Journal of Human Evolution 57 (2009) 229–240

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Journal of Human Evolution

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New discoveries of blacki teeth from Chuifeng Cave in the Bubing Basin, Guangxi, south China

Wei Wang

Natural History Museum of Guangxi Zhuang Autonomous Region, Nanning 530012, P. R. China article info abstract

Article history: Ninety two Gigantopithecus blacki teeth have recently been recovered in situ at Chuifeng Cave in the Received 10 July 2008 Bubing Basin, Guangxi, south China. The hominoid teeth are associated with a typical early Accepted 9 February 2009 fauna. In comparison with Gigantopithecus of known provenience, for which classification is problematic, the Chuifeng sample can be allocated definitively to G. blacki. The new collection represents the largest Keywords: sample of this known, with the exception of the material from Liucheng. Statistical analysis of the Bubing Basin fauna indicates that G. blacki is one of the dominant elements, comprising 9% of the fauna. Chuifeng Cave Eleven teeth (12% of the sample) of G. blacki are diagnosed as having caries. In addition, wear on M Gigantopithecus blacki 3 early Pleistocene shows that G. blacki was adapted to consume tough or fibrous food and this wear may potentially imply relative longevity. Further study of this large sample of Gigantopithecus will provide additional insight into the paleobiology of this extinct hominoid. Ó 2009 Elsevier Ltd. All rights reserved.

Introduction The cave is located at an elevation of about 227 m above sea level and 11 m above the Mohui cave, which has also produced G. blacki Gigantopithecus blacki was first named by von Koenigwald in teeth and an associated late Pliocene to early Pleistocene 1935 based on a huge lower third molar found in a pharmacy in mammalian fauna. Five weeks of excavation took place from June to Hong Kong (von Koenigswald, 1935). Subsequent in situ discoveries July in 2006. As a result, 92 G. blacki teeth were recovered, associ- were made at nine karst cave localities from 1956 to 2006, ated with 915 other mammal teeth. including Daxin, Liucheng, Wuming, Bama, Jianshi, Longgupo, Mohui, and Bijie in southern China, and Tham Khuyen in north Geology and chronology Vietnam. In addition, G. giganteus was found in India (Table 1). The previous collections of G. blacki consist of three fragmentary The Chuifeng Cave, situated in the south-east part of the Bubing mandibles and 1057 isolated teeth from Liucheng and 75 isolated Basin (233402700 N, 1070002200 E) in west Guangxi, south China teeth from eight other localities. Based on dated localities (Huang (Fig. 1), is located in a limestone tower about 77 m above the local et al., 1995; Ciochon et al., 1996; Rink et al., 2008) and biostrati- valley floor near Hetang village and 16 m below the hill top (Fig. 2). graphical analysis (Huang, 1979; Han and Xu, 1985), G. blacki is Chuifeng Cave penetrates the hill from southeast to northwest, estimated to have lived during the early to middle Pleistocene, being 19 m in length, 0.5–2 m in width and 1.5–5 m in height. The approximately from 2 to 0.3 Ma. cave is partially filled with fossiliferous sandy-clay and a few The sequence of mammalian , including hominoids, at limestone breccias, with an average depth of 1.3 m. The sediment is karst caves in the Bubing Basin has been systematically studied. Our unusually intact in this cave, unlike most of the other south China previous research indicates that there are at least six levels of caves caves in which the sediments have been largely removed by in the basin that vary by elevation. Four of the caves exhibit well- collectors or for fertilizer. All of the fossil teeth and fragmentary documented faunal changes that relate to climate change, biogeo- bones derive from sediment at the northeast part of the cave. graphic events, and evolutionary change over the past two million The Chuifeng Cave mammalian fauna consists of 24 species years (Wang et al., 2007). (Figs. 3 and 4). Among this fauna, Hystrix magna, Sinomastodon sp., Here, the author reports findings from a recently discovered and Stegodon preorientalis, Ailuropoda microta, Pachycrocuta licenti, carefully excavated locality at Chuifeng Cave in the Bubing Basin. Tapirus sanyuanensis, and Sus peii are typical early Pleistocene elements in south China, similar to the Liucheng and Longgupo faunas (Pei, 1987; Han, 1987; Huang and Fang, 1991). The Chuifeng E-mail address: [email protected] fauna can be considered to be contemporaneous with these early

0047-2484/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2009.05.004 Table 1 History of Gigantopithecus research.

Site Date of discovery Identification Summary References 1–2 Daxin 1956 Three teeth (P3,M ,M3) The first locality to yield in situ G. blacki in southwest Guangxi. This Pei and Woo, 1956 cave is also called Heidong (dark cave), where three Gigantopithecus teeth were recovered from reddish clay deposits.

Liucheng 1956–1963 Three mandibles and 1006 This cave, also named Gigantopithecus cave, was discovered by Woo, 1962 1 2 1 teeth (11I ,7I1,6I ,6I2, 24C , a local farmer, and subsequently excavated by Pei and his field team. 3 4 27C1, 65P , 89P3, 104P ,100P4, Comprises the largest sample of G. blacki associated with numerous 174M1–2, other mammalian specimens. 1–2 2 140M ,141M3, 103M3 2DM , 7DM2)

3 4 Wumin 1965 Twelve teeth (4P ,2P,1P3, Located on a mountainside in Bulali, a suburb of Nanning City, was Zhang et al., 1973 1–2 2M ,1M1–2,2M3) first found by a hydrogeological survey team, and followed by test excavation by IVPP the same year. G. blacki teeth and a small number of other mammal fossils were collected from a hard yellow sandy- clay.

Bama 1973 One M3 Discovered by the survey team that discovered Wumin. Zhang et al., 1975

Jianshi 1970, 1999–2000 Twenty-seven teeth (1I1,1C1, Located in west Hubei Province in central China. Initially discovered Xu et al., 1974; Gao, 1975; 3 4 1 2 3 1P ,1P ,3M,2M ,2M,4C1, and excavated by a team from IVPP in 1970, and more recently Zhang et al., 2004 2P3,5P4,2M1,2M2,2M3) excavated by a joint IVPP-Hubei Museum team in 1999 and 2000. Some teeth had been assigned to Australopithecus or Meganthropus, however, the phylogenetic position of these early Asian hominids are actually ambiguous and disputed.

3 4 Longgupo 1984–1988 Fourteen teeth (1C1,1P ,2P, Discovered and excavated by Huang and his team from 1984 to 1988. Huang et al.,1995; Schwartz 3P3,3P4,4M1–2) The fauna also includes a mandibular fragment and an isolated and Tattersall, 1996; Wu, incisor originally allocated to Homo. The mandible is now widely 2000 considered to be that of an ape, closely related to the hominoid Lufengpithecus.

1 3 Mohui 2002–2004 Sixteen teeth (1I1,2C,1C1,3P, Excavated from 2002 to 2004. The site has yielded 16 G. blacki teeth Wang et al., 2005, 2007 1–2 3 1P3,5M1–-2,1M ,1M ,1M3) and 25 species of other .

Bijie 2006 One P4 An isolated P4 of Gigantopithecus was found by IVPP paleontologists Zhao et al., 2006 in a drugstore in Bijie, west Guizhou Province, southwest China in 1964, but its provenience is unknown. Fortunately, when the author of this paper visited Bijie in November 2006, he recognized five G. blacki teeth, including two incisors, one canine, one upper and one lower molar. These fossils were found by a local fossil collector from a quarry at Baeryan, located 12 km west of Bijie City.

Tham Khuyen 1996 One C1 Located in northern Vietnam. The site is well-known for its co- Ciochon et al., 1996 occurrence of H. erectus and G. blacki . The canine of Gigantopithecus is morphologically and metrically similar to those at Liucheng.

India 1969 One mandible A mandible found in deposits near Haritalyangar was originally Simons and Chopra, 1969; described as G. bilaspurensis (Simons and Chopra,1969) and ascribed Simons and Ettel, 1970; to the upper levels of the Dhok Pathan zone. Later, Szalay and Szalay and Delson, 1979 Delson (1979) assigned it to G. giganteus.

Figure 1. Map of G. blacki localities and plan of the Chuifeng Cave. W. Wang / Journal of Human Evolution 57 (2009) 229–240 231

Figure 2. Landscape in the vicinity of Chuifeng Cave in the Bubing Basin. The arrow points to the cave.

Pleistocene sites based on this biostratigraphic evidence. The fauna molars and a simple occlusal pattern, differ from the low crowned is also similar to that of Mohui, which is located in the same valley and wrinkled molars of Pongo, as well as the very large teeth of G. 500 m to the northwest (Wang et al., 2007). The elevation of blacki. These teeth are provisionally assigned to Hominoidea gen. et Chuifeng is about 11 m higher than the cave at Mohui, possibly sp. indet. There are 90 teeth of very large size that can be allocated representing an earlier cave formation event and sedimentary to Gigantopithecus. In addition, two relatively small incisors can be infilling. The appearance of Equus in Chuifeng may imply the assigned to this as well. Of the 1007 mammalian teeth, occurrence of grassland environments. Gigantopithecus constitutes a relatively common element, comprising 9% of the total fauna (Fig. 4). Eleven teeth of Gigan- Methods topithecus have been identified as exhibiting caries (about 12% of the collection). Among the Gigantopithecus sample, 77 teeth have Excavation was focused on the northwest part of the Chuifeng well-preserved crowns, but their roots are mostly missing due to Cave. The poorly cemented deposits consist mainly of light-brown porcupine gnawing (Figs. 5 and 6). Brief descriptions follow. sandy clay with a few limestone breccias. On the basis of the distribution of the sediment, four excavation sites were designated Upper canines as Areas A, B, C, and D. An area of 5.5 m2 was excavated down to limestone bedrock in 10 cm intervals. Fossils were mapped, recor- The crown of two left C1s (Z008535; Fig. 5-[5] and Z008536; ded, and collected, and about 100 kg of sediment were wet-sieved Fig. 5-[6]) are in good condition, but the root of Z008535 is missing. through 0.5 mm mesh screens at a stream at the foot of the hill to Z008536 preserves a small cervical portion of the distolingual part gather microfossils. In the dark inner cave, a gas-powered gener- of the root. Z008535 is especially robust, being nearly twice as large ator was used for lighting. An Electric Distance Measure was used to as Z008536. The crowns are relatively low, slightly worn, and make the measurements. conical in shape, with a height of 23.81 mm and 14.55 mm corre- Comparisons are made between the Chuifeng sample and G. sponding to MD of 20.80 mm and 14.79 mm. The buccal surface of blacki specimens from late Pliocene to middle Pleistocene localities the crown is strongly convex. The lingual surface is slightly bulging. in China, as well as the G. giganteus mandible from the Miocene of The distal edge is comparatively sharp and the mesial side is blunt. north India (Simons and Ettel, 1970; Miller et al., 2008). Gigan- The base of the crown is surrounded by a broad cingulum, most topithecus blacki specimens from drugstores are not included in the prominently developed lingually. Two upper canines present clear comparison because their lack of provenience may potentially sexual dimorphism: the Z008535 is likely male, while Z008536 is mask important temporal and spatial variation. female. A coefficient of variation was calculated for each dimension of the G. blacki teeth of known provenience. These data were Upper third premolars compared with the Chuifeng sample and the statistical significance between samples was examined using t-tests. The null hypothesis The upper third premolar is represented by six teeth, including was rejected if p < 0.05. four left P3s (Z008585-8) and two right P3s (Z008589-90; Fig. 5-[8– 13]). The crowns are well-preserved, but the roots are mostly Hominoid sample setting and description missing. Z008588 and Z008590 are slightly worn, Z008589 is heavily worn, and the remaining three teeth are moderately worn A total of 117 isolated hominoid teeth have been identified to expose a relatively large dentine area on the apex of the paracone among the Chuifeng mammalian remains. One small tooth and smaller one on the summit of the protocone. All of these P3sare resembles that of extant Hylobates and can be assigned to this morphologically similar. The occusal outlines are rounded-trape- genus. Twenty four relatively large teeth, with high crowned zoidal; the buccal half is broader than the lingual half. The area of 232 W. Wang / Journal of Human Evolution 57 (2009) 229–240

Figure 3. Mammal fauna from Chuifeng Cave. (1) Hylobates sp.; (2) Macaca sp.; (3) Hystrix kiangsenensis; (4) Hystrix magna; (5) Sus xiaozhu; (6) Sus peii; (7) Cervus sp.; (8) Cervus cf. fengqii; (9) Bovinae Gen. et sp. indet.; (10) Muntiacus sp.; (11) Megalovis guangxiensis;(12)Equus sp.; (13) Arctonyx collaris;(14)Pachycrocuta licenti;(15)Ursus thibetanus;(16)Felis sp.; (17) and (18) Ailuropoda microta;(19)Tapirus sanyuanensis; (20) Rhinoceros unicornis;(21)Stegodon preorientalis; (22) Sinomastodon sp. the paracone is markedly greater than that of the protocone. The protocone. All of the P4s are morphologically similar. The occusal median longitudinal groove is sinuous, separating the labial and outline is rounded to rectangular. The area of the paracone is lingual cusp at the median basin, and terminating at the mesial and greater than the area of the protocone. The anterior fovea is similar distal marginal ridges. The lingual slope of the paracone is much to the posterior fovea, which is clearly represented in the slightly more receding than the buccal slope of the protocone. The mesial worn Z008595. and distal marginal ridges are well developed. The anterior fovea is deeper than the posterior fovea, which is clearly represented in the Upper first/second molars slightly worn specimen of Z008588. In lingual view, the summit of the protocone is mesially placed, while the summit of the paracone The upper first/second molars are represented by 11 teeth, occupies a more distal position. Enamel folds are present on the consisting of five left M1–2s (of Z008598 and 8602) and six right lingual and buccal faces. In mesial or distal view, the strong M1–2s (Z008603-8; Fig. 6-[1–11]). The crowns are in good condi- development of the buccal basal prominence is remarkable, tion, but the roots are mostly missing, except in Z008600, which producing a characteristic lingual inclination of the labial face. has only the apical portion missing. Four teeth (Z008598, 8605, 7 and 8) are unworn or slightly worn, six teeth (Z008599, 8600, 2, 3, 5 Upper fourth premolars and 6) are moderately worn to expose one or two dentine craters and one tooth (Z008601) is heavily worn, with the lingual half The upper fourth premolars are represented by seven teeth, exposing a large sunken area. The occusal outline of the crown is including three left P4s (Z008592, 3, 5) and four right P4s (Z008591, somewhat rhomboidal in shape and generally buccolingually 4, 6, 7; Fig. 5-[14–20]). The crowns are well-preserved and the roots elongated. The protocone, hypocone, paracone, and metacone are are mostly or entirely missing. In occlusal view, the teeth of well developed. The protocone and paracone are slightly higher Z008591, 3, 5, 6 are slightly worn; Z008597 is moderately worn to than the distal cusps. The paracone and metacone rise steeply, expose two dentine areas on the apices of the paracone and while the protocone and hypocone are inclined buccally. The two W. Wang / Journal of Human Evolution 57 (2009) 229–240 233

worn with no dentine exposed (Fig. 6-[12–13]). The roots are entirely missing in the former and partially preserved in the latter. The occlusal outline is rounded-trapeziform and is much smaller than in the M1–2. Four cusps are present; the two mesial cusps are much larger than their distal counterparts. The paracone projects vertically, markedly protruding buccally. The protocone is the largest cusp, followed by the paracone and metacone, and the hypocone is the smallest cusp. The metacone and the hypocone are greatly reduced, forming an asymmetrical shaped crown mesio- distally. Thus, only a small portion of the distal moiety remains. The slopes of the cusps have ridges generally similar to those of M1–2. In mesial and distal views of Z008611, the highest cusp is the paracone, and the hypocone is the lowest. The mesial face is broad and moderately convex, marked with an interproximal wear facet. The distal face is small and strongly convex. Viewed lingually, the lingual slope recedes buccally. The root of Z008609 is divided into three branches, two buccal and one lingual. The lingual branch is buccolingually compressed, the mesiobuccal branch is mesiodistally compressed, and the dis- tobuccal branch is missing.

Lower deciduous first molar

The lower deciduous first molar is represented by a single tooth, Figure 4. Proportions of mammalian taxa from Chuifeng Cave. Z008540 (Fig. 5-[1]). This tooth is distinctly smaller than its cor- responding permanent tooth. The crown of the DM1 is sub- rectangular in occlusal outline, with a length-breadth index of lingual cusps, as well as the two buccal cusps, are clearly separated 105.5. Its occlusal surface is moderately worn. The metaconid is the by a vertical furrow, which extends to the cervix of the crown on the largest cusp. The protoconid is similar to the hypoconid in size, but buccal and lingual surfaces, respectively, being connected by larger than the entoconid, which has a cavity at its distal face. The a median longitudinal groove in the central basin. The protocone buccal face sways markedly towards the lingual side. The meta- and paracone are larger than the hypocone and metacone in conid and entoconid are more elevated than the two buccal cusps; general, while the hypocone is the smallest cusp. The four cusps are the hypoconulid is absent. The roots are missing. delimited by a Y-shaped fissure. In the unworn or slightly worn teeth, the mesial and distal Lower deciduous second molar marginal ridges are prominent, and the posterior fovea is deeper than the anterior fovea. On the slope of the paracone a large The second lower deciduous molar (Z008539) is rectangular in wrinkle extends from the apex of the cusp to the median longitu- occlusal view, with a length-breadth index of 139.5 (Fig. 5-[2]). The dinal groove, and is linked with the mesiobuccal marginal ridges, occlusal surface is lightly worn, with five cusps. The metaconid is forming a ‘‘flower-bud pattern.’’ On the slope of the metacone the largest and most elevated cusp, while the hypoconulid is the a main ridge extends from the apex of the cusp toward the Y-sha- smallest. The lingual side is straight, while the buccal side is ped fissures, giving out smaller secondary wrinkles. On either side lingually oblique for the protuberant base. Numerous crests origi- of the main ridge is the distinctly crenulated rim or marginal ridge, nate from the five main cusps and extend in various orientations, producing a ‘‘clover-leaf pattern.’’ On the slope of the protocone the forming a typical Gigantopithecus occlusal pattern. The mesial fovea main ridge is divided into two or three branches as it extends is transversely and moderately narrow, the distal fovea is much toward the longitudinal groove. On either side of these are marginal smaller and triangular. The roots are broken off at the base and are ridges. The wrinkle of the hypocone is similar to that of the pro- much more weakly developed than in the permanent tooth. tocone, but smaller in size. These characteristics are similar to those described for the Liucheng specimens (Woo, 1962). There is a slight Lower lateral incisor basal prominence on all sides of the crown. The prominence is more conspicuous on the lingual side. The lingual slope is much more The lower lateral incisor is represented by two right I receding than the buccal face. 2s (Z008533; Fig. 5-[3]) and a left I (Z008534; Fig. 5-[4]). Their In Z008600 the root consists of three separate branches, two 2 crowns are well preserved, but most of the roots are missing. The buccal and one lingual. The lingual branch is buccolingually lateral incisors are heavily worn and expose the dentine. In lingual compressed, extending lingually. The mesiobuccal and distobuccal view, the basal crowns are convex and inclined buccally. Viewed branches diverge from each other at the cervical enamel junction. buccally, the labial face is vertically convex and inclined lingually. The branches are buccolingually broad, mesiodistally compressed, generally vertical, and inclining lingually near the apex. Lower canine

Z008537 is the only lower canine, from the right side. The crown Upper third molars is well-preserved, but the root is missing (Fig. 5-[7]). This tooth is slightly worn. In mesial and distal view, the crown is wedge-like, The upper third molars comprise two teeth, one left M3 with a sharp cutting edge. The buccal surface is strongly convex. (Z008611) and one right M3 (Z008609). The crowns are well- The base of the crown is surrounded on all sides by a cingulum. preserved. Z008611 is slightly worn, while Z008609 is moderately Viewed lingually, the lingual tubercle extends to the tooth’s apex. 234 W. Wang / Journal of Human Evolution 57 (2009) 229–240

Figure 5. Incisors, canines, premolars, and deciduous teeth of G. blacki (occlusal view) from Chuifeng Cave. (1) LDM1, Z008540; (2) LDM2, Z008539; (3) RI2, Z008533 (lingual view); 1 1 3 3 3 (4) RI2, Z008534 (lingual view); (5) LC , Z008535 (lingual view); (6) LC , Z008536 (lingual view); (7) RC1, Z008537 (distal view); (8) LP , Z008585; (9) LP , Z008586; (10) LP , Z008587; (11) LP3, Z008588; (12) RP3, Z008589; (13) RP3, Z008590; (14) LP4, Z008592; (15) LP4, Z008593; (16) LP4, Z008595; (17) RP4, Z008591; (18) RP4, Z008594; (19) RP4, 4 Z008596; (20) RP , Z008597; (21) LP3, Z008545; (22) LP3, Z008546; (23) LP3, Z008547; (24) LP3, Z008548; (25) LP3, Z008549; (26) LP3, Z008550; (27) LP3, Z008551; (28) LP3, Z008552; (29) RP3, Z008543; (30) RP3, Z008544; (31) LP4, Z008553; (32) LP4, Z008554; (33) LP4, Z008555; (34) LP4, Z008556; (35) LP4, Z008557; (36) LP4, Z008558; (37) LP4, Z008559; (38) RP4, Z008560; (39) RP4, 8561; (40) RP4, 8562; (41) RP4, Z008563.

Lower third premolars than the latter. The posterior half of the crown is much wider than the anterior half; the two cusps are separated by the median The lower third premolars are composed of ten teeth, including longitudinal groove. The anterior and posterior foveae are deep, but eight left P3s (Z008545-52) and two right P3s (Z008543-4; Fig. 5- the anterior is a small pit, whereas the posterior fovea is a large [21–30]). The crowns of these teeth are well-preserved, but the transverse groove. The crown is surrounded by a basal swelling of roots of the Z008545 and 51 are partially missing, those of Z008543, cingulum. In lingual view, the basal part of the crown is slightly 7, 8, 9, and 52 are mostly missing, and those of Z008544, 6, and 50 convex and inclined towards the buccal. Viewed buccally, the labial are entirely missing. Slight occlusal wear occurs on six specimens of face is also slightly convex, but distinctly inclined lingually, with Z008544, 5, 7, 9, 50, and 51, moderate wear is present on Z008548 enamel elongation basally. and 52, and heavy wear occurs on Z008543 and 6, exposing The roots are mostly preserved in Z008545 and 51. In these a continuous dentine plane between the protoconid and specimens, the roots are divided into an anterior and a posterior metaconid. branch. The former is compressed buccolingually, with shallow The outline of the crown forms an irregular pyramid and is longitudinal grooves on the mesial and distal sides. The latter is asymmetrical. The main axis runs obliquely from mesio-buccal- flattened buccolingually, but its buccolingual dimension is much ward to disto-lingualward. The protoconid is well-developed and broader than the former, with a wider longitudinal groove devel- markedly larger than the metaconid. The protoconid and meta- oped mesially. In the broken apex of the roots of Z008545, one conid are conical, the former is higher and situated more mesially small rounded canal occurs in the anterior branch, and two small W. Wang / Journal of Human Evolution 57 (2009) 229–240 235

Figure 6. Molars of G. blacki (occlusal view) from Chuifeng Cave. (1) LM1–2, Z008598; (2) LM1–2, Z008599; (3) LM1–2, Z008600; (4) LM1–2, Z008601; (5) LM1–2, Z008602; (6) RM1–2, 1–2 1–2 1–2 1–2 1–2 3 3 Z008603; (7) RM , Z008604; (8) RM , Z008605; (9) RM , Z008606; (10) RM , Z008607; (11) RM , Z008608; (12) LM , Z008611; (13) RM , Z008609; (14) LM1–2, Z008564; (15) LM1–2, Z008565; (16) LM1–2, Z008566; (17) LM1–2, Z008567; (18) LM1–2, Z008568; (19) LM1–2, Z008569; (20) LM1–2, Z008570; (21) LM1–2, Z008571; (22) LM1–2, Z008572; (23) RM1–2, Z008573; (24) RM1–2, Z008574; (25) RM1–2, Z008575; (26) RM1–2, Z008576; (27) RM1–2, Z008577; (28) RM1–2, Z008578; (29) RM1–2, Z008579; (30) RM1–2, Z008580; (31) LM3, Z008581; (32) LM3, Z008584; (33) RM3, Z008582; (34) RM3, Z008583; (35) LM3, Z008610.

canals appear on the buccal and lingual sides of the posterior root. strong wear exposing large areas of dentine on the protoconid Ten P3s present sexual dimorphism; Z008541, 48, 49, 50, and 52 are occurs on six (Z008555, 56, 57, 59, 60, and 61). markedly large in dimensions, whereas Z008540, 44, 45, 47, and 48 The outline of the crown is nearly rounded quadrate, with its are relatively small, representing male and female, respectively. buccolingual dimension slightly greater than the mesiodistal length, except in Z008553. The protoconid and metaconid are cone- Lower fourth premolars shaped, located on the mesial two-thirds of the crown, and the remaining distal one-third of the crown is the talonid. The trigonid The lower fourth premolars consist of 11 teeth, including seven is higher than the talonid. The protoconid is slightly larger and left P4s (Z008553-9) and four right P4s (Z008560-3; Fig. 5-[31–41]). lower than the metoconid. These two cusps are separated by The crowns of these teeth are well-preserved, but roots of six teeth a median longitudinal groove, which is mesiodistally prolonged, (Z008554, 55, 58-61) are mostly missing, five teeth (Z008553, 56, intersecting the mesial and distal marginal ridge. The anterior and 57, 62, and 63) are partially missing the apical portion of the root. posterior foveae are transversely developed. However, the former is Slight occlusal wear is present on three specimens (Z008554, 58, smaller than the latter. On the whole, the P4 shows a high degree of and 62), moderately wear is present on two (Z008553 and 63), and molarization. In both lingual and buccal views the surfaces of the Table 2 Measurements of height and breadth (mesiodistal [MD] and buccolingual [BL]) of G. blacki teeth from Chuifeng Cave (in mm).a

Number Dentition MD BL Height Number Dentition MD BL Height

Z008533 RI2 7.50 9.83 – Z008551 LP3 14.39 17.57 12.30* Z008534 RI2 7.47 9.68 – Z008552 LP3 16.51 19.54 – 1 Z008535 LC 20.80 17.99 23.85* Z008553 LP4 17.41 15.26 14.17* 1 Z008536 LC 14.79 14.52 14.55* Z008554 LP4 13.77 14.68 14.26* 1 Z008537 RC 11.97 17.35 23.01 Z008555 LP4 15.17 17.35 – 3 Z008585 LP 13.13 16.54 – Z008556 LP4 13.76 16.60 – 3 Z008586 LP 13.84 18.49 – Z008557 LP4 15.01 18.95 – 3 Z008587 LP 13.55 18.77 – Z008558 LP4 15.75 16.77 15.71 3 Z008588 LP 15.46 20.60 17.33 Z008559 LP4 14.30 16.39 – 3 Z008589 RP 15.27 21.36 – Z008560 RP4 13.41 15.57 – 3 Z008590 RP 15.70 21.64 15.48* Z008561 RP4 16.44 19.85 – 4 Z008591 RP 13.05 19.75 13.22 Z008562 RP4 14.67 17.14 15.41* 4 Z008592 LP 12.59 19.86 – Z008563 RP4 17.36 18.20 15.16* 4 Z008593 LP 11.63 19.34 – Z008564 LM1–2 18.92 15.43 12.97 4 Z008594 RP 13.11 – – Z008565 LM1–2 19.44 17.32 – 4 Z008595 LP 14.27 21.50 16.25 Z008566 LM1–2 18.73 15.81 11.17* 4 Z008596 RP 13.70 21.93 16.11* Z008567 LM1–2 20.54 18.37 – 4 Z008597 RP 13.07 22.63 – Z008568 LM1–2 19.39 18.01 – 1–2 Z008598 LM 17.95 19.65 12.60 Z008569 LM1–2 18.25 17.03 – 1–2 Z008599 LM 17.12 20.20 – Z008570 LM1–2 19.24 15.87 – 1–2 Z008600 LM 15.95 20.86 – Z008571 LM1–2 17.83 14.69 – 1–2 Z008601 LM 17.66 22.59 – Z008572 LM1–2 18.59 16.30 – 1–2 Z008602 LM 16.35 20.78 Z008573 RM1–2 22.52 19.09 – 1–2 Z008603 RM 20.38 22.95 14.35* Z008574 RM1–2 20.91 17.83 12.38* 1–2 Z008604 RM 15.90 18.67 – Z008575 RM1–2 18.47 16.87 – 1–2 Z008605 RM 16.88 18.75 14.96 Z008576 RM1–2 17.25 14.04 – 1–2 Z008606 RM 16.81 20.30 – Z008577 RM1–2 18.81 16.66 – 1–2 Z008607 RM 17.98 17.00 – Z008578 RM1–2 17.38 15.79 – 1–2 Z008608 RM 21.08 23.97 19.35 Z008579 RM1–2 19.02 16.20 – 3 Z008609 RM 14.82 17.58 12.91* Z008580 RM1–2 22.84 20.00 14.45* 3 Z008611 LM 17.29 17.53 12.93 Z008581 LM3 21.65 19.24 12.18* Z008543 RP3 14.78 18.32 – Z008582 RM3 24.41 21.11 15.65 Z008544 RP3 13.87 17.13 13.42* Z008583 RM3 20.30 17.25 11.32* Z008545 LP3 14.90 17.43 13.60 Z008584 LM3 20.99 17.21 11.12* Z008546 LP3 18.45 20.33 – Z008610 LM3 18.19 16.48 – 2 Z008547 LP3 13.81 15.03 13.48* Z008538 LDM 12.45 14.24 – Z008548 LP3 15.73 16.71 – Z008539 LDM2 17.16 12.30 9.10 Z008549 LP3 17.89 20.40 16.18* Z008540 LDM1 14.43 13.68 – Z008550 LP3 16.92 21.43 15.45 a Values indicated by an asterisk indicate measurements from slightly worn crowns.

Table 3 Crown dimensions of the maxillary teeth of G. blacki (in mm).a

Liucheng Longgupo Mohui Jianshi Wuming Chuifeng

Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n C1 MD 19.05 (14.1–23.9) 17 15.00 (14.4–15.6) 2 15.3 1 17.80 (14.79–20.80) 2 BL 16.87 (13.0–20.8) 15 14.00 (13.5–14.5) 2 14.5 1 16.23 (14.52–17.99) 2 Area 325.49 (183.30–497.17) 14 210.3 (194.4–226.2) 2 221.9 1 244.47 (214.75–274.19) 2 L/B 112.98 (101.88–131.14) 14 107.15 (106.7–107.6) 2 105.5 1 108.74 (101.86–115.62) 2

P3 MD 14.65 (12.5–17.4) 39 15.6 1 14.4 (13.6–14.8) 3 15.5 1 16.4 (15.8–17.1) 4 14.49 (13.13–15.70) 6 BL 18.98 (16.3–22.6) 39 19.7 1 19.2 (18.2–20.6) 3 21.2 1 23.2 (22.1–24.0) 4 19.57 (16.54–21.64) 6 Area 279.47 (213.75–393.24) 39 307.32 1 276.87 (247.5–304.9) 3 328.6 1 380.51 (369.07–410.4) 4 285.3 (217.17–339.75) 6 L/B 77.13 (67.68–85.00) 39 79.19 1 75.1 (71.9–78.7) 3 73.1 1 70.75 (67.52–75.57) 4 74.25 (71.49–79.38) 6

P4 MD 14.29 (11.0–15.6) 63 14.15 (13.6–14.7) 2 15.4 1 17.5 1 13.06 (11.63–14.27) 7 BL 19.43 (17.0–24.0) 63 20.15 (20.1–20.2) 2 24.7 1 25 1 20.84 (19.34–22.63) 6 Area 259.47 (187.00–345.60) 63 285.15 (273.36–296.94) 2 380.4 1 437.5 1 271.64 (224.92–306.81) 6 L/B 68.45 (60.00–79.06) 63 70.22 (67.66–72.77) 2 62.3 1 70 1 62.70 (57.76–66.37) 6

M1–2 MD 17.81 (14.4–21.9) 105 16.5 1 18.5 (16.8–20.8) 3 21.45 (20.8–22.1) 2 17.64 (15.90–21.08) 11 BL 19.72 (16.5–24.0) 105 19.5 1 20.19 (18.2–22.5) 3 24.7 (24.2–25.2) 2 20.52 (17.00–23.97) 11 Area 353.07 (247.50–489.95) 105 321.8 1 374.78 (322.1–449.28) 3 530.14 (503.6–556.92) 2 363.93 (296.85–505.29) 11 L/B 90.43 (78.69–108.47) 105 84.6 1 91.78 (82.4–97.3) 3 86.83 (85.95–87.70) 2 86.38 (76.46–105.76) 11

M3 MD 17.12 (13.2–20.4) 88 18.4 1 19.1 1 16.06 (14.82–17.29) 2 BL 19.1 (15.2–24.0) 87 21.8 1 21.95 (21.2–22.7) 2 17.56 (17.53–17.58) 2 Area 329.97 (229.46–475.20) 87 401.1 1 433.6 1 254.82 (206.54–303.09) 2 L/B 89.83 (67.69–105.92) 87 84.4 1 84.1 1 91.47 (84.30–98.47) 2

a Individual tooth values from other sites are: P4 of Bijie: MD ¼ 12.8, BL=19.6, Area ¼ 250.9; M1-2 of Daxin: MD ¼ 17.7, BL ¼ 19.2, Area ¼ 229.84. W. Wang / Journal of Human Evolution 57 (2009) 229–240 237

Table 4 Crown dimensions of the mandibular teeth of Gigantopithecus (mm).a

Liucheng Longgupo Mohui Jianshi Wuming Chuifeng India

Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n Mean (Range) n Mean

I2 MD 7.86 (7.2–8.3) 10 7.6 1 7.49 (7.47–7.50) 2 BL 10.30 (9.6–11.3) 10 10.6 1 9.76 (9.68–9.83) 2 Area 81.06 (70.56–93.79) 10 80.6 1 73.02 (72.31–73.73) 2 L/B 76.42 (71.43–84.38) 10 71.7 1 76.74 (76.30–77.17) 2

C1 MD 12.97 (10.4–15.0) 15 12.6 1 13.3 1 10.93 (9.8–12.1) 4 11.97 1 9.3 BL 17.14 (14.5–21.0) 14 17.2 1 18.4 1 15.15 (12.8–17.5) 4 17.35 1 16 Area 221.90 (152.88–306.60) 12 216.72 1 244.7 1 167.63 (125.4–211.8) 4 207.68 1 148.8 L/B 72.98 (69.23–82.22) 12 73.26 1 72.3 1 72.45 (69.1–76.6) 4 68.99 1 58.13

P3 MD 15.83 (12.9–18.8) 67 17.57 (16.6–18.8) 3 15.2 1 17.27 (16.2–18.3) 3 16.5 1 15.73 (13.81–18.45) 10 11.8 BL 15.97 (12.9–21.0) 67 13.33 (12.4–14.3) 3 14 1 18.26 (16.2–19.5) 3 17 1 18.39 (14.22–21.43) 10 14.6 Area 254.21 (174.20–378.00) 67 234.92 (205.84– 3 212.8 1 315.2 (282.0–347.7) 3 280.5 1 286.36 (207.56– 10 172.28 268.84) 375.09) L/B 99.36 (85.71–111.64) 67 131.81 (130.08–133.87) 3 108.57 1 95.17 (83.1–106.1) 3 97.06 1 85.70 (78.96–94.14) 10 80.82

P4 MD 14.73 (12.1–17.3) 120 16.10 (14.3–17.5) 3 16.98 (16.0–18.6) 4 15.19 (13.41–17.41) 11 13.1 BL 15.79 (13.1–18.7) 119 15.67 (14.4–17.1) 3 18.7 (17.6–20.2) 5 16.98 (14.68–19.85) 11 14.5 Area 233.99 (170.30–317.90) 119 253.64 (205.92– 3 322.7 (280.0–366.4) 4 258.63 (202.14– 11 189.95 299.25) 326.33) L/B 93.40 (81.25–103.23) 119 102.70 (99.31–106.45) 3 89.58 (82.2–94.4) 4 89.86 (79.21–114.09) 11 90.35

M1–2 MD 19.13 (16.0–23.6) 41 19.48 (17.9–20.6) 4 19.12 (16.9–20.8) 4 21.18 (18.5–23.7) 4 21 1 19.30 (17.25–22.84) 17 18.05 BL 16.55 (13.1–20.5) 41 17.50 (16.2–18.8) 4 164.75 (14.8–17.6) 4 17.98 (15.1–20.2) 5 20 1 16.78 (14.04–20.00) 17 15.1 Area 319.14 (209.60–475.6) 41 341.96 (317.72–289.98) 4 316.38 (250.1– 4 374.09 (279.4–478.74) 4 420 1 325.98 (242.19– 17 272.56 352.0) 456.80) L/B 115.61 (105.64–127.74) 41 111.28 (109.58–113.81) 4 116.05 (112.6– 4 121.18 (117.33–124.2) 4 105 1 115.81 (107.16–122.86) 17 119.55 123.8)

M3 MD 19.19 (16.0–23.0) 51 19.2 1 22.47 (19.0–25.0) 3 23.00 (20.00–26.00) 2 21.11 (18.19–24.41) 5 19.6 BL 16.07 (13.7–19.0) 51 16.4 1 19.0 (16.0–21.2) 3 20.55 (18.1–23.0) 2 18.26 (16.48–21.11) 5 16.2 Area 310.43 (228.79–430.10) 51 314.9 1 431.7 (304.0–496.1) 3 480 (362–598) 2 379.61 (299.77– 5 317.52 515.30) L/B 119.55 (105.26–128.57) 51 117.1 1 118.48 (110.4–126.3) 3 111.77 (110.5–113.04) 2 115.64 (110.38–121.96) 5 120.99

a Dimensions of Individual teeth from other localities are: C1 of Tham Khuyen: MD ¼ 13.0, BL ¼ 18, Area ¼ 234; P3 of Daxin: MD ¼ 19.3, BL ¼ 17.6, Area ¼ 339.68; M3 of Daxin: MD ¼ 22.0, BL ¼ 18.3, Area ¼ 402.6; M3 of Bama: MD ¼ 25.2, BL ¼ 19.9, Area ¼ 501.5. crown are divided by vertical furrows into a larger mesial and Fig. 6 – [14–30]). The crowns of these teeth are all well-preserved. a smaller distal portion. The furrow on the buccal surface is longer The roots of five of these (Z008567, 70, 72, 75, and 79) are missing, and deeper than on the lingual surface. The roots of Z008563 are seven (Z008565, 68, 69, 71, 73, 74, and 80) are partially preserved, mostly-preserved and divided into an anterior and posterior and five (Z008564, 66, 76, 77, and 78) are mostly preserved. The branch. Both branches are flat and wide with the buccolingual diameter much larger than the mesiodistal diameter.

Lower first and second molars

The lower first/second molars consist of 17 teeth, including nine left M1–2s (Z008564-72) and eight right M1–2s (Z008573-80;

Figure 7. Comparisons of upper teeth of G. blacki. Figure 8. Comparisons of lower teeth of Gigantopithecus. 238 W. Wang / Journal of Human Evolution 57 (2009) 229–240

Table 5 Coefficient of variation for crown dimensions of teeth of G. blacki.

Liucheng Longgupo Mohui Jianshi Wuming Chuifeng

CV n CV n CV n CV n CV n CV n C1 MD 4.65 17 0.72 2 18.06 2 BL 11.76 15 0.50 2 6.02 2 P3 MD 1.55 39 0.48 3 0.37 4 1.23 6 BL 3.01 39 1.56 3 0.63 4 3.91 6 P4 MD 1.45 63 0.61 2 0.83 7 BL 3.57 63 0.01 2 1.85 6 M1–2 MD 2.91 105 1.91 3 0.85 2 2.86 11 BL 3.16 105 2.20 3 0.50 2 4.21 11 M3 MD 3.84 88 3.05 2 BL 3.66 87 0.00 2

I2 MD 0.16 10 0.00 2 BL 0.32 10 0.01 2

C1 MD 3.13 15 1.55 4 BL 5.25 14 5.28 4

P3 MD 1.86 67 1.26 3 1.10 3 2.74 10 BL 2.35 67 0.90 3 2.96 3 3.96 10

P4 MD 1.98 120 2.68 3 1.27 4 1.98 11 BL 2.14 119 1.84 3 1.48 5 2.45 11

M1–2 MD 4.07 41 1.82 4 3.02 4 4.99 4 2.51 17 BL 3.66 41 1.37 4 1.46 4 4.50 5 2.39 17

M3 MD 5.13 51 9.65 3 18.00 2 5.09 5 BL 3.31 51 7.24 3 12.01 2 3.59 5

occlusal wear on four teeth (Z008564, 66, 74, and 80) is slight sample shows that the metaconid is the highest cusp, followed by without dentine exposure, on ten teeth (Z008565, 67, 68, 69, 72, 73, the protoconid, hypoconid, entoconid, and hypoconulid. 75, 76, 77, and 79) is moderate, whereas, on three teeth (Z008570, The root of Z008576 is the best preserved of the M1–2s, having 71, and 78) it is heavy, with relatively large areas of dentine been only slightly gnawed by a porcupine at the apex. The roots exposed. divide into two branches, a mesial and a distal branch. Both The occlusal outline of these teeth is mesiodistally elongated, branches are mesiodistally compressed, declining distalward. and rounded oblong. The five cusps, arranged in a Y-pattern, are Measurements of the mesial branch are as follows: mesiodistal from largest to smallest: metaconid, protoconid, entoconid, hypo- diameter of 6.02 mm, buccolingual diameter of 13.11, and height of conid, and hypoconulid. These cusps are rounded and blunt, with 26.57 mm. The distal branch is 7.43 mm in mesiodistal diameter, a few narrow furrows. The buccal, lingual, and median longitudinal 13.30 mm in buccolingual diameter, and 26.15 mm in height. grooves, and the three foveae are visible, while the buccal and lingual grooves separate the protoconid and hypoconid, and met- Lower third molars aconid and entoconid, respectively. The buccal surfaces of the protoconid and hypoconid have a weak cingulum. The lightly worn The lower third molars include three left M3s (Z008610, 8581, and 8584) and two right M3s (Z008582 and 83; Fig. 6–[31–35]). The Table 6 crowns are in good condition. The roots of the four teeth (Z008581, Results of G. blacki metrical comparisons of teeth from Chuifeng and other localities.a 8583, 8584, and 8610) are partially preserved, but the roots of Z008582 are missing. The occlusal wear of Z008582 is minimal, that Liucheng Longgupo Mohui Wuming on three others (Z008581, 83, and 84) is slight to moderate, and that pp ppon Z008610 is extremely heavy, exposing a large dentine concavity. 1 C MD 0.785 – 0.458 – The occlusal outline of these teeth is mesiodistally elongated. BL 0.272 – 0.338 – The trigonid is broader than the talonid as the posterior part of the P3 MD 0.708 0.398 0.901 0.015 BL 0.592 0.953 0.782 0.009 crown is buccolingually compressed. Five cusps are separated by P4 MD 0.555 0.181 – 0.006 a Y-shaped system of fissures. The metaconid is the largest cusp, BL 0.155 0.524 – 0.036 followed by the protoconid, hypoconid, entoconid, and hypo- 1–2 M MD 0.009 – 0.533 0.012 conulid. There is a very small accessory internal tubercle between BL 0.294 – 0.645 0.019 M3 MD 0.168 – 0.471 – the metaconid and entoconid of Z008584 and Z008582, and a small BL 0.235 – 0.007 – tubercle exists also between the entoconid and the hypoconulid on I2 MD 0.235 – 0.142 – Z008581, 82, and 84. Three foveae are visible. Observation from the BL 0.217 – 0.097 – minimally worn tooth of Z008582 shows that the metaconid is the C MD 0.571 – – – 1 highest cusp, followed by the protoconid, hypoconid, hypoconulid, BL 0.990 – – – and entoconid. P3 MD 0.894 0.103 0.769 0.666 BL 3.310 0.002 0.065 0.523 In mesial view, the mesial face is vertically flattened and buc- P4 MD 0.555 0.352 – – colingually elongated. The interproximal wear facet is well marked. BL 0.053 0.214 – – Viewed distally, the distal face is remarkably convex and bucco- M1–2 MD 0.290 0.823 0.649 0.313 BL 0.765 0.398 0.993 0.060 lingually compressed. The lingual and buccal surfaces are convex.

M3 MD 0.237 – – 0.452 The root of Z008610 divides into two branches. The mesial branch is BL 0.035 – – 0.287 vertically flat and mesiodistally compressed, with two independent a Bold values indicate that G. blacki in Chuifeng Cave are significantly different radicular canals at the open root apex. The distal branch is mostly from those of the comparative samples (p < 0.025, after Bonferroni correction). missing due to porcupine gnawing. W. Wang / Journal of Human Evolution 57 (2009) 229–240 239

Measurement and comparison Longgupo is also close to the latter three samples, but the Jianshi specimens are smaller and are even smaller than those in Measurements are limited to the 77 G. blacki teeth with well- G. giganteus. The posterior lower teeth from Chuifeng, Liucheng, preserved crowns. Their mesiodistal and buccolingual dimensions Longgupo, and Mohui do not vary much in size. However, they have been taken. Crown height has been measured on 14 unworn are much smaller than the corresponding teeth from Jianshi, and 20 slightly worn teeth for comparison. All of the measurements Wuming, Bama, and Daxin, but larger than those of G. giganteus. are listed in Table 2. Thus, comparisons of the lower posterior teeth of Gigantopithecus In general, the morphology of the Chuifeng G. blacki teeth confirm a similar trend in size variation as that seen in the upper resembles that of the previously known samples of this species. The teeth. main similarities include: the posterior teeth are markedly larger than the anterior teeth; the incisors are small and peg-like; the Statistical analyses canines are reduced in size and low-crowned; the unworn posterior teeth are polycuspidate; the premolars are broad, flat, and molar- The results of the CV analyses are present in Table 5. The upper ized; and the molars are low-crowned and flat, with very thick canines from Liucheng and Chuifeng are notably variable, although enamel caps. those from Mohui are not, probably due to the small sample size. Metrically, the crown areas of the Chuifeng incisors are smaller 1 The lower canines of Liucheng and Jianshi are generally variable. than that those from Liucheng and Mohui. The crown size of C is The lower third molars from Liucheng, Jianshi, Wuming, and close to the Mohui and Jianshi samples, but smaller than the mean Chuifeng appear to be more variable in length and breadth than of the Liucheng sample. The crown area of C1 is distinctly larger that of other postcanines. than G. giganteus, but smaller than that of the Longgupo, Mohui, The results of T-test analysis (Table 6) show that the dimensions Tham Khuyen, and the mean of the Liucheng samples. Its breadth- of canines from Liucheng and Mohui are not significantly different length ratio is less than any G. blacki previously known, being much from those from Chuifeng. The length and breadth dimensions of P3 more mesiodistally compressed, while the G. giganteus is even less 4 3 and length of P from Wuming are larger than those of the corre- compressed. The size of the P crowns are close to those from sponding teeth from Chuifeng. M1–2 length in the Liucheng sample Liucheng, Longgupo, Mohui, and Jianshi, but distinctly smaller than 1–2 4 and the length and breadth of M from Wuming are larger than those from Wuming. The P crowns match those from Liiucheng, those from Chuifeng. The breadth of M3 from Chuifeng is smaller Bijie, and Longgupo, but are somewhat smaller than those from than that from Mohui. The breadth of P3 from Longgupo is smaller Jiangshi and much smaller than their Wuming counterpart. As in than that of Chuifeng. the Liucheng sample, the crowns of P3 from Chuifeng exhibit great range of variation in size. The mean of the crown area is larger than all of the other Gigantopithecus samples, except that from Jianshi. Conclusions The crown area of P4 is close to that from Liucheng and Longgupo, but smaller than that from Jiangshi and larger than G. giganteus. The Recently recovered in situ material, comprising 92 teeth, from sizes of M1–2 are close to those from Liucheng, Mohui, and Jianshi. the Chuifeng Cave of Bubing Basin, west Guangxi, south China can The mesiodistal and buccolingual diameters fall well within the definitively be assigned to Gigantopithecus blacki. These teeth, range of the sample from Liucheng, but close to the lower limit of associated with a typical early Pleistocene mammalian fauna of the Wuming specimens. The Chuifeng M1–2 are quite similar to south China, are morphologically similar to known G. blacki Liucheng and Mohui in mean values and ranges, but smaller than samples. The crowns of 78 teeth of Gigantopithecus are well- those in the Jianshi and Wuming samples. The crown of M3 from preserved; the roots have mostly been lost as a result of porcupine Chuifeng is relatively smaller than those from Liucheng, Mohui, and gnawing. The size of these teeth is similar to that of early Pleisto- Jianshi, and its mean falls within the lower limit of the range of cene G. blacki. However, the posterior teeth are distinctly smaller Liucheng. The M3 is clearly smaller than that of Jianshi, Wuming, than those of G. blacki in later time periods, and larger than those of Bama, and Daxin, but larger than that from Liucheng and Mohui the earlier species, G. giganteus. This potentially implies temporal (Tables 3 and 4). variation in the evolution of the lineage. The difference in size of the upper teeth of G. blacki from With the exception of Liucheng, the specimens from Chuifeng Chuifeng, Liucheng, Mohui, Jianshi, and Wuming are presented in Cave represent the largest sample, far exceeding the specimen Figure 7. Comparisons of the lower teeth of G. blacki from Chuifeng, counts from other reported localities. Liucheng, Longgupo, Mohui, Jianshi, and Wuming and G. giganteus Gigantopithecus plays a dominant role in the Chuifeng fauna, of India are shown in Figure 8. representing 9% of the total collection of mammal teeth (n ¼ 1007). In these comparisons, the C1 from Chuifeng is smaller than This frequency possibly shows that the habitats in East Asia were that from Liucheng, but slightly larger than the means of the highly suitable for this primate during the early Pleistocene. Jianshi and Mohui specimens. The P3 and P4 from Chuifeng are Eleven teeth (12% of the total) of Gigantopithecus show signs of very close to those from Liucheng, but markedly smaller than caries. Extremely heavy wear on M3 from this locality indicates that those from Jianshi, and much smaller than their Wuming coun- the extinct ape was adapted for eating tough, fibrous food, or terparts. Comparisons of M1–2 show that Chuifeng and Liucheng, possibly had a relatively long life span. The upper canines and lower and Mohui and Jianshi, are relatively close in crown size third premolar show sexual dimorphism. These new discoveries compared to the largest specimen from Wuming. Regarding the from Chuifeng Cave provide additional insight into the evolution M3, the size trend is clear: the Jianshi sample is the largest, fol- and paleobiology of the largest primate. lowed by Mohui, Liucheng, and Chuifeng. Overall, the anterior teeth of G. blacki from all localities do not vary in size, whereas Acknowledgement the posterior teeth show a distinct trend to increase in size from Chuifeng, Liucheng, Mohui to Jianshi and Wuming. This trend First, I would like to thank F. Tian from Tiandong Museum of potentially corresponds to temporal variation. Guangxi, S.W. Xie, and S.F. Qin from the Natural History Museum of In the lower dentition, the I2 of Chuifeng, Liucheng, and Guangxi Zhuang Autonomous Region for their participation in the Mohui are very close in the size of their crowns. The C1 of excavation. Also, special thanks to Terry Harrison, Susan Anto´ n, Eric 240 W. Wang / Journal of Human Evolution 57 (2009) 229–240

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