.0 88.1 87.9 Pliopithecus bii 88.3 Pliopithecus piveteaui 87.2 88.7 3.2. Included species Pliopithecus platyodon 100.0 85.5 88.9 B ¼ breadth; L¼ length. Monotypic. Type species, Fanchangia jini sp. nov. a Sources: Zapfe (1961b); Bergounioux and Crouzel (1965); Ginsburg and Mein (1980); Harrison et al. (1991); Welcomme et al. (1991); Harrison and Gu (1999); Fanchangia jini sp. nov. Kordos and Begun (2001); Wu et al. (2003); Moya-Sol a et al. (2001); Alba et al. (2010); Alba and Moya-Sol a (2012); Chaimanee et al. (2015); Sankhyan et al. Etymology d Named in honor of Professor Changzhu Jin for his (2017); authors' unpublished data. important discoveries at Laili Mountain, including the renowned b B/L Index ¼ (buccolingual breadth/mesiodistal length) 100. Arranged in order Paleolithic site of Renzidong, as well as for his outstanding contri- of B/L index for M2. butions to the study of the Neogene mammals of China. c Estimated value (Alba and Moya-Sol a, 2012).

Holotype d IVPP V18114. Left M2. Unworn and well-preserved. See Fig. 6l. talonid basin, a slightly larger hypoconulid (at least on M2), the Type locality d Tangkou fissure, Laili Mountain, Fanchang, presence of a crest linking the hypoconulid and entoconid (absent Anhui Province, eastern China. 31 05.38ʹ N, 118 05.77ʹ E. in Crouzelia) and delimiting the distal fovea; and greater size dif- Age and distribution d Late Early Miocene, ~18-17 Ma. Known ferential between M1 and M2 (Table 3). only from the type locality. Fanchangia differs from Plesiopliopithecus in the following fea- Referred specimens d In addition to the holotype, 107 isolated tures: lower incisors lower crowned (apicobasal height 100/MD teeth are referred to this species. These include examples of all of breadth of I1 is 181.3 in Fanchangia versus 225.8 in Plesiopliopithe- the permanent dentition except I2, and several deciduous teeth cus), with a weaker lingual cingulum; P3 with a well-developed (SOM Table S1; Figs. S1 and S2). metaconid (absent in Plesiopliopithecus); P4 relatively broader d (Table 4); P3 much smaller than P4 (the average occlusal area of P3 is Diagnosis As for genus. only 70% of that of P4 in Fanchangia, whereas the lower premolars are subequal in area [98%] in Plesiopliopithecus); and M1 with lower 3.3. Description and more rounded cusps, a relatively shorter mesial fovea, a less protuberant mesial margin, a shorter and less oblique cristid obli- Dental morphology The I1 crown is tall, relatively narrow mesio- qua, a larger and more buccally placed hypoconulid, and a larger distally, and bilaterally symmetrical (Fig. 6aec). The apicobasal distal fovea. height/MD breadth index is 181.3. The crown has its greatest MD Fanchangia differs from Egarapithecus in the following features: breadth at the apex, and it tapers basally. The crown exhibits a P3 with a less well-developed metaconid (prominent in Egar- distinct MD waisting toward the base, giving the crown an apithecus); P4 relatively broader (Table 4) and more ovoid (crown inverted flask-shaped outline in buccal view. This feature is a rectangular in Egarapithecus), with less prominent distal tubercles, unique characteristic of pliopithecoids. The lingual face is concave and a less pronounced mesial protuberance and mesiobuccal cin- apically and convex basally. The mesial and distal marginal ridges gular rib; P3 much smaller than P4 as in Egarapithecus, but the and the lingual cingulum on the lingual face of the crown are differential being more marked in Fanchangia (the average occlusal weakly developed. There is no lingual pillar. The buccal face is area of P3 is only 70% of that of P4 in Fanchangia, whereas the value smooth and featureless. The root is incomplete, but the basal in Egarapithecus is 78%); lower molars with broader crowns portion is stout relative to the size of the crown. fl (Table 4), a greater degree of buccal are, more rounded cusps, a The two lower canines in the collection differ in size and distinct pliopithecine triangle (the lower molars of Egarapithecus morphology, and they presumably represent those of a female and are worn, but it is evident that the pliopithecine triangle was male individual, respectively (Fig. 6d, e). The size difference be- weakly developed or absent), a less oblique crest linking the pro- tween the two canines indicates a marked degree of canine size toconid and metaconid, a broader mesial fovea, a shorter and less dimorphism. The male:female mean MD length ratio is 1.45, similar oblique cristid obliqua, a less prominent mesial protuberance and to that of extant Gorilla gorilla (1.43) and Macaca mulatta (1.48; mesiobuccal cingular rib, a hypoconulid positioned closer to the Zhang and Harrison, 2017). The canine of the female individual is midline of the crown; a less well-developed buccal cingulum; smaller, with a relatively lower crown than the canine of the male greater size differential between M1 and M2; and M3 smaller than individual. The crown is ovoid in cross section with a moderate M2 (M3 larger than M2 in Egarapithecus; Table 3). degree of bilateral compression (breath/length index ¼ 71.9). In 10 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838

Figure 6. Lower dentition of Fanchangia. Representative teeth only (see SOM Fig. S1 for images of additional specimens). (aec) V 18115.26, right I1, (a) lingual view, (b) mesial view, (c) buccal view. (d) V 18115.30, right C1 (female), buccal view. (e) V 18115.31, left C1 (male), buccal view. (f) V 18115.32 left P3, buccal view. (g) V 18115.98, left P4, occlusal view. (h) V 18115.28, right P4, occlusal view. (i) V 18115.1, left M1, occlusal view. (j) V 18115.17, left M1, occlusal view. (k) V 18115.2, right M2, occlusal view. (l) V 18114 (holotype), left M2, occlusal view; (m) V 18115.10, left M3, occlusal view. (n) V 18115.11, right M3, occlusal view; (o) V 18115.88, left M3, occlusal view. (geo) Mesial margin at top. All to the same scale. occlusal view, the apex is placed centrally. The mesial crest is short, distobuccal tubercle. The distal heel is short. The lingual cingulum and it terminates at the junction with the lingual cingulum at a low is narrow, but well defined. It originates at the junction with the rounded tubercle. The distal crest and distolingual crest are long mesial crest and undulates distally around the distolingual crest to and rounded. The lingual cingulum is narrow, and it continues join the distal marginal ridge. A trace of a cingulum is also present around the distolingual crest to become continuous with the distal on the mesiobuccal face of the crown. The buccal face bulges marginal ridge. The distal heel is short, with a shallow distal basin. slightly around the base of the crown. The root, which is incom- The root is long and slender. The tip of the root is slightly rounded, plete, is relatively stout and bears a shallow groove on its lingual but it can be estimated to have been about twice as long apicoba- face. sally as the crown height. The P3 is relatively short and narrow (Fig. 6f). The protoconid is The canine of the male individual is tall and stout, with mod- tall, mesiodistally compressed, and placed slightly closer to the erate bilateral compression (breadth/length index ¼ 68.8). The mesial end of the tooth in occlusal view. The mesial crest is short crown is relatively strongly distally recurved. The sharp mesial crest and sharp. The mesiobuccal honing face of the crown is mesiodis- descends from the apex to end basally at a prominent lingual tally short and relatively tall and steep, which is a specialized cingulum. It is bordered distolingually by a shallow groove. The feature of pliopithecoids. The sharp distal crest is longer than the distolingual and distal crest are slightly worn but appear to have mesial crest and ends basally at the distal marginal ridge. The been low and rounded. The distal crest ends basally at an irregular distolingual crest is long and sharp with a small but prominent T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 11 metaconid. A narrow and rounded lingual cingulum originates at postentocristid arcs distally and buccally to join the distal marginal the basal end of the mesial crest and passes distally around the base ridge. The hypoentocristid, together with the postcristid, form a of the crown to reach the distolingual crest. The distal marginal well-developed distal transverse crest connecting the hypoconulid ridge is low and rounded and encloses a narrow crescent-shaped and entoconid. The distal transverse crest is oriented at about distal basin. 30e45 to the transverse axis of the crown. A shallow fissure in- The P4 is an ovoid or rhomboid-shaped tooth, longer than broad terrupts the distal transverse crest midway along its length and (mean breadth/length index ¼ 89.2; range ¼ 80.9e96.4) with an allows the distal fovea to communicate with the talonid basin. The obliquely oriented long axis (Fig. 6g, h). The two main cusps are distal fovea is a well-defined D-shaped basin subequal in area to the subequal in size. The protoconid is placed slightly more mesially mesial fovea. The talonid basin is long and narrow and relatively than the metaconid. The two cusps are connected by a short deep. The pliopithecine triangle forms a small accessory fovea in transverse crest divided in the midline by a longitudinal fissure. The the talonid basin between the protoconid and hypoconid that preprotocrista is relatively robust and often forms a distinct mesial opens lingually toward the center of the basin. A pliopithecine protuberance. The premetacrista is short and rounded. There is a triangle is present on all M1s except V18115.87. The floor of the narrow crease-like mesial fovea, which sometimes opens mesio- talonid basin is marked by a simple Y-shaped fissure pattern that lingually. The talonid basin is moderately short, ovoid and relatively deviates around the pliopithecine triangle, but is otherwise smooth shallow, with a smooth or finely wrinkled enamel surface. The and generally lacks secondary wrinkling. The buccal cingulum is talonid is much longer than the trigonid, with an average trigonid poorly developed and restricted to vestiges on the buccal side of the length/talonid length index Moya-Sol a et al., 2001 of 37 (n ¼ 5, cristid obliqua, on the mesiobuccal face of the protoconid, and on range ¼ 25e43). The postprotocristid is rounded and ends at a low the distobuccal aspect of the hypoconid. rounded distobuccal tubercle. Similarly, the postmetacristid, which The M2 is morphologically similar to M1 but differs in the is sharper than the postprotocristid, ends in a small distolingual following features: the crown is much larger (the occlusal area of tubercle. A rounded distal marginal ridge links the two tubercles. M1 is only 68.6% of that of M2) and relatively broader (mean Traces of a cingulum are present on the buccal aspect of the crown, breadth/length index ¼ 79.8; range ¼ 77.9e81.1); the cusps are with a distinct rib-like enamel fold on the mesiobuccal face. more voluminous; the mesial fovea is relatively broader; the cristid The M1 is subrectangular in occlusal outline with rounded cor- obliqua is less obliquely oriented; the buccal cingulum is slightly ners (Fig. 6i, j). It is relatively narrow and tapers slightly mesially. better developed; and the crown has greater buccal flare. As in M1, The mean breadth/length index is 77.9 (range ¼ 74.4e82.7). There the crown is subrectangular to ovoid, with rounded and conical is little or no BL waisting. The tooth is relatively high crowned with cusps and sharp and elevated crests (Fig. 6k, l). The mesial fovea is rounded conical cusps and a high degree of buccal flare (index of relatively elongated and separated from the talonid basin by an MF ¼ 0.53; Singleton, 2003). The occlusal crests linking the cusps obliquely oriented mesial transverse crest. Occasionally (e.g., are sharp and elevated. The protoconid is slightly taller and more V18114 and V18115.7), the protoconid and, less commonly, the voluminous than the metaconid. The preprotocristid is short and metaconid have an accessory transverse crest in addition to the arcs mesiolingually to become continuous with the mesial marginal mesial transverse crest, which descends into the mesial fovea. The ridge. The hypoprotocristid is short and passes transversely. The pliopithecine triangle is well developed. The hypoconid is larger postprotocristid originates from the apex of the protoconid and than the protoconid. There is relatively strong buccal flare (mean passes distolingually. It bifurcates midway along its length, with the MF ¼ 0.51; range ¼ 0.48e0.54). The hypoconulid and entoconid are buccal arm passing distobuccally to form the mesial portion of the relatively small, with an oblique distal transverse crest connecting cristid obliqua. The lingual arm is directed distolingually to form a them. The distal fovea is well developed and subequal in area to the well-developed mesial arm of the pliopithecine triangle. The met- mesial fovea. The lingual notch between the metaconid and ento- aconid is tall and conical. The premetacristid curves mesiobuccally conid is relatively shallow (mean LNH index ¼ 0.74; to join the elevated mesial marginal ridge. The mesial fovea is range ¼ 0.71e0.81). The buccal cingulum is variably developed, relatively elongated and narrow, only slightly broader than long, ranging from weakly to moderately developed, with several quite deep, and lenticular in shape. The hypometacristid joins the discontinuous traces around the buccal aspect of the crown. The hypoprotocristid at an elevated transverse crest linking the proto- talonid basin is long and narrow and relatively deep. It has a simple conid and metaconid and delimiting the posterior wall of the Y-shaped fissure pattern with occasional minor secondary mesial fovea. The floor of the mesial fovea is slightly to moderately wrinkling. more elevated than that of the talonid basin. The postmetacristid is The M3 is variable in size, but the occlusal area is slightly smaller long and directed slightly distobuccally. The hypoconid is larger on average than that of M2 (mean area of M3 is 92% of M2 area). The than the protoconid. The long prehypocristid forms a slightly crown is long and narrow, tapering distally, with a moderate degree oblique cristid obliqua. A low rounded mesiolingual crest originates of buccal flare (Fig. 6m, o). The breadth/length index is 70.9 from the apex of the hypoconid and passes into the talonid basin; (range ¼ 66.3e73.8). The mesial face of the crown is oblique, with a this forms the distal arm of the pliopithecine triangle. The post- slightly bulging mesiobuccal margin. The cusps are conical with hypocristid is short but well developed. well-developed crests connecting them. The protocone is low and The hypoconulid is small, forming a low conical cusp on the rounded; it is positioned close to the mesial margin of the crown, distal margin. It is positioned slightly to the buccal side of the resulting in a short preprotocristid. The postprotocristid is robust longitudinal midline of the crown. The prehypoconulid cristid is and rounded. The hypoprotocristid is sharp and elevated. Origi- short and meets the posthypocristid to form an elevated crest nating from the distolingual base of the protocone is a well- linking the hypoconulid and hypoconid. A short posthypoconulid developed mesial arm of the pliopithecine triangle. The meta- cristid curves distally and lingually to join the low distal marginal conid is smaller than the protoconid, with a well-developed pre- ridge. The postcristid is usually high and sharp, but this crest is metacristid, hypometacristid, and postmetacristid. The mesial absent in V 18115.87 as an atypical variant. The entoconid is small, transverse crest, comprising the hypometacristid and the hypo- being subequal in size to the hypoconulid. The preentocristid is protocristid, is elevated and slightly oblique relative to the trans- shorter than the postmetacristid. A narrow transverse fissure sep- verse axis of the crown. The mesial fovea is a narrow elliptical to arates the latter two crests and passes through the shallow lingual subcircular depression. It is bordered mesially by an elevated notch (mean LNH index ¼ 0.75, range ¼ 0.71e0.80). The mesial marginal ridge. Unlike on M1 and M2, the hypoconid is 12 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 smaller than the protoconid. The cristid obliqua is slightly obliquely side of the mesial crest is a slightly wrinkled mesial groove. The oriented relative to the long axis of the crown. The posthypocristid lingual pillar is well developed. A narrow and irregular cingulum is relatively short. A moderately well-developed distal arm of the passes around the base of the crown from the mesial crest to the pliopithecine triangle originates from the base of the hypoconid lingual pillar. There is a slight trace of a buccal cingulum basally on and passes obliquely into the talonid basin. The latter crest is less the mesiobuccal aspect of the crown. The distal crest is well well developed than the mesial arm of the pliopithecine triangle. developed and widely spaced from the lingual pillar. The promi- The hypoconulid is small, perched on the distal margin of the nent distal heel accommodates a broad and shallow distal basin crown, and buccally placed in relation to the midline long axis of that is bordered by a prominent distal marginal ridge. The root is the crown. In some case, the buccal cusps are mesiodistally aligned, relatively long (almost twice as long as the crown height) and but usually, the hypoconulid is slightly lingually offset. The pre- slender, with a shallow groove along its lingual face. hypoconulid cristid is short and ends at the junction with the The P3 is triangular in occlusal outline, tapering lingually posthypocristid. The posthypoconulid cristid arcs distally and (Fig. 7g, h). The crown is buccolingually much broader than long, lingually to become continuous with the distal marginal ridge. The with a mean breadth/length index of 69.1 (range ¼ 65.9e72.4). The postcristid is short, obliquely oriented, and forms the mesial wall of paracone is tall and buccolingually compressed. The protocone is the small distal fovea. Occasionally, this latter crest is absent or small, rounded, and peripherally placed on the mesiolingual weakly developed and the distal fovea and talonid basin are margin of the crown. It is much less elevated than the paracone, confluent. In several specimens (e.g., V18115.10, V18115.11, being only ~60% of the apicobasal height of the paracone. The V18115.18, V18115.25, V18115.88), low accessory transverse crests preparacrista and postparacrista are sharp and well developed; the originate from the hypoconid and/or entoconid and almost meet to latter is slightly shorter than the former. The preprotocrista is short produce a small triangular fovea in the distal part of the talonid and rounded and becomes continuous mesially with the rounded basin. The distal fovea is similar in size to the mesial fovea. Occa- mesial marginal ridge. The postprotocrista is low and weakly sionally, a small tuberculum sextum is present on the distolingual developed. In V18115.93, the postprotocrista terminates at a small margin of the crown. The buccal cingulum is poorly developed, tubercle on the distal marginal ridge. A prominent hypoprotocrista being restricted to traces on the mesobuccal face of the protoconid, passes buccally from the base of the protocone to meet the pre- between the protoconid and hypoconid, and on the buccal face of paracrista or the base of the paracone to form a mesial transverse the hypoconulid. The entoconid is small, being subequal in size to crest. It is positioned far mesially and runs parallel to the mesial the hypoconulid. The postentocristid and hypoentocristid are short. marginal ridge to delimit a short and broad mesial fovea. Distal The entoconid is well separated from the metaconid, so the transverse crests originating from the paracone and protocone are preentocristid and the postmetacristid are relatively long. The absent or weakly developed. The distal basin is broad and relatively latter two crests are separated by a transverse fissure that passes shallow, with occasional secondary wrinkling; an elevated distal through a shallow lingual notch. The talonid is narrow but well marginal ridge borders it distally. There is no lingual cingulum. In defined and relatively deep. There is a Y-shaped fissure system on V18115.93, prominent ribs on the mesiobuccal and distobuccal the talonid floor. Secondary wrinkling of the talonid basin and margin represent traces of a buccal cingulum. distal fovea is variable, ranging from absent to slight. The P4 is short and broad, with an ovoid occlusal outline that The I1 crown is relatively narrow (Fig. 7aec). The apicobasal tapers slightly distally (Fig. 7i, j). The paracone is buccolingually height is greater than the MD breadth, with a mean height/breadth compressed with a well-developed preparacrista and post- index of 135.0 (range ¼ 132.8e137.1). In buccal view, there is a paracrista of equal length. The preparacrista terminates mesially at slight basal waisting of the crown. The lingual face is spatulate, with a small tubercle or parastyle. The protocone is conical and volu- a slightly convex mesial margin and a more strongly convex distal minous but less elevated than the paracone. The low and rounded margin bordered by a prominent distal marginal ridge. The base of preprotocrista arcs mesially and buccally to become continuous the crown is robust, with a well-developed lingual cingulum that with the mesial marginal ridge. The postprotocrista is short and extends one-third to one-half of the way to the apex. The lingual sharp and passes distally to meet the distal marginal ridge. A face is finely crenulated, and the lingual pillar is either absent or rounded mesial transverse crest connects the paracone and pro- poorly developed. The root is long and slender, being more than tocone mesially to delimit a small fissure-like mesial fovea. Short twice the crown height. subsidiary crests from the protocone and paracone form an The I2 is low crowned and spatulate. It is relatively narrow incomplete and indistinct distal transverse crest. A deep S-shaped compared with the I1 (the mean MD breadth is only 67% of that of fissure courses longitudinally across the distal basin to divide the I1). The crown is bilaterally asymmetrical in buccal view with a latter two crests. The distal basin is shallow and relatively narrow rhomboidal outline. There is a mesial angulation about two-thirds and bordered distally by an elevated distal marginal ridge. A narrow from the crown base and a distal angulation about one-third from lingual cingulum is present on the lingual face of the protocone, the base. The mesial and distal margins meet apically at a ~90 being best developed distolingually. There is no trace of a buccal angle to form a pointed incisive apex. A moderately well-developed cingulum. and rounded cingulum loops around the base of the lingual face of The M1 is broader than long (mean length/breadth index ¼ 83.4; the crown connecting the mesial and distal angulations. There is no range ¼ 80.0e87.7), rectangular in occlusal outline, with a mod- lingual pillar. The buccal face is mesiodistally and apicobasally erate degree of lingual flare (Fig. 7km). The mesial and distal convex and featureless. The slender, bilaterally compressed root is margins of the crown are subparallel, or the crown tapers slightly about twice as long as the crown height. lingually. The protocone is pyramidal in shape and voluminous. The Two upper canines are known, and judging from their preprotocrista is rounded and passes obliquely mesiobuccally to morphology and size, they are both presumed to have belonged to meet the hypometacrista at a low protoconule. The postprotocrista female individuals. The canine is low crowned, with a triangular is sharp and well developed. The conical paracone is more elevated section that broadens distally (Fig. 7e, f). The crown is moderately than the protocone but less voluminous. The preparacrista is short bilaterally compressed, with an average breadth/length index of and sharp; it terminates at the mesial marginal ridge at a low, 84.7. The protocone is more or less centrally placed in occlusal view. rounded parastyle. The hypoparacrista is represented by a fine crest The mesial crest is short and terminates basally at a small tubercle running parallel to the mesial marginal ridge. The mesial fovea is perched on the mesial rim of the lingual cingulum. On the lingual very short mesiodistally and quite narrow. The postparacrista is T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 13

Figure 7. Upper dentition of Fanchangia. Representative teeth only (see SOM Fig. S2 for images of additional specimens). (aec) V 18115.64, left I1, (a) lingual view, (b) distal view, (c) buccal view. (d) V 18115.63, right I2, lingual view. (eef) V 18115.76, left upper canine (female), (e) buccal view, (f) lingual view. (geh) V 18115.93, left P3, (g) occlusal view, (h) mesial view. (iej) V 18115.70, right P4, (i) occlusal view, (j) mesial view. (k) V 18115.33, left M1, occlusal view. (l) V 18115.39, right M1, occlusal view. (m) V 18115.54, left M1, occlusal view. (n) V 18115.51, left M2, occlusal view. (o) V 18115.35, left M2, occlusal view. (p) V 18115.73, right M3, occlusal view. (g, i, kep) Mesial margin at top. All to the same scale. sharp and well developed. The metacone is conical and slightly short and sharp and arcs distolingually to become continuous with smaller than the paracone. The premetacrista and postparacrista the distal marginal ridge. The trigon basin is broader than long and are separated by a narrow transverse fissure that passes through a forms a deep and well-defined triangular basin. The floor of the shallow buccal notch. The hypometacrista passes mesiolingually to basin has a simple Y-shaped fissure pattern. The hypocone is low meet the postprotocrista to form the well-developed crista obliqua. and conical and much smaller than the protocone. The pre- There is no development of a metaconule. The postmetacrista is hypocrista passes obliquely mesiobuccally to end at the base of the 14 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 proptocone, close to its junction with the crista obliqua. The post- cingulum passes around the base of the triangular lingual face of hypocrista is short and curves distobuccally to join the distal the crown. The buccal aspect of the tooth is convex apicobasally and marginal ridge. The distal marginal ridge is low and sharp and mesiodistally. distally convex. The distal fovea is a large triangular basin, only The dP4 is similar in its basic morphology to M1, but smaller and slightly smaller in area than the trigon basin, with a smooth floor or lower crowned. It also differs in the following features: the crown is only minor wrinkling. In more than half of the specimens, a low and relatively narrower, with a more convex lingual margin; the crown rounded accessory transverse crest originates from the apex of the tapers more strongly lingually, with a relatively small protocone hypocone and passes into the distal fovea. The lingual cingulum and hypocone; the cusps are more conical with finer crests; the ranges from moderately to weakly developed. It is either contin- trigon is narrower; the lingual cingulum is less well developed, uous around the mesial and lingual aspect of the protocone and being restricted to a narrow and discontinuous ledge around the lingual face of the hypocone (27% of specimens) or discontinuous lingual aspect of the protocone; and the buccal cingulum is weaker. (73% of specimens). Traces of a buccal cingulum are present on the Dental size and body mass estimation In terms of dental size, F. jini mesiobuccal face of the paracone, the distobuccal face of the met- is one of the largest species of pliopithecoids. The lower molars are acone, and the buccal wall between the paracone and metacone. slightly smaller than those of K. krishnaii and cf. A. priensis and 2 1 The M is morphologically similar to M (Fig. 7n, o). It differs in similar in size to those of Plat. jianghuaiensis, but larger than those 1 2 being larger in overall size (mean M area is 72% of that of M ) and in all other pliopithecoids (Table 5). ¼ relatively slightly narrower (mean length/breadth index 84.1; Based on the mean areas of M1 and M2, the estimated body mass 2 range ¼ 79.0e89.5). In addition, M typically has a continuous is 11.89 ± 1.71 kg and 13.65 ± 1.57 kg, respectively (Table 5). 2 lingual cingulum (continuous in 100% of M s compared with only However, given that F. jini exhibits marked sexual dimorphism in 1 27% of M s), and the hypocone is relatively larger. canine size comparable with that of M. mulatta and G. gorilla (see 3 Unfortunately, all of the examples of M are worn, rolled, or the aforementioned information), it is reasonable to infer that the damaged, which partially obscures the morphological details mean body mass in males would have been considerably greater (Fig. 7p). Nevertheless, it is possible to discern the main anatomical than in females. Using body mass sexual dimorphism in M. mulatta 3 1 1 3 features. The M is larger on average than M (M area is 92% of M (median body mass ¼ ±17.9%) and G. gorilla (median body 2 3 2 area), but smaller than M (M is 78% of M area). The crown is mass ¼ ±38.3%) as models (Smith and Jungers, 1997), the estimated relatively narrow and tapers distally. The distal cusps are relatively body mass of female and male F. jini would be ~8e11 kg and reduced compared with the protocone and paracone. The proto- ~15e18 kg, respectively. If these estimates represent accurate ap- cone and metacone are connected by a low crista obliqua. The proximations, Fanchangia would have been most comparable in lingual cingulum is continuous around the lingual aspect of the body mass and degree of sexual dimorphism with Papio kindae crown but narrows distally. The distal basin is relatively small, with (mean female and male body mass are 9.8 kg and 16.0 kg, respec- a moderate degree of wrinkling. Traces of a buccal cingulum are tively; Delson et al., 2000) among extant catarrhines. present on the mesiobuccal aspect of the paracone and between the paracone and metacone. 4. Comparisons with other pliopithecoids and stem The dP4 is generally similar in morphology to M1, but smaller in catarrhines overall size. It also differs in the following features: the crown is relatively narrower; the crown tends to have a more prominent When compared with other stem catarrhines, such as proplio- mesial protuberance where the preprotoconid meets the oblique pithecids and dendropithecids, pliopithecoids are characterized by mesial margin; the cristid obliqua is more obliquely oriented; the a suite of craniodental and postcranial features that are best mesial transverse crest is more oblique; the trigonid is more interpreted as unique specializations of the clade (Andrews et al., elevated in relation to the talonid; the hypoconulid is tiny and 1996; Harrison, 2013). Dental specializations include the placed on the distal margin close to the midline; and the distal following: relatively slender lower incisors at least twice as high as fovea is small to vestigial. In addition, the buccal cingulum forms a their MD length (except Anapithecus, which has broader and more discontinuous ledge around the buccal aspect of the crown. The spatulate incisors); lower central incisors waisted toward the base pliopithecine triangle is variably developed. In V18115.15 and of the crown, giving the teeth a distinctive flask-shaped outline in V18115.21, the pliopithecine triangle has a well-developed mesial buccal view; P3 mesiodistally short and high crowned, with a arm in conjunction with a weaker distal arm. In V18115.20, the steeply inclined mesiobuccal face that does not extend far basally to pliopithecine triangle is absent. form an elongated honing flange, even in presumed males; P4 and The dI1 is mesiodistally broad and relatively low crowned. The lower molars relatively long and narrow; M1 and M2 with a rela- mesial margin is slightly convex, while the distal margin is mark- tively small hypoconulid; lower molars with a relatively long edly convex. The incisive apex is broad with a rounded distal angle. mesial fovea and a pliopithecine triangle in the talonid basin (the As in the permanent central incisors, the base of the crown is incidence of occurrence increases from M1 to M3; a pliopithecine relatively robust. The moderately well-developed lingual cingulum triangle is vestigial in Plio. vindobonensis and occurs variably or originates mesially close to the apex and then arcs obliquely around incipiently in Egarapithecus, Krishnapithecus, and Laccopithecus); the lingual face of the crown to end distally close to the base of the lower molars increase in size from M1 to M3 (except in dio- crown. A low lingual pillar is located just mesial to the midline axis nysopithecids, Plio. piveteaui, and possibly Plio. bii), with a marked of the lingual face. A smaller secondary pillar is situated slightly size differential between them (Table 3); M3 with buccal cusps more distally. In addition to the lingual pillars, the lingual face is arranged more or less in line; I1 slightly mesiodistally waisted near lightly wrinkled. the base of the crown; upper premolars and molars relatively short 1 The dC is low crowned and bilaterally compressed. The apex of and broad; and M2 (and often M3) considerably larger than M1 the protocone is situated in the center of the crown in occlusal view. (Table 3; Andrews et al., 1996; Harrison and Gu, 1999). Fanchangia Sharp mesial and distal crests originate from the apex of the pro- shares this suite of features with other pliopithecoids, except that tocone. A shallow mesial groove is located mesiolingually between M3 tends to be smaller in occlusal area than M2 (Table 3). These the mesial crest and the rounded lingual pillar. A weak and irregular features (especially hallmark features such as waisted lower T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 15

Table 5 a Comparison of dimensions of M1 and M2 and body mass estimates for Fanchangia and other pliopithecoids (in descending size order).

b Taxon M1 M2 Body mass estimate (kg)

2 2 n Area, mean (mm ) n Area, mean (mm ) Based on M1 area Based on M2 area Krishnapithecus krishnaii 1 75.1 14.44 Fanchangia jini 8 49.3 6 71.9 11.89 13.65 cf. Anapithecus priensis 1 53.8 13.55 Platodontopithecus jianghuaiensis 3 52.9 3 70.8 13.21 13.37 Pliopithecus zhanxiangi 1 49.6 3 67.5 12.00 12.56 Anapithecus hernyaki 4 49.3 3 58.9 11.89 10.50 Laccopithecus robustus 16 45.9 18 57.8 10.68 10.25 Pliopithecus bii 1 52.4 9.01 Pliopithecus platyodon 13 40.7 10 47.5 8.93 7.92 Pliopithecus vindobonensis 2 35.2 3 46.0 7.19 7.46 Pliopithecus piveteaui 2 34.7 1 43.3 7.03 7.02 Egarapithecus narcisoi 2 39.8 2 42.6 8.63 6.87 Pliopithecus antiquus 4 30.7 2 39.7 5.86 6.26 Dionysopithecus orientalis 1 28.9 5.35 Dionysopithecus shuangouensis 5 27.9 2 36.3 5.08 5.57 Pliopithecus canmatensis 7 28.4 7 34.7 5.21 5.25 Plesiopliopithecus lockeri 1 28.2 5.16 Crouzelia auscitanensis 1 27.1 1 32.6 4.86 4.84 Barberapithecus huerzeleri 1 26.2 1 32.6 4.62 4.84 Crouzelia rhodanica 1 23.5 3.15

a Sources: Zapfe (1961b); Bergounioux and Crouzel (1965); Ginsburg and Mein (1980); Harrison et al. (1991); Welcomme et al. (1991); Harrison and Gu (1999); Kordos and Begun (2001); Wu et al. (2003); Moya-Sol a et al. (2001); Alba et al. (2010); Alba and Moya-Sol a (2012); Chaimanee et al. (2015); Sankhyan et al. (2017); unpublished data. b Calculated using the regression formulas of Gingerich et al. (1982).

incisors, mesiodistally short and apicobasally tall P3 with a steep Barberapithecus), and a somewhat larger hypoconulid that is mesiobuccal honing face, and a pliopithecine triangle on the lower slightly more buccally placed (except Egarapithecus). molars) offer compelling evidence to support the inclusion of Comparisons with the upper molars of other crouzeliids are Fanchangia in Pliopithecoidea. more limited because these teeth are only known for Anapithecus, Moreover, among pliopithecoids, Fanchangia shares distinctive Laccopithecus, and Barberapithecus. However, as in the lower mo- features of the dentition with the Crouzeliidae (that exclude it from lars, Fanchangia retains a more primitive pattern than the other Pliopithecidae, Dionysopithecidae, and Krishnapithecidae). These crouzeliids. The upper molars of Fanchangia resemble dio- features include the following: I1 and I2 with a well-developed nysopithecids and pliopithecids in having more rounded and vol- lingual cingulum and buccolingually expanded crown base; upper uminous cusps and more rounded crests, a less well-developed and incisors relatively small compared with the length of the upper less protuberant parastyle, a less lingually tapering crown with a molar series; upper premolars with a buccolingually compressed more mesially placed protocone, a shorter preprotocrista, and a paracone (more conical in pliopithecids and dionysopithecids); P4 better developed lingual cingulum. with a less well-developed lingual cingulum; M1 and M2 with In addition, Fanchangia retains a number of primitive features of relatively narrow crowns (except Barberapithecus), a narrow trigon, the lower premolars shared with dionysopithecids, pliopithecids, a less prominent lingual cingulum, and a distal basin with little or and crouzeliines that differentiate it from the more derived con- no wrinkling; I1 lacking a lingual pillar with a weakly developed dition in anapithecine crouzeliids: P3 is narrower and lacks a lingual cingulum; P4 with a long and robust preprotocristid ending distinct protostylid, and P4 is more ovoid, with a less pronounced in a distinct protostylid and forming a prominent mesial protu- mesial protuberance, a shorter mesial fovea, and weaker distal berance bordered by a rib-like buccal cingulum (except Crouzelia tubercles. auscitanensis); lower molars long relative to breadth (Table 4); M1 From these comparisons, it can be inferred that Fanchangia narrowing mesially; M1 and M2 with an oblique distal wall to the shares key apomorphic traits with crouzeliid pliopithecoids and mesial fovea, a more elevated trigonid basin in relation to the that it is most parsimoniously included as a member of this clade. talonid basin, a relatively small hypoconulid placed close to the However, Fanchangia retains a suite of primitive pliopithecoid midline of the crown (more buccally placed in Egarapithecus and features that indicate that it is a stem member of the clade and Crouzelia rhodanica), a relatively small distal fovea (lacking the represents the sister taxon of all other crouzeliids. This inference is mesial wall in Crouzelia), and a relatively weak buccal cingulum; further supported by the phylogenetic analysis presented in the and M3 with a more oblique distal wall to the mesial fovea (char- following section. acteristic features adapted from Andrews et al., 1996; Harrison and It is pertinent here to reference the pliopithecoid isolated upper Gu,1999). This complex of dental features shared with the currently molar from the late Middle Miocene (12.1 Ma) of Siziwangqi recognized genera of crouzeliids provides strong support for the (Damiao 01) in Inner Mongolia (Zhang and Harrison, 2008; Qiu inclusion of Fanchangia in the Crouzeliidae. et al., 2013; Kaakinen et al., 2015). The crown is identical in size However, Fanchangia differs from all other crouzeliids, and ap- and proportions to M1sofFanchangia, and the overall morphology proximates the more primitive condition in dionysopithecids and is similar. As noted by Zhang and Harrison (2008), the Damiao pliopithecids, in several key features of the lower molars. These molar shares a number of morphological features with crouzeliids include more rounded and voluminous cusps (rather than sharp (i.e., a relatively narrow crown that tapers lingually, a well- and buccolingually compressed), a shorter mesial fovea, a less developed protuberant parastyle, an incompletely developed pronounced mesial protuberance and mesiobuccal cingular rib, a lingual cingulum, and a distal basin with minimal wrinkling). We less elevated trigonid in relation to the talonid basin, shallower concur with the initial assessment of Zhang and Harrison (2008) occlusal basins, a shorter and less oblique cristid obliqua (except 16 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 that the Damiao specimen probably belongs to a primitive crou- zeliid that is less advanced than Anapithecus and Laccopithecus. Comparisons of the Damiao upper molar with that of Fanchangia serve to highlight important differences. In Fanchangia, the M1 crown does not narrow so markedly lingually; the cusps are more rounded and less buccolingually compressed; the protocone is less distally displaced; the parastyle is much less protuberant; the hypocone is relatively smaller; the lingual cingulum is better developed and more continuous around the lingual aspect of the AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusCrouzeliaPlesiopliopithecusLaccopithecusAnapithecusEgarapithecus protocone; the buccal cingulum is not as prominent. From the limited evidence available, we infer that the Damiao specimen is a crouzeliid that was more derived than Fanchangia, but more primitive than Anapithecus and Laccopithecus, and similar in grade to Barberapithecus (which is a relatively primitive crouzeliid and shares the greatest similarity in its upper molar morphology to the Damiao specimen). Clearly, additional specimens are needed to determine the precise taxonomic affinities of the Damiao pliopi- thecoid, but the currently available evidence indicates that it belonged to an unnamed genus and species of crouzeliid.

5. Phylogenetic relationships

The results of the phylogenetic analysis are illustrated in

Figures 8 and 9. Six most parsimonious trees with equal length Figure 9. Phylogenetic relationships of Fanchangia. Strict consensus tree derived from (tree length ¼ 58) were acquired based on the Wagner parsimony the most parsimonious Wagner trees (see Fig. 8). criterion (Fig. 8). All of the most parsimonious trees and the resulting strict consensus tree (Fig. 9) place Fanchangia as the sister taxon to all other crouzeliids. Among the crown crouzeliids, the of relationships implies that the Crouzeliinae, now including Fan- highly specialized anapithecines d Anapithecus, Laccopithecus, and changia, should best be construed as a paraphyletic clustering of Egarapithecusd are united together as a terminal clade. Barbera- primitive crouzeliids. The placement of Pliopithecus as the sister pithecus, which Alba and Moya-Sol a (2012) view as a primitive taxon to all other pliopithecoids is somewhat surprising given the anapithecine, falls outside this clade as the sister taxon to the weight of compelling evidence presented by Harrison and Gu remaining crouzeliids. Barberapithecus and (1999) showing that Dionysopithecus and Platodontopithecus Crouzelia þ Plesiopliopithecus are successive sister taxa to the ana- represent a basal pliopithecoid clade. Moreover, as reflected in the pithecines, thereby forming a basal Hennigian comb. This latter set most parsimonious trees (Fig. 8), the integrity of the

AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusCrouzeliaPlesiopliopithecusLaccopithecusAnapithecusEgarapithecus AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusCrouzeliaPlesiopliopithecusLaccopithecusAnapithecusEgarapithecus AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusPlesiopliopithecusCrouzeliaLaccopithecusAnapithecusEgarapithecus

a b c

AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusCrouzeliaPlesiopliopithecusLaccopithecusAnapithecusEgarapithecus AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusCrouzeliaPlesiopliopithecusLaccopithecusAnapithecusEgarapithecus AegyptopithecusDendropithecusPliopithecusPlatodontopithecusDionysopithecusFanchangiaBarberapithecusPlesiopliopithecusCrouzeliaLaccopithecusAnapithecusEgarapithecus

d e f

Figure 8. Phylogenetic relationships of Fanchangia. Six most parsimonious trees (tree length ¼ 58) based on Wagner parsimony criterion. T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 17

Neogene Europe China Chinese European Time Scale Age Polarity pliopithecoids pliopithecoids (Ma) Chron Epoch Stage/Age LMA MN LMA C2 17 Nihewanian 2.59 16 3 Piacenzian Vilanyian Maze- C2A gouan 3.60 15 4 Pliocene ian

Zanclean Gao- zhuang-

Early Late 14 5 C3 Ruscinian 5.33 5.3

6 Anapithecus priensis Messinian 13

C3A Laccopithecus robustus + cf. Baodean 7

Turolian ? C3B 7.25 12

8 C4 11 Egarapithecus narcisoi Plesippliopithecus lockeri Pliopithecus vindobonensis + Late

9 +

C4A Miocene

10 Pliopithecus bii Tortonian Bahean + 10 Anapithecus hernyaki Vallesian

9 Barberapithecus huerzeleri Crouzelia rhodanica

C5 Damiao + Laogou Pliopithecus canmatensis 11 11.2 Pliopithecus antiquus 11.63 12 7+8 Pliopithecus platyodon Crouzelia auscitanensis C5A Serravallian Platodontopithecus jianghuaiensis 13 Pliopithecus zhanxiangi C5AA + C5AB 6

13.82 Astaracian 14 C5AC Middle Pliopithecus piveteaui C5AD Miocene Langhian 15 C5B 5 16 15.97 C5C

4 Fanchangia jini 17 C5D 18 Dionysopithecus shuangouensis Orleanian C5E Burdigalian 3 Shanwangian Tunggurian 19 C6 19.5 Early

20 Miocene 20.44 C6A 2 21 C6AA

Aquitanian Xiejian

22 Agenian C6B 1 23 C6C 23.03

Figure 10. Chronological distribution of Eurasian pliopithecoids. Sources: Harrison et al. (1991); Andrews et al. (1996); Mein (1999); Moya-Sol a et al. (2001); Wu et al. (2003); Deng (2004); Deng et al. (2004); Alba et al. (2010, 2019); Hilgen et al., (2012); Alba and Moya-Sol a (2012); Qiu et al. (2013); Qiu and Qiu (2013); Kaakinen et al. (2015); Pinero~ et al. (2018). LMA, Land Mammal Age; MN, Mammal Neogene unit.

Dionysopithecidae as monophyletic lacks universal support; Dio- The cladogram illustrated in Figure 9 is consistent with our nysopithecus and Platodontopithecus are either placed as sister taxa understanding of the distribution and polarity of morphological to each other (Fig. 8ac) or placed as successive sister taxa to the features presented in greater detail previously, and we regard this crouzeliids (Fig. 8df). as the most plausible set of relationships among the 18 T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838

Pliopithecoidea. The recognition that Fanchangia represents the 7. Conclusions primitive sister taxon of all other crouzeliids has important impli- cations for understanding the zoogeographic relationships and A collection of isolated teeth of a large pliopithecoid recovered evolutionary history of the pliopithecoids. from a fissure filling at Laili Mountain, Fanchang, China, is described here as a new genus and specie dF. jini. Comparisons demonstrate that Fanchangia can be distinguished from all other pliopithecoids by its unique suite of dental features. The species has an estimated 6. Zoogeographic relationships body mass of ~12e14 kg, which exceeds that of all other pliopi- thecoids except K. krishnaii and cf. A. priensis. Detailed comparisons The earliest pliopithecoids, the dionysopithecids, are known and the results of a phylogenetic analysis indicate that Fanchangia is from Sihong in eastern China, dating to ~19e18 Ma (equivalent to a stem member of the Crouzeliidae and represents the primitive MN 3; Harrison and Gu,1999). Although there is a good fossil record sister taxon of all other known crouzeliids. The recognition of a new to document the Early Miocene (MN 3e4) of Europe, there is no species of pliopithecoid provides additional evidence to document evidence of the presence of pliopithecoids. The implication is that the remarkable diversity of the clade in China and elsewhere in pliopithecoids initially entered Asia from Africa via the Arabian Asia. Peninsula and followed a circumeIndian Ocean route into sub- Fanchangia is one of the oldest known pliopithecoids, dating to tropical and tropical regions of south and eastern Asia (Bernor et al., ~1817 Ma (late Early Miocene based on correlations of the rodent 1988; Tassy, 1989; Rogl,€ 1999). The presence of Fanchangia, astem fauna); only the dionysopithecids from Sihong in eastern China are crouzeliid in China at ~18e17 Ma, well before the appearance of older (~1918 Ma). The occurrence of Fanchangia in China at least crouzeliids in Europe, supports the inference that pliopithecids and three million years before the appearance of crouzeliids in Europe crouzeliids diverged in Asia and migrated into Europe during the supports the inference that eastern Asia was a key center for the late Early Miocene or Middle Miocene (Fig. 10). The primitive fea- initial radiation of pliopithecoids and, furthermore, that pliopi- tures of the dentition retained by Fanchangia indicate the crouze- thecids and crouzeliids diverged in Asia and migrated into Europe liids initially diverged from a pliopithecid- or dionysopithecid-like during the late Early Miocene or Middle Miocene. ancestor. The earliest pliopithecid in Europe (and in Eurasia) is Plio. Pliopithecids become extinct in Eurasia by the early Late piveteaui from the Loire region of central France (MN 5, Miocene (~11 Ma), whereas the more specialized crouzeliids ~16e15 Ma), and diverse species of Pliopithecus became well continued in Europe until the middle of the Late Miocene (~9 Ma). established across Europe and China by MN 6 onward. The extinction of pliopithecoids in Europe coincides with cooler The earliest crouzeliids in Europe, Crouzelia, Plesiopliopithecus, conditions, increased seasonality, and a shift to predominantly and Barberapithecus (MN 6e8, ~14e11.2 Ma), are all more derived deciduous broad-leaved woodlands and grasslands. Laccopithecus than Fanchangia and occur at least three million years after the robustus, a late-surviving crouzeliid (~7 Ma), persisted in the sub- latter genus appeared in China. Although known only from an tropical forests of southern China well after the extinction of plio- isolated upper molar, the pliopithecoid from Damiao (MN 7 þ 8, pithecoids in Europe. Laccopithecus appears to be more closely ~12.1 Ma) in northeastern China appears to represent a primitive related to the specialized anapithecine crouzeliids from Europe crouzeliid (Zhang and Harrison, 2008), and if so, it would indicate than to Asian stem crouzeliids, which implies that Laccopithecus that crouzeliids continued to diversify in Asia during the Middle migrated to China during the later Miocene, rather than being Miocene. Pliopithecids become extinct in Eurasia by the end of the derived autochthonously from a lineage with deep ancestral roots Astaracian (MN 7 þ 8, ~11.2 Ma), whereas specialized crouzeliids in Asia. continued in Europe into the Vallesian (MN 9 and MN 10, ~119 Ma; Fig. 10). The anapithecine crouzeliid, Egarapithecus, from Declaration of competing interest the Late Miocene of Spain, is the last surviving pliopithecoid in Europe at ~9 Ma (Alba et al., 2010). The extinction of pliopithecoids The authors declare that there is no conflict of interest. in Europe corresponds to the onset of cooler conditions and increased seasonality, which led to a shift from subtropical ever- Acknowledgments green woodlands to predominantly deciduous broad-leaved woodlands and grasslands (Fortelius et al., 2014). Laccopithecus This work was supported by the Strategic Priority Research robustus from Shihuiba in Yunnan (~7 Ma) survived in the sub- Program of the Chinese Academy of Sciences (grant no. tropical forests of southern China well after the extinction of Eu- XDB26000000), the State Key Laboratory of Palaeobiology and ropean pliopithecoids and was the last surviving pliopithecoid. Stratigraphy (Nanjing Institute of Geology and Palaeontology, CAS; Laccopithecus is a specialized crouzeliid with close affinities to the grant no. 173132, 183122), the People's Government of Fanchang European anapithecines, Anapithecus and Egarapithecus. The avail- County, Anhui Province (The Excavations and Comprehensive able evidence indicates that this latter group (as well as its suc- Research Project on Paleolithic Site of Renzidong Cave in Fanchang cessive sister taxa, Plesiopliopithecus, Crouzelia, and County, Anhui Province), and an NYU travel grant. We would like to Barberapithecus) originated and diversified in Europe during the express our profound gratitude to Prof. Changzhu Jin of the Insti- Late Miocene. If so, the lineage leading to Laccopithecus was prob- tute of Vertebrate Paleontology and Paleoanthropology (IVPP) for ably derived from an ancestral anapithecine in Europe that inviting us to describe this important material and for supporting migrated to China in the later Miocene, rather than from a lineage our research. We are grateful to the Anhui team members for their that had deep ancestral roots among the known Asian crouzeliids. efforts and endeavors in the field. We thank the directors and staff The fossil record of pliopithecoids from China provides impor- of the following institutions for access to fossil material in their tant insights into the evolutionary history and zoogeography of the care: American Museum of Natural History, New York, USA; Beijing group. It shows that eastern Asia was an important center for the Museum of Natural History, Beijing, China; Geological Survey, early diversification of the pliopithecoids before the arrival of Bangkok, Thailand; IVPP, Beijing, China; Institut Catala de Paleon- pliopithecids and crouzeliids in Europe and that the subtropical tologia Miquel Crusafont, Barcelona, Spain; Universalmuseum € regions of southern China continued as a refugium after the demise Joanneum, Graz, Austria; Magyar Allami Foldtani€ Intezet, Budapest, of pliopithecoids in the rest of Eurasia. Hungary; Museum National d’Histoire Naturelle, Paris, France; T. Harrison et al. / Journal of Human Evolution 145 (2020) 102838 19

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