Gigantopithecus: A Reappraisal ofDietary Habits

IntroductionOne of the many problems central to

paleoplimatology and paleoanthropologyconcerns the reconstruction of dietarybehaviours and adaptations in fossil species.Such endeavours become particularly difficultwhen the fossil species has left no livingdescendents. It thus becomes vital to identifyplausible living analogs for the purpose ofinferring possible behaviours in the fossilspecies. Unfortunately, there are times when aproper living analog does not exist (for

example, Gigantopithecus). The dietary habits ofextinct and extant primate species provideadditional insights into the socioecological natureof the species. Differing dietary adaptations arepaItially responsible for many of the behaviouraland ecological differences that separate extanttaxa, and by extension, must obviously affect thedifferentiation of fossil taxa. Aspects of diet,living or fossil, can be used to infer metabolicrate, body mass, ecological niche and rangingpatterns, among other things. This paper, adoptsa broad comparative approach in order to moreaccurately reconstruct the diet of one ofpaleoplimatology's greatest enigmas, the fossilMiocene ape, Gigantopithecus.

History of Discovery & TaxonomicConsiderations

For thousands of years, Chinese chemistshave been using "dragon's teeth" as medicinal

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ingredients. In 1935 a Dutch paleontologist,G.H.R. von Koenigswald, discovered thesemassive molars within Chinese pharmacies.Following the discovery, von Koenigswaldidentified the molars as belonging to a new,giant ape species, which he calledGigantopithecus blacki (von Koenigswald1952). Since its original discovery, additionalGigantopithecus fragments have beendiscovered, both in the field and in Chinesedmgstores. To date, only teeth and mandibularpieces have been recognized among thesefragments.

Despite the ever increasing sample ofGigantopithecus remains, its taxonomic positionremains elusive. Numerous taxonomic andevolutionary statements have been forwardedconcerning the genus Gigantopithecus. Amongthese, two major positions have been positedregarding the affinities of Gigantopithecus: i) asa unique pongid, and ii) as either an extinct sidebranch, or ancestral stock, of later Asianhominids. Those favouring the former positioninclude Ti-Cheng (1962), Simons and Pilbeam(1965, 1972, 1978), Simons and Chopra (l969a,1969b), Simons and Ettel (1970), Pilbeam(1970, 1972), Simons (1972, 1978), Cormccini(1975), and Delson and Andrews (1975). Thelatter position is held by Weidenreich (1945,1946), von Koenigswald (1952), Dart (1960),Woo (1962), and Eckhardt (1972). Althoughdivided, the literature now seemingly pointstowards a pongid relation (Klein 1999; Fleagle1999). A-note of caution is waITanted: if dietunderlies the behavioural and ecologicaldifferences that separate extant taxa~ the reverse- that taxonomic differences may reflectdiffering dietary adaptations - may also holdtme. If this is the case, determining the correcttaxonomic position of Gigantopithecus may berequired prior to any accurate dietaryreconstmction. With this in mind, thecomparati ve approach taken here assesses thedietary skeletal evidence of Gigantopithecusbased on the underlying assumption that it doesbelong to a pongid (albeit a unique one), ratherthan a hominid, classification.

Dietary ConsiderationsIdentifying the anatomical evidence for

diet among modern primates contributesimmensely to our understanding of fossilprimate dietary habits. Following Ungar(2002), two lines of evidence can be used toinfer the diets of extinct primates. First,adaptive evidence "concerns analyses of the

sizes and shapes of jaws and teeth, and thethickness and stlUcture of tooth enamel" (Ungar2002:261). Second, material that pel1ains to theactual foods consumed by the individualrepresents nonadapti ve evidence. This wouldinclude studies such as dental microwear, stableisotopes, and trace element analyses.

One of the first attempts to reconstmctthe diet of Gigantopithecus occurred followingthe discovery of a giant ape's jaw bone in acliffside cave, south of the Yangtze River inChina. Within this cave, the skeletal remains ofdeer, boar, tapir, stegodon and rhinoceros werefound in association with the Gigantopithecusremains (Wen-Chung 1957). These associatedremains were all hoofed animals, and thus, couldnot have possibly climbed the 270-foot, nearverticaL cliff to access the cave' s mouth (Wen-Chung 1957). The only alternative, then, was thatthe giant ape canied these hoofed animals into thecave for food. Based on this evidence, and thesurface of the newly discovered teeth, Wen-Chung (1957:836) concluded that. "it wasobvious that the animal had a mixed diet of meatand vegetables, quite different from that ofmodem apes which live on fmit." This earlyreconstruction has been turned upside-down inlight of relatively recent paleoenvironmentalevidence which indicates that, "what are nowcliffside caves were sinkholes in a limestoneplateau when the giant apes flourished" (Simonsand Ette! 1970:83).

Thus, it seems apparent that the dietaryreconstmction of Gigantopithecus will not be assimplistic as previously thought. The traditionaland most conservative view, based upon jaw andteeth morphology, postulates thatGigantopithecus probably foraged on hard,fibrous matelial (Conroy 1990). This view, likeWen-Chung's (1957) inference. may also be injeopardy based on the numerous findings elicitedfrom studies of adaptive and nonadaptiveelements. The results of such studies shall bediscussed below.

Dental MOIphologySince it is the enormous size of the

Giganropithecus teeth that have so captured theattention of the primatological world, it is notsurprising that most adaptive studies have focusedon this particular aspect. It has long beenassumed that relative tooth size reflects functionalspecialization (Ungar 2002). Additionally,variations in relative tooth shape may reflect ameans of adapting to changes in the internal

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characteristics of foods, such as the strength andtoughness (Teaford and Ungar 2000).

Gigantopithecus teeth are colossal,exceeding all known primate teeth in theirdimensions. Even those of an adult male gOlillaare dwarfed by comparison. Data taken fromvon Koenigswald (1952) indicate that the thirdlower molars range from 22.3 to 23.1 nun inlength. Such dimensions place crown volumeestimates at about six times larger than modernhumans, and twice as large as adult malegorillas, when compared to the correspondingteeth of such individuals (Weidenreich 1944).Yon Koenigswald (1952:311) also rep0l1s that,"in addition to the usual five main cusps, thetooth pattern includes vil1ually all the secondarycusps that might possibly occur in the molar ofa higher primate." As in anthropoid apes, thesecond molar of the Gigantopithecus specimensare larger than the first molar. Additionally, themolars are distinctly longer than broad (Strauss1r. 1957). In Gigantopithecus bilaspurensis,cheek tooth cusps show little relief and areflattened and plate-bke rather than conical;occlusal surfaces are relatively broad and flat(Pilbeam 1970). Gigamopithecus blacki, whilesharing similar characteristics, tends to havehigher crowned and more cuspidate molars(Pilbeam 1970). It would seem that the molarsof Gigantopithecus are extremely large, withhigh, blunt cusps separated by deep, nan'owfUlTOWS(von Koenigswald 1952). It is alsoworth mentioning that the lower anteliorpremolar is relatively broad (as in Homosapiens) rather than elongated (Fleagle 1999).In other words. the premolars have become"molarized" - that is, broad and flattened.

With such enormous molars. it wouldcome as no surprise for this gigantism trend tocontinue into other teeth. However, relative tocheek tooth size, the incisors ofGigantopithecus are small and closely packedbetween the canines (Pilbeam 1970).Furthermore, the incisors appear to be almostpeg-bke, rather than chisel-like (Strauss. J r.1957).

The canine morphology ofGigantopithecus is also unique. Although thefront premolar was clearly bicuspid. the trigonidwas still quite developed, suggesting that theupper canine continued to shear against theantelior face of the lower premolars (Pilbeam1970). However, recovered canines show littleprojection beyond the plane of the cheek teeth.In both sexes, it seems, the canines wore do\\'nrapidly at the tips, even at an early dental age

(Pilbeam 1970). Thus, a stout and broadmorphology appears to characterizeGigantopithecus canines. Since the lower caninesof Gigantopithecus are truncated more thansharpened, it is reasonable to infer that themaxillary canine was not large, and not similar infunction compared to that of the gOlilla (Frayer1973). Frayer (1973:418) also reasoned that"since the mandibular canine was truncatedduring life, masticatory actions peIformed at thecanine appear to be more involved with grinding,than with sheming and gripping functions." Thus,the canines seem to have been essentiallygrinding teeth additional to the premolars andmolars. It should come as no surprise, then, thatthese morphological characteristics of the teeth ofGigantopithecus have led many to assume andpredict a diet of hard, tough, and rigid food itemsthat required heavy grinding and crushing.

Enamel ThicknessAttempts at dietary reconstructions have

also examined the implications of enamelthickness. Two adaptive explanations forpossessing thick enamel have been offered: eitherto prolong the use-life of teeth in an abrasive diet;or to minimize the lisk of crown damage givenhigh occlusal forces caused by a diet includingvery hard objects (Ungar 2002). In recent years,it has become common to describeGigantopithecus molars as thick enameled(Fleagle 1999; Klein 1999). If such is true. andthe assumptions underlying the adaptive functionof thick enamel are true. possession of thickenamel buttresses the data from dentalmorphology, which together suggests a hard,abrasive diet. possibly consisting of nuts, seeds,and subterranean tubers.

Mandibular MorphologyMandibular fragments are among the

most common bony remains found on fossilplimate sites. It is assumed that the architectureof this bone has been adapted to withstand thestresses and strains associated with oral foodprocessing. If this is the case. its morphologyprobably reflects (at least indirectly) some aspectsof diet.

The mandibular body of Gigantopithecusis extremely deep and highly robust. PoweIfulmastication is indicated by numerous features.The symphysis was long. deep and poweIfullybuttressed by large tOli (Yinyun 1982). That theface was short is indicated by the M, originationposition of the ascending ramus (Pilbeam 1970).Powerful mastication is also indicated by the

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everted and buttressed mandibular gonial angles(Pilbeam 1970). Short, deep faces are adaptedto powerful mastication (Klein 1999).

Dental orientation within the jaw isalso of interest. The "Theropithecus complex"describes a unique orientation whereby grindingteeth dominate - and may include adaptedpremolars and canines (Pilbeam 1970). Here,large grinding cheek teeth and powerfulmasticatory muscles are vital, while incisors arerelatively unimportant. Gelada baboons(Theropithecus), which subsist on small seedsand nuts - thus, grinding - represent a livingmember of the "Theropithecus complex."Surprisingly, if we review the aforementioneddata on Gigamopithecus dentition, we see that itshares many characteristics that belong to the"Theropithecus complex." Such dentalorientation within the mandible ofGigamopithecus may indicate a similar diet tothe gelada baboon - nuts and seeds. A study byGroves (1970) also shows "Theropithecuscomplex" -like characteristics in the modernmountain gorilla, whose differences with theeastern and western lowland gorilla mimic thosedifferences found between Gigantopithecus andits dryopithecine forbearers. Groves (1970)concludes that these similarities may provide abasis for inferring the Gigamopithecus dietfrom the mountain gOlilla diet. which includesroots, bark, and similar hard-wearing, bulkyitems.

Dental MicrowearAssociations between aspects of diet,

tooth use, and microwear in living primateshave been used to infer diets of fossil primates.Primates that often use their antelior teethduring ingestion have high densities andfrequencies of microwear striations on theirincisors (Teaford and Ungar 2002). Highincidences of long narrow scratches on molarspoint towards folivores. where as frugivorestend to have more pits than scratches (Teafordand Ungar 2002).

Yon Koenigswald's (1952:317)original assessment of the gross dental wear onGigamopithecus molars, "suggests the kind ofattrition found in man." It was later determinedby Strauss (1957 :685) that, "the occlusalsurfaces of the teeth appear to have been worndown considerably, so that the crown patternsof the molars cannot be made out." Such astatement lends critical support to the idea ofmassive gtinding and crushing as the primarymode of mastication in Gigantopithecus. These

early specimens were poor representatives; morerecent finds provide a better basis for micro wearanalysis.

Daegling and Grine (1987) examinedocclusal microwear in a sample ofGigantopithecus blacki teeth, and concluded thatthis species was unlikely to have been a hard-object specialist. In a recent study by Daeglingand Grine (1994), occlusal microwear on the teethof Gigantopithecus were compared withmicrowear on the molars of two extant bamboospecialists. Hapalemur griseus (the gentle lemur)and Ailuropoda melanoleuca (the giant panda).Bamboo feeding within the extant species doesnot produce a consistent pattern of microwear.Thus, because of these dissimilar patterns, dentalmicrowear on Gigamopithecus provides littledirect evidence for bamboo feeding. It does,however. appear unlikely that Gigantopithecllssubsisted exclusively on this particular resource(Daegling and Grine 1994). Rather, themicro wear patterns found on the molars ofGigamopithecus most closely resemble that of thepredominantly frugivorous Pan troglodytes. thecommon chimpanzee (Daegling and Grine 1994).This would suggest a diet of a broad range offruits and fibrous materials.

An alternative technique, based on theidentification of opal phytoliths found bonded tothe enamel surfaces of the teeth of fossil species,allows for the identification of individual plantremains eaten prior to death (Ciochon et al.1990b). The phytoliths found on Gigantopithecusmolars derive from two distinct taxonomic groupsand from different plant organs: i) the vegetativeparts of grasses, and ii) the fruits and seeds ofdicotyledons, specifically of a species in theMoraceae, or a closely related, family (Ciochon etal. 1990b). The relative roles of grasses and fruitsin the diet of Giganropithecus are difficult toestimate. However. judging from the presentfrequency of dental phytoliths in Gigantopithecus,fruits may have constituted a significant portionof the diet (Ciochon et al. 1990b).

Locomotor LimitationsWhile these dental aspects provide

invaluable clues to the diet of Giganropithecus,one must not forget the limitations imposed bybody size. Body size and stature estimates forGigantopithecus have ranged greatly: 10 feet talland 1200 pounds (Ciochon et al. 1990a), 9 feettall and 600 pounds (Simons and Ettel 1970), aheight of twelve or more feet (Wen-Chung 1957),and more recently, possessing a body mass of 300kg (Fleagle 1999). Given its size,

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Gigantopithecus must have been completelytelTestlial. Recently, the long bones ofGigantopithecus have been determined to be20-25% longer and more robust, on average,than those of living gOlillas (Johnson, Jr. 1979).Even this increase in size would most likelyresuict Gigantopithecus to a ground-dwellinglocomotor repertoire. This restriction hasnumerous implications on the type of foodstuffsavailable to Gigantopithecus, and will bediscussed in the next section.

DiscussionThe preceding sections discussed the

various lines of evidence that appear in thefossil record which indicate, either directly orindirectly, the diet of Gigantopithecus. Anoften overlooked vmiable in discussions offossil diets is the influence of the environmentalcontext in determining resource availability.Knowledge of the paleoenvironment dUling theexistence of Gigantopithecus providesadditional clues to its diet. Earlypaleoenvironment reconstructions, based mainlyon associated fauna, indicated a grassland andopen woodland terrain (Pilbeam 1970; White1975). More recent reconstructions (Ciochon etal. 1990a; Jablonski et at. 2000) suggestsubtropical-seasonal to tropical forest settings.Like the range of dietary reconstructionsdiscussed earlier, paleoenvironmental settingsalso vary, depending on the source. This shiftin reconstruction to a tropical forest setting isundoubtedly related to recent micro wear andphytolith analyses.

Locomotion and resource acquisitionare intimately linked, and are the directconsequences of environmental setting. Withsuch enormous mass. the only plausiblelocomotor pattem for Gigantopithecus wouldinvolve some form of tenestrial, quadrupedalmovement. This great size would, no doubt,eliminate any form of arboreality. Thus,foodstuffs in the diet of Gigantopithecus wouldbe limited to low-lying (or often-falling) items.As a general rule, fruits require high light togrow. and thus, in tropical forests most fruitsare found on terminal branches high-up in thecanopy ceiling (Campbell 1996). Suchpositioning would obviously be problematic fora ground-dwelling species such asGigantopithecus.

A number of explanations are possible.Perhaps body size reconstructions have beengrossly over-estimated. Until post-cranialremains are recovered which permit more

accurate body stature estimates, currentreconstructions must be taken with a grain of salt.It is unlikely, however, that a post-cranialskeleton indicating arboreality would beassociated with a dental and mandibularmorphology of such extreme size. Thus, itappears that a consensus has been reachedregarding the habitually ten-estlial nature ofGigantopithecus. Assuming this, how then wouldfruit become a vital pm1 of the diet, as wasposited by the phytolith analysis of Ciochon et al.(l990b)? First, perhaps some species of theMoraceae family represent low-lying fruits.Second, overly lipe fruit may fall to the ground,upon which Gigantopithecus could subsist.However, the chances of enough fruit, left toover-lipen by other frugivores, to sustainGigantopithecus is slim. Similm-ly. it is alsounlikely that Gigantopithecus could havesubsisted on fruit dropped by clumsy arborealfrugivores. Lastly, perhaps the indication ofheavy frugivory suggested by phytolithfrequencies is elToneous. For instance, only fourteeth, among thousands now discovered, wereselected for inclusion in the phytolith study. Suchselecti ve sampling introduces a large amount ofbias. Schwartz (1991) concurs, and notes that agreat deal is being drawn from the analysis offour teeth, only two of which were found to beharboming phytoliths, with the greatestconcentration on only one. Clem'ly, largersamples of teeth need to be similarly analyzed.Fm1hermore, phytolith accumulation propertiesneed additional investigation. Perhaps thephytoliths associated with various speciesaccumulate at differing rates and concentrations;or perhaps, the phytoliths only represent the last,or most recent, foods consumed by the individualprior to death, and not necessarily the mostcommonly eaten item.

The importance of Ciochon et al.· s(I990b ) phytolith study must not be entirelydismissed. As argued by Daegling and Grine(1994), bamboo feeding may be a possible dietaryadaptation for Gigantopithecus. Supportingevidence for such a scenario may be found inCiochon et al.' s (1990b) analysis. In addition tophytoliths belonging to the Moraceae family,phytoliths of vegetative grasses were alsorecovered. The recovery of such phytoliths maybe of some importance in that the bamboo familyis a vegetative grass. As stated earlier, additionalphytolith analysis need to be conducted.

A cautionary note is also wan-anted forthe dental micro wear analyses of diet ofGigantopithecus. Microwear features do not

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necessarily reflect specific food items per se,but rather the mechanical properties of thosefoodstuffs, or the constituents of thosefoodstuffs. Thus, the varying degrees ofmechanical properties of food items willproduce varying degrees of micro wear on thedentition of a specimen. This may createsomewhat of a "dental paradox", in that themicrowear found on dentition may not reflectthe most common, or distinctive, dietaryfoodstuff of that species, but merely the mostabrasive item within the dietary range of thatindividual (Plavcan et al. 2002).

As discussed earlier, one of the mostfrequently cited enamel con-elations is betweenthe consumption of hard, and abrasive, fooditems, and thick molar enamel. However, thickenamel by itself does not necessarily provideprotection against hard objects. whichcommonly cause fractures in the enamel layer(Ungar 2002). To prevent fractming, the bestprotection is "prism or crystallite decussation orinterweaving" within the enamel structure(Teaford and Ungar 2000:13508). Thus, it maybe the structure, and not the thickness ofenamel. that provides clues to the abrasive andrigid nature of dietary foodstuffs. If this is thecase, the thick enamel layers onGigantopithecus teeth may not provide evidencefor a diet that consists of hard, tough objects.To date, analyses of Gigantopithecus enamelstructure have not been conducted. and thus.statements and inferences about such propertiescannot be made.

It should also be noted that theenormous molar size found in Gigantopifhecusmay fall victim to the aforementioned "dentalparadox." During its existence. theenvironmental setting of GigallfOpifhccllShabitats experienced increasing aridity andseasonality (Jablonski ef al. 2(00). Theseclimatic fluctuations may have createdshortages of prefen-ed food items. The largemolars and increased occlusal surface may haveevolved as a coping strategy during periods ofstress in which dietary habits wen: forced toswitch to tough, fibrous, and hard food items.In this way, the enormous gross morphology ofGigantopithecus dentition is an adaptation forcritical feeding, and not a reflection of the mostcommon dietary foodstuffs.

ConclusionsReconstructing behaviour in any fossil

species requires an equiliblium hetween theenthusiastic inference of how organisms )jyed in

the past, and the skepticism necessary tounderstand the boundary between supportedhypothesis and unsuppOlted speculation (Plavcanef al. 2002). Reconstructions of the behaviour offossil species are limited to the evidence availablein the fossil record. This includes adaptive(dental morphology, mandibular form, andenamel structure) and nonadaptive (dentalmicrowear, gross tooth wear, stable isotope, andtrace element analyses) lines of evidence.

Though the cun-ent nonadaptiveevidence regarding the diet of Giga11fopithecus issomewhat controversial, the adaptive evidencedoes provide some important clues. Themandibles are deep - top to bottom - and areextremely thick. The molars are low-crownedand flat, with very thick enamel caps. The canineteeth are not sharp and pointed - as is the case inother apes - but are more similar to premolars inmorphology, while the incisors are small. peg-like, and closely packed. The features of theteeth, combined with the massive. robust jaws,lead to the conclusion that Giganfopithecus wasadapted to the consumption of tough, fibrousfoods through the extensive use of crushing andgrinding. However. as discussed previously, thisconclusion should be met with the same amountof skepticism as that given to other dietaryreconstructions, namely those of Ciochon et af.(1990b) and Daegling and Gline (1994).

Taken cumulatively. primary foodstuffsrelating to the reconstructed diet ofGiga11fopifhecus have included nuts. seeds.tubers. and other hard objects, grasses. bamboo,and fruit. Various skeletal elements point towardsdiffering sets of food items. It can only beconcluded that the most distinctive characteristicof the diet of Gigalltopifhccus is the enormousrange and variability of that diet. SinceGigantopithecus represents an ape unlike anyother in primate history. it should come as nosurprise that its diet should reflect its enigmaticpersona. As such. it probably exploited numerousresources, including all those mentioned here. Interms of dietary foodstuffs. the range of resourcesexploited by Gig({lIfOpifhcc/ls most closelyresembles that of anatomically modem Homosapiens (i.e. an opportunistic omnivore). Thisextensive range in diet. along with a giganticstature, are just two of the many characteristicsthat make GigQlIfollifhcc/ls an enigmatic ape.

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