example, Gigantopithecus). The dietary habits of extinct and extant species provide Gigantopithecus: A Reappraisal of additional insights into the socioecological Dietary Habits of the species. Differing dietary adaptations are paItially responsible for many of the behavioural and ecological differences that separate extant taxa, and by extension, must obviously affect the differentiation of fossil taxa. Aspects of diet, Introduction living or fossil, can be used to infer metabolic One of the many problems central to rate, body mass, and ranging paleoplimatology and paleoanthropology patterns, among other things. This paper, adopts concerns the reconstruction of dietary a broad comparative approach in order to more behaviours and adaptations in fossil species. accurately reconstruct the diet of one of Such endeavours become particularly difficult paleoplimatology's greatest enigmas, the fossil when the fossil species has left no living Miocene , Gigantopithecus. descendents. It thus becomes vital to identify plausible living analogs for the purpose of History of Discovery & Taxonomic inferring possible behaviours in the fossil Considerations species. Unfortunately, there are times when a For thousands of years, Chinese chemists proper living analog does not exist (for have been using "dragon's teeth" as medicinal

T()TI':~I yol II :21111:2-:21111.; Coprrighl II) :211111'1'( l'1V.\!: The U\X'() Journal of .\nlhropologl' ingredients. In 1935 a Dutch paleontologist, sizes and shapes of jaws and teeth, and the G.H.R. von Koenigswald, discovered these thickness and stlUcture of " (Ungar massive molars within Chinese pharmacies. 2002:261). Second, material that pel1ains to the Following the discovery, von Koenigswald actual foods consumed by the individual identified the molars as belonging to a new, represents nonadapti ve evidence. This would giant ape species, which he called include studies such as dental microwear, stable Gigantopithecus blacki (von Koenigswald isotopes, and trace element analyses. 1952). Since its original discovery, additional One of the first attempts to reconstmct Gigantopithecus fragments have been the diet of Gigantopithecus occurred following discovered, both in the field and in Chinese the discovery of a giant ape's jaw bone in a dmgstores. To date, only teeth and mandibular cliffside cave, south of the River in pieces have been recognized among these China. Within this cave, the skeletal remains of fragments. deer, boar, , and were Despite the ever increasing sample of found in association with the Gigantopithecus Gigantopithecus remains, its taxonomic position remains (Wen-Chung 1957). These associated remains elusive. Numerous taxonomic and remains were all hoofed , and thus, could evolutionary statements have been forwarded not have possibly climbed the 270-foot, near concerning the Gigantopithecus. Among verticaL cliff to access the cave' s mouth (Wen- these, two major positions have been posited Chung 1957). The only alternative, then, was that regarding the affinities of Gigantopithecus: i) as the giant ape canied these hoofed animals into the a unique pongid, and ii) as either an extinct side cave for food. Based on this evidence, and the branch, or ancestral stock, of later Asian surface of the newly discovered teeth,Wen- hominids. Those favouring the former position Chung (1957:836) concluded that. "it was include Ti-Cheng (1962), Simons and Pilbeam obvious that the had a mixed diet of meat (1965, 1972, 1978), Simons and Chopra (l969a, and vegetables, quite different from that of 1969b), Simons and Ettel (1970), Pilbeam modem which live on fmit." This early (1970, 1972), Simons (1972, 1978), Cormccini reconstruction has been turned upside-down in (1975), and Delson and Andrews (1975). The light of relatively recent paleoenvironmental latter position is held by Weidenreich (1945, evidence which indicates that, "what are now 1946), von Koenigswald (1952), Dart (1960), cliffside caves were sinkholes in a limestone Woo (1962), and Eckhardt (1972). Although plateau when the giant apes flourished" (Simons divided, the literature now seemingly points and Ette! 1970:83). towards a pongid relation (Klein 1999; Fleagle Thus, it seems apparent that the dietary 1999). A-note of caution is waITanted: if diet reconstmction of Gigantopithecus will not be as underlies the behavioural and ecological simplistic as previously thought. The traditional differences that separate extant taxa~ the reverse and most conservative view, based upon jaw and - that taxonomic differences may reflect teeth morphology, postulates that differing dietary adaptations - may also hold Gigantopithecus probably foraged on hard, tme.If this is the case, determining the correct fibrous matelial (Conroy 1990). This view, like taxonomic position of Gigantopithecus may be Wen-Chung's (1957) inference. may also be in required prior to any accurate dietary jeopardy based on the numerous findings elicited reconstmction. With this in mind, the from studies of adaptive and nonadaptive comparati ve approach taken here assesses the elements. The results of such studies shall be dietary skeletal evidence of Gigantopithecus discussed below. based on the underlying assumption that it does belong to a pongid (albeit a unique one), rather Dental MOIphology than a hominid, classification. Since it is the enormous size of the Giganropithecus teeth that have so captured the Dietary Considerations attention of the primatological world, it is not Identifying the anatomical evidence for surprising that most adaptive studies have focused diet among modern contributes on this particular aspect. It has long been immensely to our understanding of fossil assumed that relative tooth size reflects functional primate dietary habits. Following Ungar specialization (Ungar 2002). Additionally, (2002), two lines of evidence can be used to variations in relative tooth shape may reflect a infer the diets of extinct primates. First, means of adapting to changes in the internal adaptive evidence "concerns analyses of the

'1'(),I'I':~I \"01 1111l1J2-1111n Corl'right © 11111,)TU1V~1: The UWO .Iournnl of ,\nthmro\og" characteristics of foods, such as the strength and (Pilbeam 1970). Thus, a stout and broad toughness (Teaford and Ungar 2000). morphology appears to characterize Gigantopithecus teeth are colossal, Gigantopithecus canines. Since the lower canines exceeding all known primate teeth in their of Gigantopithecus are truncated more than dimensions. Even those of an adult male gOlilla sharpened, it is reasonable to infer that the are dwarfed by comparison. Data taken from maxillary canine was not large, and not similar in von Koenigswald (1952) indicate that the third function compared to that of the gOlilla (Frayer lower molars range from 22.3 to 23.1 nun in 1973). Frayer (1973:418) also reasoned that length. Such dimensions place volume "since the mandibular canine was truncated estimates at about six times larger than modern during life, masticatory actions peIformed at the humans, and twice as large as adult male canine appear to be more involved with grinding, , when compared to the corresponding than with sheming and gripping functions." Thus, teeth of such individuals (Weidenreich 1944). the canines seem to have been essentially Yon Koenigswald (1952:311) also rep0l1s that, grinding teeth additional to the and "in addition to the usual five main cusps, the molars. It should come as no surprise, then, that tooth pattern includes vil1ually all the secondary these morphological characteristics of the teeth of cusps that might possibly occur in the of Gigantopithecus have led many to assume and a higher primate." As in anthropoid apes, the predict a diet of hard, tough, and rigid food items second molar of the Gigantopithecus specimens that required heavy grinding and crushing. are larger than the first molar. Additionally, the molars are distinctly longer than broad (Strauss Enamel Thickness 1r. 1957). In Gigantopithecus bilaspurensis, Attempts at dietary reconstructions have cheek tooth cusps show little relief and are also examined the implications of enamel flattened and plate-bke rather than conical; thickness. Two adaptive explanations for occlusal surfaces are relatively broad and flat possessing thick enamel have been offered: either (Pilbeam 1970). Gigamopithecus blacki, while to prolong the use-life of teeth in an abrasive diet; sharing similar characteristics, tends to have or to minimize the lisk of crown damage given higher crowned and more cuspidate molars high occlusal forces caused by a diet including (Pilbeam 1970). It would seem that the molars very hard objects (Ungar 2002). In recent years, of Gigantopithecus are extremely large, with it has become common to describe high, blunt cusps separated by deep, nan'ow Gigantopithecus molars as thick enameled fUlTOWS(von Koenigswald 1952). It is also (Fleagle 1999; Klein 1999). If such is true. and worth mentioning that the lower antelior the assumptions underlying the adaptive function is relatively broad (as in of thick enamel are true. possession of thick sapiens) rather than elongated (Fleagle 1999). enamel buttresses the data from dental In other words. the premolars have become morphology, which together suggests a hard, "molarized" - that is, broad and flattened. abrasive diet. possibly consisting of nuts, seeds, With such enormous molars. it would and subterranean tubers. come as no surprise for this gigantism trend to continue into other teeth. However, relative to Mandibular Morphology cheek tooth size, the of Mandibular fragments are among the Gigantopithecus are small and closely packed most common bony remains found on fossil between the canines (Pilbeam 1970). plimate sites. It is assumed that the architecture Furthermore, the incisors appear to be almost of this bone has been adapted to withstand the peg-bke, rather than chisel-like (Strauss. J r. stresses and strains associated with oral food 1957). processing. If this is the case. its morphology The canine morphology of probably reflects (at least indirectly) some aspects Gigantopithecus is also unique. Although the of diet. front premolar was clearly bicuspid. the trigonid The mandibular body of Gigantopithecus was still quite developed, suggesting that the is extremely deep and highly robust. PoweIful upper canine continued to shear against the mastication is indicated by numerous features. antelior face of the lower premolars (Pilbeam The symphysis was long. deep and poweIfully 1970). However, recovered canines show little buttressed by large tOli (Yinyun 1982). That the projection beyond the plane of the cheek teeth. face was short is indicated by the M, origination In both sexes, it seems, the canines wore do\\'n position of the ascending ramus (Pilbeam 1970). rapidly at the tips, even at an early dental age Powerful mastication is also indicated by the

.!( f11·\1 ",III ~II[)~ ~1I11) Cllpl'fightr£. 21))))T( lTI \1. 1·11, ,.\\( l.l',urIl.d "t.\nthropolog\· everted and buttressed mandibular gonial angles early specimens were poor representatives; more (Pilbeam 1970). Short, deep faces are adapted recent finds provide a better basis for micro wear to powerful mastication (Klein 1999). analysis. Dental orientation within the jaw is Daegling and Grine (1987) examined also of interest. The "Theropithecus complex" occlusal microwear in a sample of describes a unique orientation whereby grinding Gigantopithecus blacki teeth, and concluded that teeth dominate - and may include adapted this species was unlikely to have been a hard- premolars and canines (Pilbeam 1970). Here, object specialist. In a recent study by Daegling large grinding cheek teeth and powerful and Grine (1994), occlusal microwear on the teeth masticatory muscles are vital, while incisors are of Gigantopithecus were compared with relatively unimportant. Gelada baboons microwear on the molars of two extant (Theropithecus), which subsist on small seeds specialists. Hapalemur griseus (the gentle lemur) and nuts - thus, grinding - represent a living and Ailuropoda melanoleuca (the ). member of the "Theropithecus complex." Bamboo feeding within the extant species does Surprisingly, if we review the aforementioned not produce a consistent pattern of microwear. data on Gigamopithecus , we see that it Thus, because of these dissimilar patterns, dental shares many characteristics that belong to the microwear on Gigamopithecus provides little "Theropithecus complex." Such dental direct evidence for bamboo feeding. It does, orientation within the of however. appear unlikely that Gigantopitheclls Gigamopithecus may indicate a similar diet to subsisted exclusively on this particular resource the gelada baboon - nuts and seeds. A study by (Daegling and Grine 1994). Rather, the Groves (1970) also shows "Theropithecus micro wear patterns found on the molars of complex" -like characteristics in the modern Gigamopithecus most closely resemble that of the mountain , whose differences with the predominantly frugivorous troglodytes. the eastern and western lowland gorilla mimic those common chimpanzee (Daegling and Grine 1994). differences found between Gigantopithecus and This would suggest a diet of a broad range of its dryopithecine forbearers. Groves (1970) fruits and fibrous materials. concludes that these similarities may provide a An alternative technique, based on the basis for inferring the Gigamopithecus diet identification of opal found bonded to from the mountain gOlilla diet. which includes the enamel surfaces of the teeth of fossil species, roots, bark, and similar hard-wearing, bulky allows for the identification of individual plant items. remains eaten prior to death (Ciochon et al. 1990b). The phytoliths found on Gigantopithecus Dental Microwear molars derive from two distinct taxonomic groups Associations between aspects of diet, and from different plant organs: i) the vegetative tooth use, and microwear in living primates parts of grasses, and ii) the fruits and seeds of have been used to infer diets of fossil primates. dicotyledons, specifically of a species in the Primates that often use their antelior teeth , or a closely related, family (Ciochon et during ingestion have high densities and al. 1990b). The relative roles of grasses and fruits frequencies of microwear striations on their in the diet of Giganropithecus are difficult to incisors (Teaford and Ungar 2002). High estimate. However. judging from the present incidences of long narrow scratches on molars frequency of dental phytoliths in Gigantopithecus, point towards . where as frugivores fruits may have constituted a significant portion tend to have more pits than scratches (Teaford of the diet (Ciochon et al. 1990b). and Ungar 2002). Yon Koenigswald's (1952:317) Locomotor Limitations original assessment of the gross dental wear on While these dental aspects provide Gigamopithecus molars, "suggests the kind of invaluable clues to the diet of Giganropithecus, attrition found in man." It was later determined one must not forget the limitations imposed by by Strauss (1957 :685) that, "the occlusal body size. Body size and stature estimates for surfaces of the teeth appear to have been worn Gigantopithecus have ranged greatly: 10 feet tall down considerably, so that the crown patterns and 1200 pounds (Ciochon et al. 1990a), 9 feet of the molars cannot be made out."Such a tall and 600 pounds (Simons and Ettel 1970), a statement lends critical support to the idea of height of twelve or more feet (Wen-Chung 1957), massive gtinding and crushing as the primary and more recently, possessing a body mass of 300 mode of mastication in Gigantopithecus. These kg (Fleagle 1999). Given its size,

TOTI':~! ml II 2(J(12-2(J(J~ Cop\'right © 21111.) TOTh\!: The UWO .Journal of .\nthropolog\· Gigantopithecus must have been completely accurate body stature estimates, current telTestlial. Recently, the long bones of reconstructions must be taken with a grain of salt. Gigantopithecus have been determined to be It is unlikely, however, that a post-cranial 20-25% longer and more robust, on average, skeleton indicating arboreality would be than those of living gOlillas (Johnson, Jr. 1979). associated with a dental and mandibular Even this increase in size would most likely morphology of such extreme size. Thus, it resuict Gigantopithecus to a ground-dwelling appears that a consensus has been reached locomotor repertoire. This restriction has regarding the habitually ten-estlial nature of numerous implications on the of foodstuffs Gigantopithecus. Assuming this, how then would available to Gigantopithecus, and will be fruit become a vital pm1 of the diet, as was discussed in the next section. posited by the analysis of Ciochon et al. (l990b)? First, perhaps some species of the Discussion Moraceae family represent low-lying fruits. The preceding sections discussed the Second, overly lipe fruit may fall to the ground, various lines of evidence that appear in the upon which Gigantopithecus could subsist. fossil record which indicate, either directly or However, the chances of enough fruit, left to indirectly, the diet of Gigantopithecus. An over-lipen by other frugivores, to sustain often overlooked vmiable in discussions of Gigantopithecus is slim. Similm-ly. it is also fossil diets is the influence of the environmental unlikely that Gigantopithecus could have context in determining resource availability. subsisted on fruit dropped by clumsy arboreal Knowledge of the paleoenvironment dUling the frugivores. Lastly, perhaps the indication of existence of Gigantopithecus provides heavy frugivory suggested by phytolith additional clues to its diet. Early frequencies is elToneous. For instance, only four paleoenvironment reconstructions, based mainly teeth, among thousands now discovered, were on associated fauna, indicated a grassland and selected for inclusion in the phytolith study. Such open woodland terrain (Pilbeam 1970; White selecti ve sampling introduces a large amount of 1975). More recent reconstructions (Ciochon et bias. Schwartz (1991) concurs, and notes that a al. 1990a; Jablonski et at. 2000) suggest great deal is being drawn from the analysis of subtropical-seasonal to tropical forest settings. four teeth, only two of which were found to be Like the range of dietary reconstructions harboming phytoliths, with the greatest discussed earlier, paleoenvironmental settings concentration on only one.Clem'ly, larger also vary, depending on the source. This shift samples of teeth need to be similarly analyzed. in reconstruction to a tropical forest setting is Fm1hermore, phytolith accumulation properties undoubtedly related to recent micro wear and need additional investigation. Perhaps the phytolith analyses. phytoliths associated with various species Locomotion and resource acquisition accumulate at differing rates and concentrations; are intimately linked, and are the direct or perhaps, the phytoliths only represent the last, consequences of environmental setting. With or most recent, foods consumed by the individual such enormous mass. the only plausible prior to death, and not necessarily the most locomotor pattem for Gigantopithecus would commonly eaten item. involve some form of tenestrial, quadrupedal The importance of Ciochon et al.·s movement. This great size would, no doubt, (I990b ) phytolith study must not be entirely eliminate any form of arboreality. Thus, dismissed. As argued by Daegling and Grine foodstuffs in the diet of Gigantopithecus would (1994), bamboo feeding may be a possible dietary be limited to low-lying (or often-falling) items. adaptation for Gigantopithecus. Supporting As a general rule, fruits require high light to evidence for such a scenario may be found in grow. and thus, in tropical forests most fruits Ciochon et al.' s (1990b) analysis. In addition to are found on terminal branches high-up in the phytoliths belonging to the Moraceae family, canopy ceiling (Campbell 1996). Such phytoliths of vegetative grasses were also positioning would obviously be problematic for recovered. The recovery of such phytoliths may a ground-dwelling species such as be of some importance in that the bamboo family Gigantopithecus. is a vegetative grass. As stated earlier, additional A number of explanations are possible. phytolith analysis need to be conducted. Perhaps body size reconstructions have been A cautionary note is also wan-anted for grossly over-estimated. Until post-cranial the dental micro wear analyses of diet of remains are recovered which permit more Gigantopithecus. Microwear features do not

'1'0'1'1 ':1\1 \"1)1 II 211112-21111.1 CopHight © 21111.'1'( )'1'1 ':1\1: The UWO -'ollmnl of. \nthropology necessarily reflect specific food items per se, the past, and the skepticism necessary to but rather the mechanical properties of those understand the boundary between supported foodstuffs, or the constituents of those hypothesis and unsuppOlted speculation (Plavcan foodstuffs. Thus, the varying degrees of ef al. 2002). Reconstructions of the behaviour of mechanical properties of food items will fossil species are limited to the evidence available produce varying degrees of micro wear on the in the fossil record. This includes adaptive dentition of a specimen. This may create (dental morphology, mandibular form, and somewhat of a "dental paradox", in that the enamel structure) and nonadaptive (dental microwear found on dentition may not reflect microwear, gross tooth wear, stable isotope, and the most common, or distinctive, dietary trace element analyses) lines of evidence. foodstuff of that species, but merely the most Though the cun-ent nonadaptive abrasive item within the dietary range of that evidence regarding the diet of Giga11fopithecus is individual (Plavcan et al. 2002). somewhat controversial, the adaptive evidence As discussed earlier, one of the most does provide some important clues. The frequently cited enamel con-elations is between are deep - top to bottom - and are the consumption of hard, and abrasive, food extremely thick. The molars are low-crowned items, and thick molar enamel. However, thick and flat, with very thick enamel caps. The canine enamel by itself does not necessarily provide teeth are not sharp and pointed - as is the case in protection against hard objects. which other apes - but are more similar to premolars in commonly cause fractures in the enamel layer morphology, while the incisors are small. peg- (Ungar 2002).To prevent fractming, the best like, and closely packed. The features of the protection is "prism or crystallite decussation or teeth, combined with the massive. robust jaws, interweaving" within the enamel structure lead to the conclusion that Giganfopithecus was (Teaford and Ungar 2000:13508). Thus, it may adapted to the consumption of tough, fibrous be the structure, and not the thickness of foods through the extensive use of crushing and enamel. that provides clues to the abrasive and grinding. However. as discussed previously, this rigid nature of dietary foodstuffs. If this is the conclusion should be met with the same amount case, the thick enamel layers on of skepticism as that given to other dietary Gigantopithecus teeth may not provide evidence reconstructions, namely those of Ciochon et af. for a diet that consists of hard, tough objects. (1990b) and Daegling and Gline (1994). To date, analyses of Gigantopithecus enamel Taken cumulatively. primary foodstuffs structure have not been conducted. and thus. relating to the reconstructed diet of statements and inferences about such properties Giga11fopifhecus have included nuts. seeds. cannot be made. tubers. and other hard objects, grasses. bamboo, It should also be noted that the and fruit. Various skeletal elements point towards enormous molar size found in Gigantopifhecus differing sets of food items. It can only be may fall victim to the aforementioned "dental concluded that the most distinctive characteristic paradox." During its existence. the of the diet of Gigalltopifhccus is the enormous environmental setting of GigallfOpifhccllS range and variability of that diet. Since habitats experienced increasing aridity and Gigantopithecus represents an ape unlike any seasonality (Jablonski ef al. 2(00). These other in primate history. it should come as no climatic fluctuations may have created surprise that its diet should reflect its enigmatic shortages of prefen-ed food items. The large persona. As such. it probably exploited numerous molars and increased occlusal surface may have resources, including all those mentioned here. In evolved as a coping strategy during periods of terms of dietary foodstuffs. the range of resources stress in which dietary habits wen: forced to exploited by Gig({lIfOpifhcc/ls most closely switch to tough, fibrous, and hard food items. resembles that of anatomically modem Homo In this way, the enormous gross morphology of sapiens (i.e. an opportunistic omnivore). This Gigantopithecus dentition is an adaptation for extensive range in diet. along with a gigantic critical feeding, and not a reflection of the most stature, are just two of the many characteristics common dietary foodstuffs. that make GigQlIfollifhcc/ls an enigmatic ape.

Conclusions Reconstructing behaviour in any fossil species requires an equiliblium hetween the enthusiastic inference of how organisms )jyed in

I (111\1,,,1 II 211112-21111.' Coprnglll' 2",,;'1 ( ) II \1·'1 h, LI\\,( Jjournal "i .\nthropojogr Frayer, D.W. 1973. Gigantopithecus and its Campbell, N.A, ed. 1996. Biology. New Relationship to . York: The Benjamin/Cummings Amelican Joumal of Physical Publishing Company, Inc. Anthropology 39(3):413- 426.

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Conroy, G.C 1990. Primate Evolution. New Klein. RG. 1999. The Human Career: York: W.W. Norton and Co. Human Biological and Cultural Origins. Chicago and London: The Corruccini, RS. 1975. Gigantopithecus and University of Chicago Press. hominids. Anthrop. Anz. 35:55-57. Pilbeam, D.R 1970. Gigantopithecus and the Daegling, D,1., and FE. Grine. 1987. Tooth Oligins of . Nature wear, gnathodental scaling and djet in 225:516-518. Gigantopithecus blacki. Amelican Joumal of Physical Anthropology 1972. The Ascent of Man. New York: 72:191-192. MacMillian.

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TUIVI\! YO] II ~IIIJ~-~IIII.) Copyright © ~1I11.)'1'()'IVI\[: Thl' U\X'() .Iournal o! .\nthropo!o!-,,,' Simons, E.L., and S.R.K. Chopra. 1969a. Simons, E.L., and P.c. Ettel. 1970. Gigantopithecus (Pongidae, Gigantopithecus. Scientific Hominoidea), a new species from American 222:77-85. North India. Postilla 138:1-18.

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