Geomorphology to paleoecology: Gigantopithecus reappraised

Tim D. White Dqwment of Anthrqkhgy, Uniumity of Michipn, Ann Arbor, Michigan 48104, U.S.A. Received 31 May 1974 and accepted 23 September 1974

Geomorphology to Paleoecology: Gigantopithecus Reappraised Critical evaluation of the geomorphological circumstances leading to the formation of and deposition within the Gigantojithmus blacki cave near Liucheng, South China is undertaken on the basis of the available data. Models for the evolution of the Liucheng tower karst and the mode of deposition of the fossiliferous Quaternary sediments are presented. Analyses concerning the behavioural, phylogenetic and taxonomic placements of G. blacki are examined in the light of the proposed models. Hominid affinity or parallel in terms of behaviour or Savannah provenance are questioned.

1. Introduction In recent months, western medical technicians have shown enthusiastic interest in the most Chinese believe the technique to be a virtual Oriental practice of acupuncture: cure-all with nearly unlimited powers when applied to human ailments. Over thousands of years, the Chinese people have given faith to another sort of medicine. Ingestion of vertebrate and invertebrate fossil material was thought to be beneficial, and it was this belief that encouraged pharmacies to secure large stocks of vertebrate and invertebrate remains. Drugstores in Chinese cities have sold three kinds of medicinal fossils for hundreds of years, brachiopods, crabs and “dragon teeth” (von Koenigswald, 1952, p. 299). It is the “dragon teeth” fossils which have captured the interest of vertebrate paleontologists, these materials representing fossil mammalian remains excavated from deposits throughout the Chinese mainland. In the early 1900’s paleontologists began sorting through “dragon bone” collections in Chinese drugstores from Hong Kong to San Francisco. Despite strong Chinese belief in terms of their therapeutic value, the fossils subsequently purchased often proved of a nature surely not concordant with such expectations for vertebrate paleontologists. Difficulties concerning chronological placement and phylogenetic afhnity of much of the Chinese “drugstore” fauna1 material stemmed primarily from the uncertain provenance of the specimens. In a successful effort to obtain vertebrate fossil material in situ from China, Walter Granger and others from the American Museum visited the commercial deposits at Yenchingkou in the early 1920’s. This locality is a remotely-situated set of clogged sinkholes in a limestone ridge 520 m above the Yangtze River (Granger, 1938, p. 264; Figure 1). The deposits contained fossil mammalian remains now widely recognized as “Middle Pleistocene” in age. Yenchingkou became the type locality for the “Stcgodon-Ailuropoda” fauna, named after the dominant fauna1 elements. While Granger had taken significant steps in obtaining “drugstore” material in a geological context, other paleontologists continued to search the pharmaceutical collections. Such investigations assumed significance for primate paleontology when, in 1935, G. H. R. von Koenigswald brought to light a few fossil teeth of a new kind of animal which he named Gigantopithms blacki. The molar teeth were incredibly large, even when compared to those of a large male gorilla. They were clearly anthropoid and provoked a good deal of speculation concerning the size (estimated to range to a 9 ft upright stance Journal of Human Euolution ( 1975) I,2 19-233

220

T. D. WHITE Figure

1. Liucheng

Gigantopithecuc locality.

by Simons & Ettel, 1970, p. 80) and phylogenetic affnity of the represented individuals. teeth, von Concerning the drugstore fossils “associated” with the Giguntopithecus Koenigswald (1952, p. 299) stated: “Generally, all bony parts, including roots of the teeth, had been gnawed by porcupines . . . . There were no horse teeth but many of the porcupine and pig.” It was a broadly “Stegodon-Ailuropoda” fauna, but as with so many other “drugstore” fossils, the provenance was again uncertain. The yellow-earth matrix adhering to many of the fossils suggested that they had been derived from cave and fissure deposits in the provinces of Kwangsi and Kwangtung, Southern China. Paleontologists forwarded diverse opinions concerning the affinities of the fossil form. Weidenreich first called it a giant orangutang, but later changed his mind, referring it to the ancestry of Homo erectus (Weidenreich, 1937, p. 145 ; 1945, p. 86). Such hominid features as molar size, shape, morphology and wear have led various authors to agree with the hypothesis that Gigantopithecus represents a hominid ancestor, while others have argued for a consideration viewing the animal as an aberrant pongid (Simons & Pilbeam, 1965). The recovery of a nearly complete mandible from the Siwalik Hills of India in 1968, thought to date back to five to nine million years (Pilbeam, 1970, p. 517) has kindled renewed argument and suggestion of hominid ancestry for the genus (Robinson & Steudel, 1973; Frayer, 1973). In 1957 a key series of discoveries important for a phylogenetic analysis of the originally described taxon began. The Institute of Vertebrate Paleontology and Paleoanthropology

CIGANTOPITHECUS

221

REAPPRAlSED

of the Academia Sinica began to explore the cave deposits of the Kwangsi area of Southern China. These investigations resulted in the recovery of three partial mandibles and over 1000 isolate teeth of Gigantopithecus blacki (Woo, 1962). These discoveries are of critical importance in the appraisal of the giant primate. Since at present no complete documentation and analysis of the geomorphological circumstances under which the fossil material was recovered is available, an attempt is made here to synthesize them, while some of the hypotheses forwarded by paleontologists and anthropologists concerning Gigantopithecus will be examined in the light of the geomorphic data and their implications. 2. Location

and Description Gigantopithecus Cave

of

Southwestern China is the largest land surface on earth covered with carbonate rock. The area involved is roughly three times the total area of Great Britain (Kowalski, 1965, p. 78). The limestones of this part of the Asian continent (Huang, 1932) display such features as sinkholes, solution valleys, cliffs, underground streams, and cavesgeomorphological features characteristic of most karstic regions of the world. Southeast Asian karst areas have evolved and continue to exist under a subtropical to tropical climate and as a consequence, present both a greater rate of development brought about by increased temperature, humidity and organic acids as well as geological features completely absent in karst developed under temperate conditions. The Turmkarst (Gm.) or tower-karst of South China and North Vietnam represents a landform conditioned by such climate (SIlar, 1965, p. 35). Indeed, introductory texts in geomorphology often present ancient Chinese paintings depicting these typical limestone “monadnocks” or towers rising above the valley floor to exemplify a limestone landscape conditioned by tropical humid weathering conditions (Bloom, 1969, p. 33). The typical tower forms of South China karst are limestone peaks penetrated by numerous caves of various dimensions (Schworm, 1958, pp. 38-40). The inactive higher-level caves may be the sites of guano accumulations and phosphates (Jennings, 1971, p. 190). It is these biogenic deposits which have been exploited as fertilizer for hundreds of years by the farmers of South China (Wang & Huo, 1945, p. 7). The discovery of the first fragmentary mandible of G. blacki was made in 1956 by a peasant digging in the phosphate deposits of such a cave, later named Gigantopithecus Cave. The Chinese paleontological team working in the Kwangsi area immediately began excavation in the cave and eventually recovered two additional mandibles and more than 1000 isolate teeth from the locality. Of the other caves subsequently examined in the Kwangsi area, 88 out of 300 yielded mammalian fossils, and only one contained further Gigantopithecus teeth (Pei, 1965a, p. 44). Gigantopithew Cave is located in the southeast face of an isolated hill (a tower in the classic sense of Turmkarst), “Leng-chai-shan” in the Liucheng District of Kwangsi Provence, South China. The Li Kiang River, which now flows 3 km from the cave is shown in Figure 1 in relation to Gigantopithecus Cave. The cave entrances are located in the near-vertical sides of the tower, approximately 90 m above the local land surface. It has been observed that the caves and solution cavities found in the limestone towers of Kwangsi are generally distributed at three discrete levels with reference to the local land surface and water table. The highest caves in the area are those like the Gigantopithecus

T. D. WHITE

Figure 2. Gigmtopithws Cave: (a) horizontal section; (b) vertical section; (c) Transverse section at (b) X-X’. Scale identical.

Cave, 90 m above the local land surface. In the Liuchow region, the high caves are generally 80-100 m above the present land surface and are “tunnel-like” in shape. The caves of the lower two levels are often interconnected as they are in “Leng-chai-shan” (Pei, 1965a, p. 45). Excavation by the Chinese team has resulted in a detailed stratigraphy for the Gigantopithecus “cave”, this actually comprising a set of interconnected tunnels with three entrances. The stratigraphy is shown in Figures 2 and 3 which have been translated and modified from Pei (1965a). Reference to these schematic diagrams will be useful in the following consideration of modes of cave formation and deposition.

GICANTOPITHECUS

223

REAPPRAISED

Figure 3. Cigant~ithccus Cave: transverse from Figure 2.

P

I

-i

I

,

sections

with

stratigraphy

p’

224

T.

D.

WHITE

3. Formation Cave formation centered

has been a subject

on the relative

long debated

importance

modes

by geomorphologists.

of corrasion

The work of Davis in the early 1930’s various

of Gigantopithecus Cave

of cave formation,

(Davis,

and corrosion

this work

being

carried

was the notion of sub-water

cave formation.

(1932)

suggestion

occurs near the top of the permanent opposed

to Bretz’s

in the last two decades

though

the term shallow phreatic

Indeed,

Swinnerton

genetic

relation

Careful

survey

formation

China

features

would

(Credner,

exert

great

p. 41) observed,

of China

that the tower-karat

strong

corrosion

a variety

by renewed tectonics

the present

towers

Silar

followed

“lateral

“horns”

it expresses aggressive

con-

regime

tropical

water

As Tielhard

et al.

China region is complex. (1962)

and Silar

(1965),

and

tower karst proceeds in the folresult in disruption

(dolines)

(4b).

shaping

and

somewhat

corrosion

by vertical

When

of the

the base level

solution

ceases and

the basal parts of the hills, and

The resulting cones and towers rise from a common Further evolution can occur as the region is

of shapes.

or increased

precipitation.

The caves found penetrating

of original

solution

only towers resting

is seen in the expansion

of glacial

remaining-the

on the valley

cirques

which

floor.

results in

such as those of the Swiss Alps.

p. 36) attributes

corrosion”

the caves found within the South This undoubtedly

seen during tower evolution.

caves, but for the [email protected] morphology,

Chinese

of

of the area.

Fauna from the late

phenomenon.

of karst depression begins,

of the climate

evolution

It is under this climatic

by Gellert

and subterranean

to the point of leaving

to this action

its

as well

a fairly stable “warm

developed-the

seem to be the only evidence

have expanded

(1965,

which

of the South

and corrasion

rejuvenated

An analogy

has long retained

the karst history of the South

surface

of cave

to determine

nature

of certain fauna1 elements.

for the Turmkarst

(1954),

they occur.

of tower-karst

on the physiographic

of Kwangsi

the deepening

base level and exhibit

remnant

in which

the formation

1961, pp. 93-94).

making their slopes steeper (4c, 4d).

dolines

zone of action.”

must be analysed.

into small hills and depressions

is reached,

lateral

than Bretz’s,

the dynamics

that the monsoonal

China

responsible

4, the evolution

substrate

of erosion intense

concerning

a “Malayan”character

by Lehmann

manner:

limestone

South

characteristic

p. 200) aptly observe, in Figure

influence

times (Kahlke,

in the area seems largely As explained

1932)

by the persistence

develops

sistently until modern

locality,

within the limestone

indicated

lowing

of the region

literature

with the Gigantofiithecus

wet” climate, Miocene

cave investi-

views rather

for this important

development

contradictory

As Pei (1960,

detailed

has been preferred

action in

school of thought

p. 159) points out, “Most Swinnerton’s

(1942).

in terms of solution

makes it obvious that each cave must be studied individually

It was early recognized

(1935,

(1971,

of the

by Bretz

table or “phreatic”

that most formation

has confirmed

to the physiographic of the often

as the solutional South

1971, p. 140).

left little room for dispute as he stressed the fact that caves bear a

In dealing

origin.

forward

water table was the important

ideas, and as Jennings

gation

(Jennings,

argument

1931) led to a fuller consideration

Basic to this school of thought Swinnerton’s

Early

formation

cave

well below

network

the ground

China towers to the explains some of the

as well as other caves exhibiting surface

is more likely.

Features

similar in the

Gigantopithecus cave accord well with those given by Jennings (1971, pp. 156-157) as indicative of solutional work in the saturated zone, without definite currents. These features

include

wall and ceiling

pockets,

continuous

rock spans across cave chambers

GIGANTOPITHECUS

REAPPRAISED

225

Figure 4. Evolution ofTower Karst at Gigant@ithecusLocality: Tentative Model. (a) Formation of 90 m cave level at phreatic-vadose interface below flat plain. (b) Base level lowering by local tectonics. Formation of solution dolines and small hills. Formation of 50 m caves begins. (c) Further uplift brings 90 m caves into surface communication. Porcupine occupation and deposition of fossiliferous sediments. Early Pleistocene. (d) Uplift ceases with consequent 15 m cave formation at phreatic-vadose interface. (e) Further uplift and associated base-level lowering allows vertical and lateral erosion, present-day tower karst geomorphology attained.

226

T. D.

WHITE

and two and three-dimensional networks of passages (see Figures 2 and 3). An analogy may be seen in the African caves described by Jennings (197 1, p. 2 15). These caves are They are located on the face of a of phreatic nature but show vadose modification. 100 m fault scarp in the Congo. Similar features at an earlier stage of development have been uncovered by mining operations in Sarawak and Malaya. The alluvial plains here have been excavated, exposing limestone bedrock which is “intricately cut up in detail by subsurface solution but planed off flat overall” (Jennings, 1971, p. 191). With further uplift and associated base-level depression in this area, these caves could establish communication with the surface in the same manner postulated for the South China caves. The age of the tower karst in South China has never been satisfactorily documented. Gellert (1962, p. 379) shows that some regions displaying the towers have basalt-sealed valley floors overlying a Mio-Pliocene fauna which could indicate a minimum age for the towers in these areas. As noted by Silar (1965, p. 42), development of the tower karst was substantially influenced by tectonic uplift. The Yenshan folding and Himalayan fault tectonics seem to have been responsible for the necessary changes in base level. Early work indicated that a Shanyuan Pliocene peneplanation occurred, with subsequent rejuvenation resulting in the “gorge-cutting period” seen in the Yangtze gorges of a more northerly area of China. Barbour (1935, pp. 46-52) noted the overzealousness with which early investigators apphed concepts of peneplanation, but reasserted the usefulness of the concept as applied to South China as well as Central China. Tielhard et al. (1935, pp. 201-203) placed the peneplain rejuvenation at the end of the Pliocene, noting that the tops of “pillars” in South China represented an old erosional plain. The dating of the inferred tectonic episodes was influenced by the assumed age of the Yenchingkou fauna, erroneously thought to have been deposited on the Yangtze floodplain. The actual date of the tectonic uplift which brought the 90 m Gigantopithecus cave at Liucheng above the watertable is presently unknown and is of minor relevance to paleontologic considerations. It is the age of initial communication of the cave with the surface which assumes importance in this regard. Such opening of what now remains of the cave network probably occurred shortly before accumulation of the fossiliferous deposits (Pei, 1965a, p. 46). Rejuvenation of the tower karst in the Liucheng area occurred with subsequent tectonic activity. The second and third levels of cave were brought to the phreaticvadose interface by these tectonic periods, and formed here during relatively quiescent periods as seen in Figure 4. As indicated by Sllar (1965, p. 42), active seismicity in the area today indicates that uplift is probably continuing. Presumably cave networks are forming in the limestone below the present tower karst valley floors in South China, and further tectonic episodes will result in further lowering of base level, subsequent erosion, and eventually, a fourth level of caves. A schematic representation of the model proposed for development of the Liucheng tower and Gigantopithem Cave is presented in Figure 4. It is of critical importance in the interpretation of the Gigantopithem locality to note that geomorphological evolution here was not at all analogous to the situation described for the Yenchingkou fauna1 locality. In a recent review of the Gigantopithecus material (Simons & Ettel, 1970, p. 83) it was stated that “What are now cliffside caves were sinkholes in a limestone plateau when the giant apes flourished”. From the work of Granger (1938, p. 266) and more recent analyses of the fauna1 material recovered from

GIGANTOPITHECUS

REAPPRAISED

227

Yenchingkou (Colbert & Hooijer, 1953, p. 10) it appears that here, true sinkhole accumulation prevailed as a mode of deposition. Solution dolines in the form of vegetationobscured, steep-sided shafts acted as traps for unwary animals at Yenchingkou. The model presented above for the evolution of the Gigantopithecus cave, the morphology of the cave, and the nature of the fossil fauna itself are not analogous to the situation of Yenchingkou. It seems reasonable to expect that the modes of deposition of the fauna1 material at Yenchingkou and the Gigantopithecus locality would have been different as well.

4. Deposition

in Giguntopithecus Cave

A generalized stratigraphy for the 90 m caves near the Gigantopithecus locality is given as follows by Pei (1963, p. 222). A nearly horizontal layer of stalagmitic crust is usually found at the base of the deposits, resting on the cave floor and succeeded by a layer of red clay. Above the red clay is a consolidated, yellowish-brown sand and clay in which fossil material is concentrated and from which the Gigantopithecus specimens were derived. Finally, the deposits are capped by another stalagmitic layer. The G. blacki cave at Liucheng has a stratigraphy which follows this general scheme as seen in Figures 2 and 3. The lowest layer of stalagmitic crust is found on the cave floor beneath a red clay. Jennings (1971, pp. 176-l 77) reports that similar red clay is found widely in limestone caves in many climates, placing doubt on Pei’s inference of a warm and humid climate on the basis of this stratum (1965a, p. 46). The Gigantopithecw Cave red clay deposits follow Jenning’s description in detail: “They are unbedded but liable to desiccation cracks, into which other material may be introduced subsequently”. The cave shows such cracks, which contain fossil twigs and leaves (Pei & Li, 1958, p. 199), indicating not only desiccation but certain communication of the cave network with the ground surface or tower side at this stage of its evolution. Kowalski (1965, p. 79) states that the concretions in this layer as well as in the fossiliferous layer are rolled, indicating deposition by running water, but Jennings (1971, p. 175) makes it clear that such features are often seen when small blocks are partly rounded by solution before detachment in caves. On the other hand, the presumed proximity of the cave opening, the climate, and the occurrence of some fossil material in constricted portions of the cave may be indicative of reworking by water. The consolidated yellow sand and clay layer which contained the G. blacki material as well as the remainder of the fossil fauna in the Liucheng cave can be chronologically placed (in a strictly relative sense) according to the fossils. Early analysis led to the conclusion that the fauna was Middle Pleistocene, contemporary with the rest of the “Stegodon-Ailuropoda” fauna in China. Chow (1957, pp. 394-399) assigned the fauna to an Early Pleistocene or even Pliocene age because of isolated chalicotherid and Mastodon sp. molars. He called it the “Gigantopithecus-Stegodon-Ailuropoda” fauna. Pei (1965a, p. 44) follows this analysis, calling the assemblage a “Gigantopithecus” fauna characterized by archaic elements such as Stegodonpre-orientalis Young, Trilophodon, Gigantopithecw, Ailuropoda microta and Equus yunnanensis. The presence of the distinctive Hyaena licenti Pei (Pei & Li, 1958, p. 199) allows correlation with the “Nihowan” fauna of North China. Chang (1968, p. 25) states that the Liucheng Gigantopithecus fauna is the temporal equivalent of the Nihowan fauna of the North. The significance of this correlation is emphasized by the finds at Hsi-hou-tu in North China. Here, a clearly artifactual stone tool assemblage

228

T. D.

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was found in “direct association” with a typical Nihowan fauna (Chang, 1968, p. 41). Kahlke (1961, pp. 83-108) described the fauna typical of the Liucheng locality as “Early Middle Pleistocene”. While the fauna does lack a “Villafranchian character” as defined for European or African assemblages, it is perhaps worthwhile to consider the case of the Djetis Formation in Java. Here, associated with hominid fossils of indeterminate (Modjokerto) and early pithecanthropine (P-IV) affinity, the fauna was called “post-Villafranchian Middle Pleistocene” by Hooijer in 1968 (pp. 86-90). Recent absolute dates for the Upper Djetis at 1.9 * O-4 MY (J acob & Curtis 1971, p. 50) have led investigators to speak of the fauna in the Upper Djetis as “javanese Villafranchian”. Comparative study may shed light on the relative dating of the Liucheng fauna, but for the present time, placement as “Early Pleistocene” seems warranted. It has been noted that most “drugstore” fossils derived from South China were isolated teeth, often without root. Early writers (Weidenreich, 1946, p. 64; von Koenigswald, 1952, p. 299) correlated this universal characteristic of the fauna1 remains with the presence of Hystrix, the porcupine. Destruction of much of the material seemed due to the gnawing of these rodents, apparently in their attempt to incorporate the calcium needed in quill production (Prater, 1965, p. 216). As Young (1932, p. 386) and Pei (1963, p. 221) have recognized, an identical fragmentary nature of the fossil fauna is present in deposits throughout South China. In a paper dealing with the state of preservation of the fossils in the Giguntopithecus Cave, Li (1960, p. 404 observes that rodents played a “predominating part in the destruction”, with plant and groundwater action The complete nature of the Yenchingkou fauna has been of secondary importance. attributed to the local absence of Hystrix (von Koenigswald, 1952, p. 301). This is As previously noted, the mode of deposition at probably not the proper explanation. Yenchingkou was almost certainly as different as was the mode of cave formation. It was precisely this mode of deposition and associated geological circumstances which excluded Hystrix as an agent of destruction and at the same time allowed for the uncommonly complete state of the fauna1 material at Yenchingkou. Simons & Ettel (1970, p. 83) state that carnivores predominate in the fauna of the Gigantopithecus Cave. Forms found here include Hystrix, Ursus, Ailuropoda, Hyaena, Cuyon and Arctonyx (Pei, 1963, p. 223). Besides being fragmentary, most of the fauna is “. . . either young with milk dentition or old individuals with teeth greatly worn down” (Pei, 1957, p. 50). Such observations have led investigators to consider the Gigantopithecus deposits as predator bone accumulations with (Dart, 1960, p. 144) and without (Simons & Ettel, 1970, p. 83) reference to the giant primate as a predator. Fairly convincing evidence in favour of non-primate predator/scavenger accumulation has been brought to light for the Swartkarans australopithecine-bearing cave breccia by Brain (1970, pp. 1112-I 118). In the case of the G. blacki locality at Liucheng, it would seem preferable to use the word “collector” instead of “predator” with specific reference to the porcupine. The habits of this animal are shown to be highly concordant with the nature of the fossil fauna, even to the point of the animal’s inhabiting similar limestone crevices and cliffs from North India (McCann, 1928, p. 214) through Mesopotamia (Pitman, 1925, p. 831) and Africa (Alexander, 1956, p. 258). Accumulation of skeletal material seems to be an integral part of this animal’s behavioural repertoire, and has been repeatedly observed in the wild (Pitman, 1925, p. 832; Prater, 1965, p. 216) as well as in captivity (Alexander, 1956, p. 258). The predomination of juvenile and old individuals in the fauna represents simply those segments of any population most susceptible to death: the agent active in

bone accumulation

is not necessarily

229

REAPPRAISED

GIGANTOPITHKCUS

the agent responsible

for the death of the organisms

it accumulates. Thus,

in terms of the actual

the Giguntopithecus

suggest that the information modes of deposition been considered the

(Simons

evidence is that

Simons

sediments

& Ettel

within

(1970,

p. 83)

does not allow a choice among the three alternative that of sinkhole

accumulation,

has

valid in the case of Yenchingkou,

it does not seem to fit

available

locality.

for

the

of water-washing

1970, p. 83;

the fossil remains

of the fossil-bearing

seems in order.

The first hypothesis,

While

mentioned & Ettel,

available

they offer.

above.

geomorphologic

alternative

mode of deposition

Cave, a reconsideration

Gigantopithecus

Pei, 1963, p. 222).

and requires

a somewhat

The

of the fauna1 material This alternative

absurd

probability

into

second the cave

ignores the nature

statement

which

of

asserts

that the caves were filled by an input of surface runoff alone, that this runoff contained solid discharge the discharge resulted priate

with incredibly was composed

in the transport to question

high proportions

of predominantly

and deposition

the lack of expected

corrasional

material

into the Giguntopithecus

Noah’s oversight Clearly,

the attribution

likely explanation

of the fossil material

perhaps

is an agency

the replacement

nearer

area

the most

of Gigantopi-

to wash the fauna1 improbable

as

represent

to, or perhaps

113), is the most

in the South China Secondary

of the

deeper into the recesses of the cave

in the filling of constricted of the Liucheng

with the documentation

arguments

caves in

washing

portions

of “Gigantopithecus-Stegodon-Ailuropoda”

combined strong

observed

entrances

agent of the porcupine,

1970, pp. 1110-I

Cave in particular.

and third levels (50 m and 15 m caves)

vanced fauna1 complement, Liucheng

required

associated

Perhaps

event as seemingly

to the collecting

(Sutcliffe,

cave or burrow

likely to have resulted

Certainly

second

the hyaena

Gigantopithecus

from the original

height.

Cave.

for the extinction

capability

of the fossil bone accumulations

and the Liucheng

material

and selective

Cave is a geological

it seems appro-

features

of the world’s largest primate.

and to a lesser extent general

and depositional

is its provision

a

and that

If water-washing

G-3 mandible,

of the Gigantopithecus

aspect of the second hypothesis

A flood of the magnitude

thecus.

skeletal material,

young and old animals.

of the immense

with such inflow in the geomorphology attractive

of vertebrate

fauna

locality of tectonic

for the original

opening

to a roof-level in the

by a more adactivity

in the

of the 90 m cave

even with, the local ground surface at the time of accumulation. 5. Discussion

Through

a comprehensive

examination

of the geomorphology

of the Liucheng

locality

and an integration

of the taphnocoenic

data with this analysis, it becomes clear that many

statements

by anthropologists

concerning

made

certainly

imaginative,

unbased

propriate

in this regard:

“It is patent,

south of Africa was inhabited thecine

series or variety

and highly

however,

during the Early

of mankind

this giant

improbable.

fossil form

The

following

are,

havioural already

statements advanced.

affinities,

Pleistocene

by hunters

(G. blacki in this case),

the fact that the data presented

has some importance Some

have stated

is ap-

that the far east of Asia, like the far of the australopi-

that they customarily

in caves and were cannibalistic . . .” (Dart, 1960, p. 144). While it is beyond the stated purpose of this paper to deal with Gigantopithecus of its phylogenetic

although

passage

for both behavioural

that the morphological

lived

in terms

above allow few firm beand phylogenetic attributes

schemes

of the creature

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“The existence of several T-complex features in argue for an open country adaptation: the dentition of Gigantopithecw sheds some light on the giant ape’s probable environment.” (Simons & Ettel, 1970, p. 82.) This morphological aspect of the Savannah provenance concepts center on the anatomical correlates of a heavily-masticated diet, often supposed to consist of cereal grains from grassland context (Jolly, 1970, pp. 5-24). In terms of habit inference, to reconstruct Gigantopithecus blacki as a Savannah dwelling primate strictly on the basis of anatomical considerations is to imply that either heavily-masticated diets are restricted to animals occupying grassland habitats, or that the evolution of an apparatus adapted to this kind of diet is impossible for forest or woodland animals. Morphology has also been employed in attempts to emphasize the “hominid” affinity of Gigantopithecus, usually citing the presence of parallels in dental and mandibular anatomy between gigantopithecines and australopithecines (Eckhardt, 1972, p. 102; Robinson & Steudel, 1973, pp. 525-526; Frayer, 1973). The proportion of caries on the teeth of G. blacki would seem to substantially discount this analogy with early hominid forms (Woo, 1962, p. 93). Of 911 Gigantopithecus teeth examined, 9.8 % displayed authentic caries, while slightly more than 1-O% of australopithecine teeth examined displayed such decay (Clement, 1956, pp. 4-7). Of more importance and demonstrable irony is the fact that the 1964 monograph by Davis on the giant panda showed nearly character for character the same complex which was later observed by Jolly (1970) for Theropitheczu and has become known as the “T-complex” referred to above. In fact, Davis made specific reference to the South African australopithecines, clearly recognizing the evolutionary parallels which exist (p. 128). Thus, the morphological similarity in this case implies evolutionary parallel instead of phylogenetic affinity as well it might in the case of Gigantopithem and the australopithecines. If the morphological characteristics observed in G. blacki allow no firm statements as to the Savannah provenance of this primate, paleoecological reconstruction of the animal’s habitat must rely upon associated fauna. The fauna1 data as published are patently inadequate in terms of this kind of definitive analysis, but this has not deterred the placement of Gigantopithecur in an “open woodland or savanna” habitat compared to such related primate stocks as Gorilla and Pan, which remain in the “tropical forest” (Simons & Ettel, 1970, p. 84). Although not enough proportional figures are yet available, indications seem somewhat clear. The hyaena and dog are extremely adaptive forms, not diagnostic in terms of forest/grassland indication, and the rhino and porcupine are represented by forest-dwelling forms in the same area of the world today (Prater, 1965). More diagnostic paleoenvironmental indicators such as the horse and chalicothere are represented by a few isolated teeth in the first case and a deciduous molar in the second. When the sampling agency involved in bone accumulation is considered for the Liucheng locality, it becomes obvious that the presence of more than 1000 isolated teeth and three mandibles of Gigantopithecus, a predomination of forest forms such as the panda and tapir, and a few isolated equid and bovid teeth cannot support the contention that “. . . one can probably infer a similar non-forested habitat for G. blacki.” (Pilbeam, 1970, p, 518.) What then can be said about G. blacki? The available data have been shown to support precious few of the speculative statements found in the literature dealing with this giant primate. Unfortunately, little can be said about this Early Pleistocene form of Southern Asia. Its late temporal placement probably excludes it from any direct ancestry of the hominid line. It probably stems from a form like C. bilaspurensis of the Dhokpathan in India-a form recently advocated as a plausible Pliocene alternative to the supposed

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hominid ancestor Ramapithecur (Frayer, 1973, p. 420). G. blacki, contrary to Frayer (1973, p. 420) can probably not be “best explained” as an aberrant hominid, as it lacks the critical evidence for either culturally-patterned behavior or bipedal locomotion. Paleoecologically, the evidence is not conclusive in terms of habitat placement of G. blacki, and allows a variety of alternative reconstructions. This large (gorilla-sized or larger) terrestrial or semi-terrestrial (in the sense of Gorilla) animal is associated with a primarily warm, semitropical forest fauna. It seems to be adapted to a diet both high in carbohydrates or starches, and requiring heavy mastication. Selective pressures seem to have led to a masticator-y pattern convergent onor parallel to thatof theaustralopithecines, with strongly buttressed mandibular morphology, relatively small anterior teeth, and large buccolingually expanded premolars and molars. It is highly interesting that the same anatomical complex adapted to heavy mastication as well as the same inferred and observed habits are characteristic of Ailuropoda, the panda. Tantalizing information regarding these ecological alternatives for G. blacki includes the fact that the dwarf panda Ailurofioda microta found in the Liucheng Gigantopithecus Cave fauna (Pei, 19656, p. 45) shows no caries. The giant panda, developing large size and inhabiting a wider geographic range concomitant with the spread of the bamboo forests in the Middle and Late Pleistocene displays 10.7 % caries (Woo, 1962, p. 93). Could it be that a primate occupation of and subsequent replacement within a panda-like niche was occurring in the forests of Southeast Asia during Lower and Middle Pleistocene times ? Rather than asserting a Savannah provenance and cereal-grain grinding masticatory complex as hominid indicators for G. blacki, it is possible to view this primate as an anthropoid adapted to a panda-like niche within a forested habitat in the Lower Pleistocene. In conclusion, I think it fair to state that a realistic evaluation of all the available data sheds a different light on, or perhaps removes some artificial illumination on the giant primate of South China, Gigantojithecus blacki. Through an examination of the geomorphological data it has been possible to place serious doubt on previous statements dealing with this extinct form. It is perhaps too obvious to stress the necessity of evaluation of fossil evidence in the light of all of the available data, but it is certainly more obvious that such evaluation has been somewhat lacking in all too many past examples. In the case of Gigantopithecur blacki, alternatives concerning behaviour, provenance and phylogenetic affinity must remain open, awaiting the addition of further anatomical and paleoecological information. I would like to thank Mr Shigeki Maruyama for his patient translation of the difficult Chinese geological descriptions. My thanks are also due to Dr D. Eschman, Department of Geology and Drs F. Livingstone and M. Wolpoff, Department of Anthropology at the University of Michigan, who critically commented on versions of this paper. Also, my sincere appreciation goes to David Frayer for his thoughtful and often stimulating discussion concerning this subject. Finally, the financial assistance of the National Science Foundation Graduate Fellowship Program is gratefully acknowledged. The conclusions and interpretations are those of the author. References Alexander, A. (1956). Bone carrying by a porcupine. South African Journal of Science 52,257-250. of China, Memoirs. Series A: Barbour, G. (1935). Physiographic history of the Yangtze. GeologicalSuq 14, 112pp. Bloom, A. (1969). The Surfme of the Earth. Englewood Cliffs: Prentice-Hall.

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