Chimpanzee subsistence technology: Materials and skills

Chimpanzee Subsistence Technology: Materials and Skills

Geza Teleki Department of Anthropology, Pennsylvania State University, 409 Social Sciences Building, University Park, Pa. 16802, U.S.A.

Beyond reviewing basic data on Primate technological behavior, the aim of this report is to document the nature of chimpanzee technical skills by examining some of the mental as well as physical expertise which chimpanzees bring to bear on subsistence activities. This task is approached along several avenues : first, an outline is drawn of current knowledge about chimpanzee subsistence technology throughout Africa; second, a sketch is made of technological variability in several chimpanzee populations; third, the results of a personal investigation into the skills needed by chimpanzees to probe for insects are provided; and fourth, a comparison is made of the baboon, chimpanzee and human techniques used to exploit termites as a food resource. Instead of focusing on the unique features of human subsistence technology, the report attempts to show that many technical skills are and probably were firmly rooted in Primate prehistory, well before the advent of the earliest hominids. An integrated model of technological achievements among extant Primates, based on a sample ofAfrican cercopithecid, pongid and hominid populations representing stages in a phylogentic sequence, is offered as a foundation for reconstructing the gradual evolution of primate subsistence technology. This approach is intended to provoke contemplation and discussion among those investigators of human behavior and society who favor the thesis that the criterion of technology, together with the cultural transmission of technical skills, separates the human from the nonhuman Primates.

Received 15 July 1974 and accepted 8 August 1974

1. Introduction Research

on the technological

a century.

capabilities

of captive

for use in specific

tasks range

from

to the spontaneous

the designed

Kijhler

(1925)

1973).

It was merely a decade ago, however,

yield information 1964;

on their

Kortlandt,

The

chimpanzees

Studies on the ability of pongids to manipulate, behavior

technological

experiments

observed

in compounds

that research

performances

describing

such technological

using” and “tool-making”

performances

fauna

Kortlandt

1963,

1965;

serve as an introduction

achievements

observed

& Kooij,

conditions

by

(1972,

began to (Goodall,

skills exhibited

and on the materials

and specifically

on those attributes

activities-namely,

any behavior

and/or

substances

preparing

among nonhuman

1963;

Primates

van Lawick-Goodall,

and other

1970) should

Only the salient features of nonhuman Primate My attention will center on the will be considered here.

technical

patterns of behavior nests with branches, attention. Three main points

is now expanding at such Thus, reviews of the “tool-

to the subject.

behavior”

habitats,

oriented, for

utilized by chimpanzees

underlie

via technological

ingestion.

this limited

living in natural

which occur in association Due

involving objects (e.g. displacement grooming

years ofwork with chimpanzees

to

extension,

this

with subsistence toward obtaining

orientation,

many

other

drumming on buttress roots, building with leaves) will receive only cursory

focus.

in Gombe National

One,

having

Park, Tanzania,

Joumal of Human Evolution (1974) 3,575-594 10

in cages

by Menzel

on wild chimpanzees

in natural

a rate that all the data cannot be reviewed in this briefreport.

“technological

and modify objects

conducted

1965).

literature

(Hall,

spans more than half

assemble

completed

nearly

two

these field experiences

576

G.

TELEKI

have inevitably funneled my interests and insights in the direction of naturalistic behavior. Two, laboratory research on the skills elicited from chimpanzees and other Primates can provide vital information about the behavioral and psychological potentialities associated with manipulation and modification of objects, but only field research can establish the extent to which these potential capabilities are actually realized in natural populations. And three, knowledge about the realized subsistence technology of extant chimpanzee populations may shed light, albeit indirectly, on the evolution of technical skills among Primates in general and among the early hominids in particular. 2. Contexts

of Technological

Behavior

In order to provide some comparative perspective to the study of chimpanzee subsistence technology, the general dimensions and contexts of both baboon and chimpanzee technical skills are outlined below. A sketch of technological behavior in these African genera is appropriate for the following practical and theoretical reasons: because (a) many populations within these genera have been thoroughly studied in field conditions (Baldwin & Teleki, 1972, 1973); (b) the basic subsistence patterns of baboons and chimpanzees are similar, both being omnivores that collect flora and prey on fauna, and their collectorpredator habits can be ranked along a graded spectrum of diversity and complexity which integrate with human gatherer-hunter habits in the same African regions (Teleki, in press) ; (c) numerous populations of these genera live in areas that lie within or near to the African rift system, in habitats that range from arid savanna to rain forest, and thus inhabit geographical and ecological zones which were associated with hominid emergence (e.g. Pfeiffer, 1969); and because (d) these more than any other Primate genera have been cited as models for reconstructing the behavior, social life and subsistence habits of early hominids (e.g. Washburn & DeVore, 1961; Mann, 1972). In comparison to humans and even to chimpanzees, wild baboons exhibit a distinctly limited repertoire of technological behavior. Although Primates in general tend to apply their technical skills mainly in contexts other than subsistence (Hall, 1963 ; Kortlandt & Kooij, 1963), the meager records of technical performance among baboons are in fact restricted to this one context. Marais (1969) observed chacma baboons break open hard fruits with stones; Kortlandt & Kooij (1963) reported accounts of baboons using objects to crush a scorpion before eating it, to widen the entrances of ant and termite nests and to stir up insects under rocks; and van Lawick-Goodall, van Lawick & Packer (1973) observed a baboon wipe her lips with a small stone after feeding on sticky pods. Several of these manipulative performances are similar in function and execution to those which will be described under chimpanzee subsistence technology, but the levels of sophistication routinely reached by chimpanzees, especially in modification and transportation actions, are qualitatively and quantitatively beyond those achieved by baboons. For example, when considering the many thousands offield observation hours spent with baboons during the past decade or so, this list is seen to contain a mere sprinkling of incidental cases which fall rather short of establishing baboons as technologically adept Primates. This sharp limitation is in some respects anomalous, for baboon subsistence patterns are in many other ways quite closely analogous to those of omnivorous chimpanzee and human populations, especially in predatory and hunting behavior (Teleki, in press). Wild chimpanzees, on the other hand, occupy a firm position as the most technically skilled of all living nonhuman Primates, at least at the time of this writing. These

CHIMPANZEE

African lation,

apes exhibit

a rich array of technological

modification

often highly

and transportation

sophisticated

other contexts,

SUBSISTENCE

technical

behavior

of many

577

TECHNOLOGY

that incorporates

types of materials;

skills are applied in subsistence as well as in many Typical agonistic actions include shaking,

both agonistic and nonagonistic.

flailing, dragging,

throwing,

hitting and thrusting

with trees, branches,

sticks, twigs, fronds,

stones, earth clods, leaves and even food items (van Lawick-Goodall, 1968).

And nonagonistic

investigate

probing,

technical

displacement

social playing-situations and other materials

performances

1971;

(Nishida,

items,

have

behavior,

Kortlandt

(van Lawick-Goodall,

1973).

1963,

also yielded 1965,

and

leaves

in order to perform

1968)) and the Kasakati-Mahali

field experiments

valuable

information

in the use of objects

1967,

1968;

leopards

on their

as weapons

Zon & Orshoven,

exposed baboons (Eimerl

(Teleki,

in Uganda & DeVore,

1973a).

1967). between

leopard

Kortlandt,

in Guinea

potential

(Albrecht

Moreover,

to a mechanized 1965;

conducted

leopard dummies and other

not yet been observed in interactions

including

but not weapon-use

Numerous

were exposed to mechanized

and especialIy

have, however,

live carnivores,

solitary

subsistence have been recorded in Uganda (Sugiyama, 1968),

sites:

Park in Tanzania

Kortlandt,

responses

body cleaning,

sticks, twigs, vines, grasses,

in contexts other than Budongo Forest

study

and Zaire, in which chimpanzees unfamiliar

nest building,

and to fetch objects from one place to another

chimpanzee

area in Tanzania

nological

Kortlandt,

task.

Technological Gombe National

grooming,

1970;

during food and water collecting,

in which stones, branches,

distances,

some premeditated at several

skills are exhibited

are manipulated and sometimes modified to suit specific functions 1970). Ch’rm p anzees are also known to transport certain objects for

(van Lawick-Goodall, considerable

the manipu-

and their varied,

for tech& Dunnett,

Similar

‘attack’

chimpanzees

similar

and

tests in which

elicited frenzied

attacks,

pers. comm.).

This catalogue of baboon and chimpanzee technical skills shows that the frequency and distribution of such performances differ remarkably in these particular cercopithecid and pongid genera. with incidents

Reports

of chimpanzee

of technological

behavior,

only sparse cases at a very limited technical

capabilities

collector-predators, ators and human

of baboons

field research

number

of sites.

and chimpanzees

of the discrepancy

The discrepancy is reminiscent,

in the capabilities

Africa

are laden

field research

contain,

observable

even though

of chimpanzee

in the:

both are

collector-pred-,

gatherer-hunters.

3. Chimpanzee The technical

skills of chimpanzees

of subsistence,

and their technological

as well as between

throughout

whereas reports of baboon

Subsistence

are highly diversified repertoire

Technology

even within the special context

seems to contain

some flexibility

within,

populations.

(a) Skills exhibited and materials utilized The technical

skills and materials

used in naturalistic

food-getting

activities

are shown

in Table 1. When summarized, these include : pounding of hard items, such as nuts and. hard-shelled fruits, with sticks and stones; probing for honey and insects, usually ants or termites, 1OA

with twigs, vines, grass stalks and bark strips;

prying open the nests of arboreal

1

probing probing

prying (var. on probing or digging)

honey ants

ants

6

7? 8

Mt Okoro Biko

Rio Muni

Zaire Tanzania

Gombe National Park

?

termites

5

near Ayimken

termites

termites

probing

probing

probing

probing

Rio Muni

Cameroon

5

pounding pounding pounding

Manipulation

Dipikar Island

nuts palm nuts hard-shelled fruits termites

Food type

sticks, twigs

twigs sticks, twigs vines, grasses

sticks, twigs

sticks, twigs

sticks, twigs

stones stones sticks and stones sticks (twigs)

Modification

chewed and worked smooth breaking to suitable length, stripping of leaves and other projections breaking to suitable length, stripping of leaves and other projections breaking to suitable length, stripping of leaves and other projections ? breaking and biting to suitable length, stripping leaves and other projections breaking to suitable length

none( ?) none noted none noted

technology

Objects used

subsistence

Rio Muni

I? 2 3

no.

Map

data on chimpanzee

4

?

?

Site

Vital

Tai Forest Reserve regional survey

(West Africa) Liberia Ivory Coast

Country

Table

& Sabater

Pi (1969)

van Lawick-Goodall (pers. observ.)

(1970), Teleki

Merfield & Miller (1956) van Lawick-Goodall (1968, 1970)

Jones

Jones & Sabater Pi (1969)

Savage & Wyman (1843-44) Beatty (1951) Rahm (1971), Struhsaker & Hunkeler (197i) ‘Struhsaker & Hunkeler (1971) Jones & Sabater Pi (1969)

References

Tanzania

Makolongo & Kabogo Pt Mahali Mtns

Tanzania

Basin

Kasakati

Tanzania

prying (var. on probing or digging) wiping (var. on sponging)

ants

ants

probing

probing probing

honey termites ants

probing

termites

ants

sponging

water

leaves

sticks, twigs vines, bark strips sticks

twigs twigs, vines

twigs, bark strips twigs, bark strips

leaves

sticks, twigs leaves

probing sponging

honey brain tissue

sticks twigs, vines grasses

digging

termites

ants

crumpled by hand into wad

none noted

breaking and biting to suitable length, stripping of leaves and projections

breaking to suitable length, stripping of leaves and other projections none noted none noted

breaking to suitable length, stripping of leaves and other projections none noted chewed into an absorbent mass chewed into an absorbent mass breaking to suitable length

breaking to suitable length

Nishida (1973)

Nishida (1973)

Nishida (1973)

Izawa & Itani (1966) Izawa & Itani (1966)

Suzuki (1969)

Suzuki (1966, 1969)

van Lawick-Goodall

van Lawick-Goodall Teleki (19738)

van Lawick-Goodall (pers. observ.) van Lawick-Goodall (pers. observ.)

(1968,

( 1970)

1970)

( 1968, 1970)) Teleki

(1970), Teleki

580

G. TELEKI

ants with sticks;

poking at unfamiliar

‘olfactory

digging into the terrestrial

aids’;

items, including

foods, with sticks or grasses used as

nests of safari ants with sticks;

from hollows in trees, and brain tissue from a skull cavity, by chewing;

and z.@ing up ants with leaves crumpled

An interesting observed ation

footnote

to use objects

have been recorded

technical

skills and predatory

among baboons;

habits

technology

the actions

skills.

then

in the application

absent

the opening

which

is somewhat

of technological

performed

comprise

as variations

chimpanzee

probing,

analogous

and human

chimpanzee

and

technique

subsistence

If prying,

themes of digging,

It is noteworthy,

limitation

poking

probing

as distinct

and

technical

among human gatherer-hunters,

but it occurs rarely,

of some gatherer-hunter

of objects

involve several types of foods in variety.

probing

having

been seen only

however,

that an activity

for insects-is

widely

But the tubers and other subsurface

Herein lies another

of modification,

to

skills to subsistence

a rift between

digging and sponging remain

to digging-namely,

by chimpanzees.

apes.

The manipulation by actions

of technical

limited

on the respective

of ant nests with sticks.

the staple resources

these omnivorous

of a link between

is on the one hand similar

behavior

are themselves

from this repertoire;

performed

183 cases of pred-

devices.

only pounding,

during

frequently

The absence

chimpanzees

Digging with sticks, which is a standard

is not entirely

in press).

by virtue of the fact that the latter hunt almost exclusively

the above examples

and wiping are regarded sponging,

among

observable

with the aid of technological and objects,

1960 (Teleki,

have not yet been

prey, even though

but on the other it serves to emphasize

subsistence

Although

or dismembering

since

other kinds of limitations human

by hand.

to the above list is the fact that chimpanzees for killing

sponging water

with wads of leaves crumpled

and

flora which

populations

cannot be exploited

highlighting

the difference

by

between

technology.

in contexts

other than subsistence

so it is doubly

significant

that

is rarely

many

accompanied

of the food-getting

technical skills exhibited by chimpanzees include a component ofpreparatory modification. The form of an object is most commonly modified when materials are prepared for probing and sponging:

twigs and vines are stripped

vines and grasses are broken

or bitten

of leaves and other projections;

off to suitable

lengths;

sticks, twigs,

and leaves are chewed

or

manually crumpled into absorbent masses. These observations suggest that there may be, and may well have been, a stronger linkage between modification and subsistence activities

than between

modification

and other patterns

of technological

behavior,

such

as aggressive use of objects. Chimpanzees sometimes carry objects about for periods of more than an hour and for Such transport behavior occurs mainly in connection distances greater than a kilometer. with

termite

mound

collecting,

as an individual

when

an appropriate

seeks new tunnels.

probe

Materials

may

be carried

from

mound

to

such as vines and grass stems are

provides usually carried in the hands or between the lips. The context of insect-collecting the only definite cases where transportation is an intermediate activity between a series of applications. But there are other situations in which transportation is an anticipatory activity, as when a chimpanzee sets off to probe for termites and then selects and prepares an appropriate vine or grass stalk some 100 or more meters before arriving at a mound. And in still other situations chimpanzees will leave one place to fetch an object from another : e.g. when getting a new probe during a ‘fishing’ session, or when getting an The apparent absence of transport behavior object to strike or throw at an antagonist. in baboons, the conditional presence of rudimentary transport behavior in chimpanzees,

CHIMPANZEE

and the routine of ordered

practice

Surveys have

of long-distance

levels of complexity of Primate

used in agonistic been

stimulus,

behavior,

as proposed by Kortlandt

of objects

probably

Similarly, Primate

whereas

the emergence

& Kooij

in humans

are most commonly

behavioral

context

for developing

was initially

(1963))

the modification

with subsistence

skills probably

of pongids.

appeared

and transportation

with

Whereas

in the very early

in technology,

An evolutionary feedback

much

perhaps

model of technological

system wherein

applied to weapon gathering

later,

with the emergence

development

modification

stages

of

until

such as a shift from

of the earliest

flora, may hominids.

would, in this case, have involved

and probably

long-distance

making long after these activities

subsistence

sticks

for use as probes.

skills may not have appeared

More subtle refinements

skills.

agonistic

and transportation

activities.

the use of sticks to expose insect nests to the use of sticks to dig for subsurface only have appeared

and widely

may, in effect, technical

associated

for use as clubs, twigs must usually be modified

modification

are suggestive

technology.

source,

of objects

in conjunction

manipulative

evolution,

This

or original

even if the manipulation

need not be modified

behavior

subsistence

1963).

581

TECHNOLOGY

skills show that objects

(e.g. Hall,

emerged

transport

in Primate

technical

situations

the Primary

However,

SUBSISTENCE

became

transportation

functional

a

were

in a collecting

or

and ecology have been conducted

in

context.

(b) Geografihical and individual variability The most intensive eastern other

Africa, parts

subsistence

of the technology

The repertoire area

studies of chimpanzee

but evidence

(including

continent.

The

Basin,

For example,

at the westernmost

has recently

geographical

Makolongo,

Kabogo

But information

been discovered

distribution

1, which supplements National

in

of chimpanzee

the data listed in Table

1,

Park and in the Kasakati-Mahali Point,

Mahali

from other regions,

Mtns)

is probably

and notably

from

portion of the Pan range, is likely to be rather incomplete.

most of the pounding

data from Cameroon

behavior

in Gombe

close to being fully documented. populations

known

is shown in Figure

of skills performed Kasakati

behavior

of technological

and Rio Muni,

data from Liberia

and Ivory Coast, and the probing

derive from the material

Figure 1. Geographical complexes.

distribution

remains of

of these activities.

chimpanzee

technological

582

G. TELEKI

Hence, an assessment of any geographical variability that may occur in technological traits rests on much firmer ground at the eastern than at the western extremes of the continent. Long-term studies in western and central Africa are sorely needed; until such have been conducted, the following regional comparisons remain highly tentative, being more an instructive exercise than a final statement of fact. The close similarities observed in the technical skills exhibited at adjacent sites, such as those in Tanzania or in Cameroon and Rio Muni, attests to the behavioral integrity of what probably were, until quite recently, larger population units. Based on this assumption, the two neighboring sites in Liberia and Ivory Coast (2 and 3 on map), the three intermediate sites in Cameroon and Rio Muni (4, 5 and 6), and the two adjacent sites in Tanzania (8 and 9) can be treated as three discrete units labeled as populations A, B and C, respectively. Table 1 showed that pounding is associated with hard fruits and nuts, probing with insects and honey, digging with ants, and sponging with water and brain tissue. These all appear to be highly functional relationships in which the actions performed and the materials utilized are closely tied to the types and locations of the resources being exploited. These functional relationships define technological complexes which consist of specific linkages in patterned actions, in materials and in foods (e.g. pounding on nuts with stones), and these complexes in turn combine to form the technological repertoire of a given population. The patterned actions may themselves be composed of behavioral sequences, such as scanning, locating, selecting, picking up, shaping and utilizing, all of which are preliminary to food consumption. Furthermore, each complex contains little or no variability from region to region-that is, the pattern of probing for termites is similarly performed wherever the complex appears-but the matrix of complexes which comprises a full repertoire may show considerable variability from one region to another. The full complement of technical skills-pounding, probing, digging and sponging-has not been documented at any single study site, nor in any one of the three labeled populations. Pounding is currently known only in population A. Despite 19 months of field surveying in the Cameroon area of population B, Struhsaker found no evidence of this trait (Struhsaker & Hunkeler, 1971); similarly, 20 months of chimpanzee surveys in the Rio Muni area of population B yielded no signs of pounding (Jones & Sabater Pi, 1971). In population C, hard-shelled fruits are opened simply by hitting the items themselves against hard surfaces, such as rocks or tree trunks (Izawa & Itani, 1966; van LawickGoodall, 1968). The pounding technique, in which one object is used to hit another, thus seems to be restricted to regions west of a zone that is approximately delineated by 5’E longitude. Probing for insects and honey has been extensively recorded in the regions inhabited by populations B and C, and this is the most widespread complex known today. Some intrapopulation variability may occur in this complex: both populations B and C probe for termites whereas only population C is known to probe for ants; and members of population B seem to favor sticks and twigs while members of population C seem to prefer grass stalks, vines and bark strips. In broad terms, the zone marked by the 5’E longitude line serves as the approximate western limit for the probing technique, as no evidence of this complex has been obtained in population A (Struhsaker & Hunkeler, 1971). Sponging up water and brain tissue, or wiping up ants, is known at present only in population C. Although the sponging complex may be a regional variation on the same scale as the pounding complex, there is some possibility that this technical skill has not

CHIMPANZEE

SUBSISTENCE

TECHNOLOGY

583

yet been observed outside Tanzania simply because of differences in the scope of field studies conducted elsewhere. The distribution of digging, which has also been observed only in Tanzania, may be similarly biased. However, when the facts are taken as they now stand, pounding is restricted to population A, sponging and digging to population C, and probing to populations B and C. The longitudinal lines of 5”E and 25”E serve as rough boundaries for the technological repertoires. It is perhaps noteworthy that the 29”E meridian, which separates the main basins of central and western Africa from thle rift zone of eastern Africa, has been cited as a rough boundary for baboon and chimpanzese predation on mammals, a phenomenon which occurs at several sites east of this lin’e (Kortlandt & Kooij, 1963; Teleki, in press). Not even the most widely exhibited technical skill, probing for insects, has been recorded at some sites lying well within the range of this complex. In Uganda, for example:, where investigators have intermittently studied chimpanzees since 1962 (Reynolds & Reynolds, 1965 ; Sugiyama, 1968, 1969), all forms of subsistence technology seem to be absent. Such negative evidence is at best dubiously reliable, but the apparent absence of certain complexes in local populations does introduce the issue of technological variability within a regional population. There is some evidence to indicate that the patterns of technological variation outlined above are less a function of opportunistic than of behavioral variability. Although termites are known to occur from the westernmost to the easternmost chimpanzee habitats where some sort of subsistence technology is practiced, these insects are not everywhere exploited via the probing technique. Similarly, the nuts of oil palm trees are apparently consumed by chimpanzees throughout Africa, yet the pounding technique has been observed only in the westernmost region. These same nuts are a favored food of population C as well, but the latter simply strip away the external pulp with their teeth and sometimes swallow the nuts whole. If further field studies show that technological complexes do vary within or between populations despite uniformly available food resources, an element of cultural heritage may play a part in the transmission of skills. The acquistion and propagation of a technical skill, such as probing for termites, is known to involve learning processes which are in turn probably affected by individua.1 differences in ability. Chimpanzee infants learn their ‘fishing’ skills by watching older individuals and then practicing the technique for several years (van Lawick-Goodall, 1968, 1970), and the level of sophistication achieved by the time adulthood is reached is noIt necessarily uniform in all members of the community, or social unit. These individual differences may lead to considerable variability in the performance of any given technical skill. Thus, the transmission of such a tradition might be dependent upon both the sophistication of the performants and the aptitude of the learners. Such individual variability is in fact observable in many contexts other than that of subsistence technology. Some members of a chimpanzee community participate regularly in predatory episodes whereas others may avoid these situations entirely (Teleki, 19733) ; some members of a group may consistently use objects in attacking a mechanized leopard whereas others exhibit no such technical skills (Kortlandt, pers. comm.). The examples are many and diverse, individual variability being a pervasive theme in most ofthe activities performed by chimpanzees wherever they have been studied in depth. Technological behavior may, therefore, be but one of many activities that are differentially performed within a single population and perhaps even within a single social unit.

584

G. TELEKI

The mechanisms whereby chimpanzee technological repertoires differentiate through time remain obscure largely because the data are still so incomplete. Models of behavioral variability in subsistence habits must therefore be sought elsewhere. Intensive studies of certain Japanese macaque populations, in which kin, consort and other social ties regulate transmission of such spontaneously acquired subsistence techniques as potatowashing and wheat-sifting (e.g. Azuma, 1973), provide one potential model. The similar social mechanisms which have been tentatively identified as regulators of chimpanzee predatory behavior (Teleki, 19736) constitute another potential model. In both of these situations, not all members of a social unit learn and apply the various subsistence techniques uniformly and unilaterally. Depending on the extent of inter-unit migration, such variability could enhance or inhibit transmission of a given behavior within and/or between populations. Most Primates can be classed as omnivores, but the degree to which any one population or social unit is omnivorous can vary tremendously. Baboon and chimpanzee populations, for example, exhibit great variability in predatory frequency, perhaps because this behavior is affected by both social and environmental contingencies (Teleki, in press). So the mechanisms regulating a subsistence activity, such as predatory behavior, and any other activity associated with subsistence, such as technological behavior, may set the stage for differential development of skills. Physical and/or psychological limitations on acquiring and transmitting technical skills may, on the other hand, offset this variability in such a way that an upper limit to technological achievement is met at some point. The balance between exploitative skills and available resources would, in effect, tend to remain at equilibrium at a given place and over a short period of time. Consequently, the Gilgil baboons (e.g. Harding, 1973) can no more be expected to exhibit the kinds of technical skills (and predatory habits) possessed by the Gombe chimpanzees (e.g. Teleki, 1973c) than can the latter be expected to exhibit the kinds of technical skills (and hunting habits) possessed by the Kalahari bushmen (e.g. Silberbauer, 1972)) even though all three populations are fully omnivorous. 4. Chimpanzee Inquiry

Termiting

Skills:

A Personal

The techniques used to probe into termite mounds and ant nests have been closely studied at the Gombe and Mahali sites (van Lawick-Goodall, 1968, 1970; Nishida, 1973). My purpose here is to supplement these descriptions with comments on the expertise needed by chimpanzees to perform this task successfully. During the course of two termite-collecting seasons, in 1968 and 1970, I followed Gombe chimpanzees on their rounds of termite mounds with the intention of learning about the following problems. (1) How does a chimpanzee find the entrances to termite tunnels which, during the diurnal foraging period of these apes, are covered by an homogeneous layer of clay? (2) How does a chimpanzee select the object to be used in probing for termites, and what are the essential properties which enable the selected object to function suitably ? And (3) what actions must be executed in order to ensure a rewarding catch of termites ? Early in my observation I had become impressed by the ease and facility with which adult chimpanzees performed the probing technique, and my queries were prompted by a suspicion that the existing descriptions of this technological complex, which seemed to focus mainly on the mechanical actions involved, did

Plate 1. Adult male chimpanzee (Faben) carefully draws a length of grass from a termite tunnel with his left hand and begins to extend his lower lip in preparation to pulling the soldier termites off the ‘fishing’ tool.

584

Plate Z.(a) Adult male chimpanzee (Leakey, on left) holds a probing tool between his lips while scraping the mound surface with a forefinger in order to expose a tunnel mouth. His estrous female (Fifi) consort watches intently. (b) Adult male chimpanzee (Figan) strips soldier termites from a grass probing tool, using his lips and teeth to remove the termites while stabilizing the tool on his wrist.

Plate 3.(a) Infant female chimpanzee (Porn) attempts to fish for termites at a tunnel just abandoned by her mother; however, the tool she uses is inappropriately stiff and thick and will not enter the tunnel (note that the grass stem is beginning to buckle). (b) Adult male chimpanzee (Figan) walks between two termite mounds, in this case more than a hundred meters apart, carrying his probing stick in his mouth (note that stick is frayed at the end, where it was broken to a suitable length and that it has been stripped of leaves).

Plate 4.(a) Juvenile male chimpanzee (Sniff) twirls a branch about with both feet in solitary play, also exhibiting a moderately intense play face. (b) Adult female chimpanzee (Flo, upper left) intently grooms a handful of leaves while two of her offspring (Flint, left foreground; Fifi, upper right) watch the lip and finger movements closely. Such ‘objectgrooming’ often serves to elicit social grooming from those who watched the performer.

CHIMPANZEE

not adequately

convey what chimpanzees

In particular,

insects.

I wished

requisites involved. One old and seemingly during

the second habits,

important

which

with regular

routinely

executed

to imitate

of Leakey,

the physical

activity,

which insight eventually pongid technical

pre-

a study of ranging

habits,

it seemed evident by then that patterned visits to certain

mounds,

efficiency

the techniques

might

under observation, experience

the ‘uniqueness’

the final

revealed

new

in this subsistence

led to the suspicion that anthropologists

skills when defining

while alone

including

involved

an

More-

in the skills so

below, these experiences factors

play

and competence.

(All aspects of the procedure, As described

and psychological

estimate

the psychological

was followed most consistently

so as to gain personal

by this individual.

about

about

Leakey,

of an individual’s

tasting of the insects, were practised.) information

in. order to ‘fish’ skillfully for these

time I was conducting

versus random

endeavored

or when in the company

585

TECHNOLOGY

something

structure-because

part in the development

over, I frequently

must k&g

to discover

highly skilled individual,

season-at

daily regimen and community ranging

SUBSISTENCE

tend to under-

of man.

(a) Setting the stage As reported

by van LawickGoodall(1968,

the following tunnel,

basic actions:

using a finger

scraping

1970), termite-probing,

or thumb;

mouth; pausing momentarily while holding the protruding extracting the object with its attached soldier and worker desired termites

(soldiers being preferred)

in one hand and stabilizing Gombe

study population

with the lips and teeth while holding the probe

activity

Termite-probing onset of rainfall reproductive the rains

is an annual

2 months

per year,

in October

when termites

and migratory

begin.

There

activity, period

evidence

cues, as they sometimes

rains.

Out-of-season

termiting

rarely

practiced

during

regularly

at Gombe

these mounds,

presumably

in other circumstances (b) Locating

tend

of all but

to show the

as the ‘fishing’ season only being

occurring

chimpanzees

closely linked

begin to examine

mounds

these

to their mounds. because

it contains

(except

but the probing

technique

It is also relevant

during the migration

Chimpanzees

sometimes

salt, a mineral

seasons,

prior to the first

and opened even during the most intense collection nocturnal

to the

the termite

about 2-3 weeks after

anticipate

of the dry or rainy seasons.

are essentially

block all entrances

that

Individuals

adults

shift also regulates

bouts do occur occasionally,

the height

tunnels must daily be relocated for termites

This climatic

cycles, with mass migration

perhaps via climatic

4-5 hr

are in season.

but not an everyday

is some

of the

hr and sometimes

skills, although

the most productive

or November.

Nearly all members

devote at least l-2

the youngest age classes exhibit the probing greatest persistence and proficiency. lasts about

end of the object in one hand; termites; and picking off the

the free end on the other wrist.

(ca. 50 individuals)

daily to this one subsistence

or ‘fishing’, consists of

away a thin soil layer that covers the entrance of a carefully poking an object into the exposed tunnel

which

is

that

period,

time)

and

suck on soil from they actively

seek

as well.

tunnels

Having repeatedly observed individuals approach a mound, make a rapid visual scan of the surface while standing on or beside it, and reach decisively out-with a high degree of predictive accuracy-to uncover a tunnel, I was soon impressed by the apparent

586

0.

TELEKI

ease with which tunnels could be located. In attempting to learn the technique, I applied several experimental procedures : examining in minute detail all crack patterns, protuberances, depressions and other “topographic” features in the clay. But, after weeks of futile searching for the essential clue, I had to resort to scraping mound surfaces with a jackknife until a tunnel was inadvertently exposed. My inability to find any physical features which could serve as visual cues eventually led me to realize that chimpanzees may possess knowledge far beyond my expectations. Consistent following of chimpanzees during the two termiting seasons, and especially among the mounds scattered throughout lower Kakombe Valley (see Teleki, 19736, for maps of the study area), revealed that some individuals routinely visit certain sets of mounds, often travelling along a prescribed network of trails from one to another in the sequence. Thus, each chimpanzee seemed to exploit a core set of mounds most heavily, although other mounds encountered during more extensive forays are of course utilized on an opportunistic basis. Presumably the mounds in the core set, which may vary in number from about 10 to 25 per individual, are more familiar to a given individual than are the mounds less regularly visited. Some or all of the core mounds may be visited on a daily basis during the collecting season, but the regularity of visits may be influenced by other behavioral opportunities (e.g. an estrous female) which may be present at the time. Many of the mounds within a core set may be shared by several individuals, so the ‘termiting ranges’ may overlap extensively. Because only 1-3 chimpanzees are likely to work a mound simultaneously, competition occurs only rarely. There is some indication, however, that adult females, and particularly mothers, who tend to have much smaller ranges than other adult classes, exhibit an unusual degree of competition for this resource (Baldwin, pers. comm.). A chimpanzee’s presumed familiarity with a core set of mounds provides the main clue to the problem of locating tunnels. The only hypothesis which, at this point, seems to reasonably account for the observed facts is that an adult chimpanzee may know (memorize?) the precise location of 100 or more tunnels in the most familiar mounds. Moreover, since intensive probing is restricted to a short annual season, the possibility that chimpanzees retain a mental map of core mound features during the intervening 10 months must also be considered. That chimpanzees require a prolonged learning period (i.e. 4-5 years) to gain proficiency in this technique (van Lawick-Goodall, 1970), and that some individuals are known to have the capability to retain specific information for many years, provides circumstantial support for this hypothesis. However, only a detailed study of termite-collecting habits, and of termite behavior, is likely to resolve the following kinds of questions. Is the error rate in locating tunnels higher at mounds that are less familiar than those in a core set ? Are the tunnels used by a given individual in a core set of mounds consistently the same for long periods of time? Is there a period of greater error in locating tunnels at the start of each season? (c) Selecting materials

An examination of the skills needed for selecting an object having the requisite properties for successful probing was the next step in the investigation. When performed by experienced chimpanzees, the selection procedure seems deceptively simple. After a brief visual scan of the nearby vegetation, a chimpanzee will usually extend a hand and defty tear off a twig, vine or grass stalk. Sometimes the individual must move a few paces away

CHIMPANZEE

from the mound selected.

These

and fetch a suitable may be rapidly

SUBSISTENCE

probe,

examined

in one, or several may be carried

587

TECHNOLOGY

and in some cases 2-3

and discarded

to the mound

objects

are initially

until some specification

for subsequent

selection.

is met

Whenever

it

occurs, the selection is made in a swift, almost casual manner, and modification is begun if necessary. Without being aware of the nuances involved, it is easy to undervalue the proficiency needed to perform these manouvers. Chimpanzees presumably have the experience can be evaluated

probes is not high.

For the novice human

sequence

holds the key to selection

apparent

only through

observer,

because

the trial-and-error

dimensions, or biting

before

motor patterns

are other physical

which

of an object

the utilization

phase of the

of an object

It is not difficult

becomes to find a

nor is it difficult to trim away leaves and

it down to the required

which entail some dexterity

properties

less easy to anticipate. stringent:

breaking

however,

of application.

other

projections

the properties

for the rate of error in selecting

the appropriateness

grass stalk or vine stem of suitable routine

whereby

before it is applied to the task of probing,

must precisely

When probing for termites,

length.

These

but little prior experience. fit the intended

the specifications

are

But there

task, and these are

are in fact surprisingly

if the vine or grass selected is too pliant, it will buckIe and collapse (accordion-, if, on the other hand, the object is too stiff or

like) when inserted into a twisting tunnel; brittle,

it will catch on the tunnel walls and either break or resist entry to the necessary

depth.

An intermediate

range

yield many termites. Despite months of observing enviable Similar

of qualities

and aping adult chimpanzees

ease, speed and accuracy, ineptness

My experiences locating, retention

suggest that selection proficiency

entail a considerable of the results.

ants from terrestrial of the

I was unable

to achieve

can only be observed in chimpanzees

individual. Behind these refinements nature

must therefore

The

amount

span

resources

being

exploited.

as they selected probes with1 their level of competence,.

may

behavior

actually

collecting foremost,

ants are large,

up with a stick.

stiff sticks or branches

into the nest without

bending

by long-term

the lifetime

used in collecting

of an

termites

and

in their properties, due in large part to the At Gombe, termite mounds (Macroterme:r

ant nests (Anomma nigricans) consist of loosely aggregated away by hand or stirred

backed

cover

belbicosus) are covered by a uniform clay Iayer which is extremely pushed

is to1

is a learned skill which must, like tunnel-

is the fact that the materials

nests differ considerably

if probing

below the age of about 4-5 years.

of trial-and-error

retention

be selected

The which

or breaking.

hard when dry, whereas

soil particles objects

which can easily be

commonly

can be forcefully

The dimensions

selected shoved,

and properties

for end of

these objects are thus quite different from materials used in probing, where the combined criteria of pliability and tensile strength are more important. To complicate the issue further,

the techniques

and materials

used to collect ants from arboreal

nests (Crematogaste:r

spp. and Camponotus spp.), as described by van Lawick-Goodall (1968) and Nishida (1973)) resemble those used in collecting termites from terrestrial mounds. Exploitation of each food resource

therefore

entails selection

procedures

geared to a specific task and resource.

Given the possibility that technological repertoires differ from region to region, each stage of a particular complex may well have to be learned via a cultural tradition that is passed from individual to individual within a given population, one of the most direct and stable routes offspring.

of learning

being

the prolonged

associations

between

mothers

and their

588

G.

TELEKI

(d) Utilizing probes Having

failed to meet chimpanzee

of termite-probing, stage,

which

probes. weeks

seemed

relatively

Consequently,

interval, particular

of aptitude

set out to acquire

simple

them

total

out again-without

failure-an

panthropologist-did tunnel

to locating probes,

getting

uncommonly

learning

(in which

termites

and selecting

for the designated

any termites.

brief

stages

in the ‘fishing’

tunnels

pausing

Only

span,

I finally begin to grasp the problems

a very rich tunnel may initially produce moderate a newly opened

and skill in the preceding

some competence

in comparison

I spent many hours inserting

and pulling of nearly

standards

I nonetheless

after some

at least

involved.

for this Although

rewards even if skill is essentially lacking,

are perhaps

still too far underground)

or a

partly ‘fished-out’ the application ful probing

tunnel (which termites may block off at some subsurface level) requires Skill is particularly necessary if the period of successof subtle techniques.

is to last more than a few minutes.

In order to collect fully and dextrously

these subterranian inserted

wrist so that the object

termites,

the probing

to a depth of about 8-16

navigates

the twisting

object

must first be care-

cm, with appropriate

channel.

The probe

turns of the

must then be gently

vibrated with the fingers during the prescribed pause, for without this movement the termites may not be stimulated into biting firmly onto the probe. However, if the vibration is performed

too lengthily

be cut through have

been

attached,

or roughly,

by the mandibles

correctly

performed,

must be extracted

If the object

is too rapidly

there is an excellent

while still in the tunnel. the probe,

chance

presumably

with

dozens

or clumsily

which then yields nothing

but not overly swift and, once started,

graceful.

If the tunnel is particularly of the probe),

of termites

now

Once again there are nuances to be observed. pulled out, the insects are likely to be scraped off

motions insertion

will

actions

from the tunnel.

along the sides of the tunnel, must be reasonably

that the probe

When these preliminary

tortuous

but a shredded

probe.

uniformly

The hand fluid and

(a feature which can be determined

during

the success of the catch can be ensured by a slow twisting

of the

wrist while the probe is pulled out. (e) Considering the results These,

then,

are the sequential

behavioral

components

of a chimpanzee

technical

skill

which, when viewed superficially, seems to be a far simpler task than those routinely performed by human gatherer-hunters. But my participant observation at Gombe unveiled

the remarkable amount of experience and finesse needed by a chimpanzee to perform this subsistence activity. When such qualitative aspects of subsistence technology are omitted from descriptions, the risk of dispensing with such skills as ‘rudimentary’ increases. Incompetent as they were, my attempts to acquire the skills of locating termites left me with a tunnels, selecting materials, inserting probes, and extracting healthy measure of respect for chimpanzee technical ability, as well as with a nagging suspicion that the physical and psychological capabilities needed to develop, apply and transmit such skills may differ in degree but not in kind from those needed by humans to locate, expose and gather insects and subsurface flora. In other words, a question about how an integrated sequence of technological behavior, where the omission of any single stage results in failure, was initiated, put together and finally fixed into the repertoire of a population, can now be directed at nonhuman as well as human Primates.

CHIMPANZEE

If these tentative gap postulated

conclusions

are confirmed

to exist between

though not entirely

SUBSISTENCE

by further field research,

the nonhuman

and human

The key word here is ‘partly’,

filled.

589

TECHNOLOGY

the technological

Primates

would be partly

for it remains

clear that the full

scope of subsistence technology probably differs considerably, in terms of both the range human gatherer-hunters and the refinement of skills, from one Primate genus to another: are unquestionably

more

than are nonhuman

dependent

on a wider array

5. Termiting The simplest way to illustrate data

on the termiting

Africa.

This

sample

exhibited

of Primate

Interspecific

skills and materials

phylogenetic

some comparative:

chimpanzees

and humans

from a cercopithecid

to mirror the increasingly

different

Comparisons

in degree is by reviewing by baboons,

populations-ranging

as well as the levels at which limitations having

Skills :

such differences

techniques

pongid to a hominid-serves primates

of technical

collector-predators.

compIex technological

arise, that can be observed

in.

through

a.

capabilities,.

in the behavior

of

histories.

(u) Baboon and chimpanzee techniques General

differences

already

in the technological

been summarized.

termite-collecting National

Park,

skills of these (1968),

baboons,

habits

The

baboons

simple

and flying insects are collected

be seen leaping methods

(These

colobus

as they emerge,

do two or more

object

Several

collect

the apes collect

as a probe,

period,

baboons

insects.

mouth.

especially

But observers

also collect

instead probe for soldiers elsewhere chimpanzee

technique.

limited

to the 2-3

simultaneously, are sometimes

by adult males, and may then using an

on a mound or abandoned termites

the insects massing

on the same mound.

As a result, the termite-collecting

weeks before and after the short migration

Baboons

have not seen baboons

some winged

in

such as the ‘leaf-eating’

may work a mound

at a single tunnel

yet they will in many cases ignore

additional

of

mass and fly away from open

Primates,

even when these have been discarded

tunnel by the chimpanzees. The Gombe chimpanzees

and

to focus on modes

up and down on mounds in an effort to grab termites

displaced from mounds by arriving chimpanzees, sit and watch

of Gombe (1971)

with hands and/or lips, and baboons

are also used by other Gombe

and the blue monkey.)

but rarely

opportunity

by Ransom

season is almost exclusively

singly or in clusters,

can sometimes

and chimpanzees

have: in the

Thus, their bulk intake consists of winged termites

occur.

tunnels. flight.

differences

in a single habitat.

that are collected, Crawling

and chimpanzees specific

have been described

a uniquely

the termiting

weeks during which migrations

of baboons

here is to highlight

Primates.

provide

a single food resource

For the Gombe

behavior

purpose

whose insect-eating

Lawick-Goodall exploiting

The

during at tunnel

in a

the migration entrances

and

Probing is clearly the standard season is extended by several

period,

and the bulk intake consists

of soldier termites. The technical exploitation of this food resource, differ markedly

skills developed by chimpanzees have clearly affected and the exploitative potentials of the two genera now in duration if notin bulk. And, although both primates show a great fancy

for termites, their respective limited-supply resource.

preferences

and methods

largely

obviate

competition

for a

590

G.

TELEKI

The value of the Gombe example lies in its exposure of what appear to be fixed patterns of behavior in baboons and to some extent in chimpanzees. What, in other words, inhibits baboons from adopting, at least to a limited degree, the techniques used by chimpanzees ? And what kinds of psychological phenomena are involved in this limitation? The questions become doubly intriguing when set against a larger regional background, for this apparent limitation is not necessarily characteristic of the genus Pa&o. According to the incidental reports collected by Kortlandt & Kooij (1963), some baboons do use pieces of wood to widen the entrances of termite nests. Whether this is a regional difference in behavior comparable to those documented for chimpanzee populations A, B and C remains unknown, as does the problem of whether the more technically skilled baboons exhibit similar levels of sophistication and persistence as do chimpanzees. Further field studies are much needed to elucidate such problems. Given the general differences between baboon and chimpanzee technological repertoires, as well as the specific discrepancies observed in termite-collecting at Gombe, it is currently tempting to postulate that the technical skills in these genera are related to varying levels of cortical complexity, and hence to differential learning and performing capacities. If so, a model of behavioral evolution might be integrated with the numerous existing models of biological evolution. But there are crucial, as yet unresolved issues: for example, to what extent are such technological limitations characteristic of species or genera, and how are the limitations adaptively significant? (b) Chimpanzee and human techniques

In a survey of human insect-eating habits, Bodenheimer (195 1) shows that termites, ants and other insects are highly favored by numerous human populations, many of which live in regions also inhabited by baboons and/or chimpanzees. Taking Africa in its entirety, human techniques of termite-gathering are extremely variable and often much more efficient than those of termite-collecting baboons and chimpanzees; on a local level, however, some of the human techniques are the same as those used by nonhuman Primates. In western Zaire, for example, members of the Baluba tribe insert palm leaves into termite tunnels, pause to stir the protruding probes, and finally withdraw the probes laden with clinging soldier termites. Elsewhere in western Zaire, large poles are driven into termite mounds to make holes about a meter in depth; the poles are withdrawn, and small “brooms” made of grass are inserted into these holes ; a noise is then made with the tongue so as to imitate the sound of raindrops falling on the clay, and the probes are turned slightly by hand so that termites will clamp onto the grass; then the probes are withdrawn, and the soldier termites are eaten one-by-one immediately, or they may be collected in baskets and taken away. A variation on the theme of imitating raindrops occurs in northern Zaire, where two pieces of wood are beaten against the mound surface. A rather common form of suberfuge, stick-beating has also been observed at Mt Elgon, on the UgandaKenya border, and other areas. The soil of termite mounds is in some places also eaten, the claim being that such soil contains higher concentrations of salt. At Gombe, where termites are also actively sought by the indigenous human population, individuals or small groups frequently go termite-gathering during the rainy season. The insects are often gathered at dusk, the following technique being commonly used: a kerosene lantern is placed on or near the mound, both to aid vision and to act as

CHIMPANZEE

SUBSISTENCE

TECHNOLOGY

591

a beacon for the insects; a large portion of the mound surface is scraped away with a panga (machette) blade to expose and stir up the insects; the exposed milling mass of termites is picked or scooped up into a dish or basket by hand and sometimes by leaves or fronds. During the 2-3 week migration season mainly winged termites are gathered, but both soldiers and workers are taken during the rest of the rainy season. On one occasion I observed a man rapping on a mound with two small sticks, presumably in order to imitate raindrops. Thus, three ways of exploiting a given resource, two of which involve technological behavior, can be compared at this one site. The human technique further extends the duration of the termite-gathering season, much beyond the periods in which baboons and even chimpanzees can collect these insects, and perhaps increases the annual yield as well. The baboon, chimpanzee and human populations in Gombe National Park clearly exhibit a series of increasingly sophisticated technical skills which reflect distinct, perhaps species-specific, levels of technological (and psychological?) capabilities. After emphasizing the remarkable degree of similarity which occurs in some chimpanzee and human termiting techniques, it should also be stressed that the above comparisons are not fully representative of the rich array of skills performed by various human populations. Whereas chimpanzees throughout much of Africa possess a single technological complex for termite-collecting, human populations exhibit extensive regional variability in techniques which can include fumigating, digging, pouring water onto mounds, placing upturned dishes onto sticks driven vertically into the clay, and so on. Moreover, only humans are known to practice the subtle subterfuge of imitating raindrops so as to entice termites to the surface. This one skill may alone extend the gathering season by weeks or months. These anecdotal cases do not provide a comprehensive picture of Primate technical skills. Numerous gaps must be filled in with further comparative field work. But the examples do suggest that subsistence activities, and to some extent even technical skills, can be arranged as overlapping segments on a behavioral continuum. When such continuities can be documented among extant cercopithecid, pongid and hominid populations, the resultant behavioral models can be used to supplement morphological trends in primate evolution and also to lay a foundation for reconstructing the probable subsistence habits of the earliest hominids. Although attention has focused here on technological behavior, this approach can be adopted with other subsistence activities, such as predatory and hunting behavior (Teleki, in press). 6. Conclusion

and Discussion

The application of technical skills to subsistence activities, and particularly to hunting game, has been widely cited as a characteristically human condition. The material remains of technological behavior, such as shaped artefacts, are often regarded by prehistorians as a sign of human cuhure, and have thus been ranked as a prime ingredient in the process of hominization (e.g. Oakley, 1959). This conceptual trend was readily adopted in other branches of the discipline, and notably by those cultural anthropologists who were interested in separating nonhuman primate society and primitive human culture (e.g. Sahlins, 1959). Now deeply ingrained in our minds and our texts, the trend continues to be propagated in these circles (e.g. Fried, 1967) despite the many advances which have begun to sway members of other subdisciplines toward an alternative position (e.g. Lancaster, 1968; Mann, 1972). Needless to say, the impact of seperatism upon

592

G. TELEKI

anthropological theories of human evolution has been immense, for the above is by no means an isolated case history. Many traits other than technology-among which can be numbered predatory behavior, nuclear family structure, division of labor by sex, kinship relationships and lineage ties, incest prohibitions, sharing and cooperation-have been routinely linked to the divergence of a savanna-dwelling, meat-hunting hominid from a forest-dwelling, plant-collecting Primate heritage. That this determination to regard humans as behaviorally and psychologically supreme (an attitude which retains a strong flavor of “special creation”) contrasts with a readiness to accept biological affinities (an attitude which often leads to scientific destruction in the name of human welfare) does not seem to have caused excessive consternation among students of human behavior and evolution. Yet it steadily becomes more obvious, as evidence accumulates about what Primates actually do in their native habitats, that nearly all of these ‘human’ traits can be identified among the nonhuman Primates. Chimpanzees are in fact rapidly becoming the prime exception to the seperatist perspective, as Pan is already known to exhibit more such traits than any extant genus other than Homo itself. Chimpanzees are, for example, known to cooperate in prey pursuit, to practice division of labor in pursuing and capturing prey, to share meat along lines reflecting kinship, consort and other social ties-all within the single context of predation (Teleki, 1973b,c). And, as shown in this report, chimpanzees are also technically skilled Primates which, as Lancaster (1968) has aptly pointed out, makes this living pongid the best candidate for mirroring the ancestor postulated for the origin of the hominid line. Similar advances continue to occur along many avenues of investigation, some of the most outstanding being family structure (van LawickGoodall, 1968, in prep.), community organization (e.g. Itani & Suzuki, 1967) and communication (e.g. Gardner & Gardner, 1969; Rumbaugh, Gill & von Glaserfeld, 1973). Once Homo erectus sites have been passed, in our search backward through time, the archeological record of artefacts becomes skimpy indeed. And, in the few cases where claims for such evidence are made, debates about authenticity continue for decades. But the evidence for technological behavior among living Primates suggests that this is one of many attributes which can be traced far into Primate prehistory. The ability to manipulate objects for specific purposes is probably one of the more ancient and pervasive trends in Primate evolution, having emerged perhaps as early as the Oligocene. Rudimentary kinds of modification and transportation, such as one sees today among baboon and chimpanzee collector-predators, may already have played a part in subsistence activities during the Miocene, although more sophisticated forms of such behavior, as performed by some contemporary human gatherer-hunters, may not have appeared until the Pliocene. Most of the materials utilized by chimpanzees (and by other Primates) in their foodgetting activities are highly maleable and perishable. These materials are in fact often indistinguishable, sometimes even after being modified, from similar materials to be found in the vi&&y. Herein lie important implications for hypotheses which maintain that technological behavior initially arose among Primates when technical skills were applied to ‘hard’ materials that are prone to long-term preservation. Debates about the authenticity of the osteodontokeratic culture may continue for many years, but the likelihood that some extinct Primates once possessed technological capabilities-expressed for perhaps millions of years only in the manipulations, modification and transportation of ‘soft’ materials-is at least circumstantially strengthened by the chimpanzee evidence.

CHIMPANZEE

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Final, unequivocal proof for such speculation must, of course, come from the discoveries of paleoanthropologists. The approach which has been taken in this report-wherein acquisition and propagation of technological behavior is viewed as a gradual evolutionary process involving many Primate species-is offered for consideration largely because it appears to be theoretically more satisfying than those which link major evolutionary developments to some single factor (e.g. hominization to predation). Consequently, it is at least concordant with the incremental and cumulative processes of gradual change upon which biological theories of Primate evolution are founded. Hopefully my investigations have also shown that studies of nonhuman Primates can be relevant and instructive to those who seek knowledge about the emergence and evolution of Homo sapiens. I am deeply indebted to the National Geographic Society and the Gombe Stream Research Centre for their financial and logistical support of my field work, and to the Government and National Parks of Tanzania for the opportunity to live in one of their newest conservation areas. I am, in addition, more than grateful to the patient and. tolerant Leakey, whose termite-collecting skills so outstripped mine. References Albrecht, H. & Dunnett, S. C. (1971). Chimpanzeesin Western Africa. Munich: Piper. Azuma, S. (1973). Acquisition and propagation of new food habits in a troop of Japanese monkeys. In (C. R. Carpenter, Ed.), Behavioral Regulators of Behavior in Primates. Lewisburg, Pa.: Bucknell University Press. Progress report of the survey of chimpanzees in their natural habitat,, Azuma, S. & Toyoshima, A. (1961-2). Kabogo Point area, Tanganyika. Primates 3,61-70. Baldwin, L. A. & Teleki, G. (1972). Field research on baboons, drills and geladas: an historical, geo graphical and bibliographical listing. Primates 13,427-432. Baldwin, L. A. & Teleki, G. (1973). Field research on chimpanzees and gorillas: an historical, geographical and bibliographical listing. Primates 14, 315-330. Beatty, E. H. (1951). A note on the behaviour of the chimpanzee. Journal of Mammalogy 32, 118. Bodenheimer, B. S. (1951). Insects as IIuman Food. The Hague: Dr W. Junk. Eimerl, S. & DeVore, I. (1965). The Primates. New York: Time-Life Books. Fried, M. (1967). The Evolution of Political Society. New York: Random House. Gardner, B. T. & Gardner, R. A. (1969). Teaching sign language to a chimpanzee. Science 165,644-672. Goodall, J. ( 1962). Nest building in free-ranging chimpanzees. Annals of New York Academy of Science 102, 455-467. Goodall, J. (1964). Toolusing and aimed throwing in a community of free-living chimpanzees. Nature 201, 1264-1266. Current Anthropolou Hall, K. R. L. (1963). Tool-using performances as indicators of behavioral adaptability. 4,479-494. Hall, K. R. L. (1965). Animals that use tools. Animals 7, 16-2 1. Harding, R. S. 0. (1973). Predation by a troop of olive baboons (Papio anubis). American Journal of Physical Anthropologv 30, 587-592. Itani, J. & Susuki, A. (1967). The social unit of chimpanzees. Primates 8, 355-381. Izawa, K. & Itani, J. (1966). Chimpanzees in Kasakati Basin, Tanganyika: I. Ecological study in the rainy season, 1963-1964. Qoto University African Studies 1, 73-156. Jones, C. & Sabater Pi, J. (1969). Sticks used by chimpanzees in Rio Muni, West Africa. Nature 223, IOO101. Jones, Cl. & Sabater Pi, J. (1971). Comparative ecology of Gorilla gorilla (Savage and Wyman) and Pun troglodytes (Blumenbach) in Rio Mum, West Africa. Bibliographica Primatologica no. 13. Basel: S. Karger. Kohler, W. (1325). The Mentality of Apes. London: Routledge & Kegan Paul. Kortlandt, A. (1963). Bipedal armed fighting in chimpanzees. XVI International Congress of Zoology Washington, lecture text. Kortlandt, A. (1965). How do chimpanzees use weapons when fighting leopards? Yearbook of American Philosphical Society pp. 327-332.

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Kortlandt, A. (1967). Experimentation with chimpanzees in the wild. In (D. Starck, R. Schneider & H. J. Kuhn, Eds), Neue Ergebnisse der Primatologie. Stuttgart: Fischer. Kortlandt, A. (1968). Handgebrauch bei freilebenden Schimpansen. In (B. Rensch, Ed.), Handgebrauch und Verstiindigung bei Affen und Friihmenschen. Bern: Huber. Kortlandt, A. & Kooij, M. (1963). Protohominid behavior in primates (preliminary communication). Symposia of Zoological Society of London 10,61-88. J. B. On the of tool-using American Anthropologist 56-70. Mann, (1972). Hominid cultural origins. 7, 379-386. E. (1969). Soul of Ape. New Atheneum. Menzel, W. (1972). invention of in a of young Folia Prima17, 87-106. E. W. Further observations the use ladders in group of chimpanzees. Folia 19,450-457. Merfield, G. & H. (1956). Were My London: Longmans. T. (1973). ant-gathering behavior the use tools among chimpanzees of Mahali Mountains. of Human 2, 357-370. K. P. Man the (2nd edit.). University of Press. Pfeiffer, E. (1969). Emergence of New York: and Row. U. (1971). d’outils par chimpanzes de de la Terre et 25, 506-509. T. W. Ecology and behavior of (Pa&o anubis) Gombe National Unpublished Dissertation. of California, Reynolds, V. Reynolds, F. Chimpanzees of Budongo Forest. (I. DeVore, Primate Behavior: Studies of and Apes. York: Holt, and Winston. D. M., T. V. van Glasersfeld, C. (1973). and sentence by a (Pan). Science 731-733. Sahlins, D. (1959). social life monkeys, apes primitive man. (J. N. Ed.), The Wayne State Press. of Man’s for Culture. Savage, S. & J. (1843-4). of the characters and of the niger. Boston of Natural 4, 362-386. G. B. The G/wi In (M. Bicchieri, Ed.), and Gatherers New York: Rinehart and Struhsaker, T. & Hunkeler, (1971). Evidence tool-using by in the Coast. Folia 15, 2 19. Sugiyama, (1968). Social of chimpanzees the Budongo Uganda. Primates 225-258. Sugiyama, (1969). Social ofchimpanzees in the Forest, Uganda. 10,197-225. Suzuki, (1966). On insect-eating habits wild chimpanzees in the woodland of Tanzania. Primates Suzuki, A. An ecological of chimpanzees a savanna Primates 10, Teleki, G. Notes on interactions with carnivores in National Park, Primates 14,407-411. G. (19736). Predatory Behavior Wild Chimpanzees. Pa.: Bucknell Press. Teleki, (19736). The chimpanzee. Scien@ 228,32-42. Teleki, (1973d). On responses to accidental death a chimpanzee Gombe National Tanzania. Folia 20,81-94. Teleki, (in press). subsistence patterns: and gatherer-hunters. of Human van Lawick-Goodall, (1968). The of free-living in the Stream Reserve. Behavior Monographs 16 l-3 van Lawick-Goodall, (1970). Tool-using primates and vertebrates. In S. Lehrman, A. Hinde E. Shaw, Advancesin Study of London: Academic van Lawick-Goodall, van Lawick, & Packer, (1973). Tool-use free-living baboons Gombe National Tanzania. Nature behavior of and early In (S. Washburn, Washburn, S. & DeVore, (1961). S Ed.), Life of Man. Chicago: Zon, J. C. J. van & Orshoven, J. van (1967). Enkele resultaten van de Zesde Nederlandse chimpanseeexpeditie. Vakblad voor Biologien 47, 161-166.