The stone-tool

technology of
capuchin monkeys: possible
implications for the evolution of
symbolic communication in
hominids

Gregory Charles Westergaard

Abstract

Researchwithcaptivecapuchinmonkeyshasdemonstratedthatthese New Worldprimatesproduce

flakedstone artifactsthattheyuse as cuttingtools, andthattheyexhibitpatternsof right-handedness
analogousto those of tool-makingPlio-Pleistocenehominids. These findingsindicate that the
cognitiveand biomechanicalconditionsof pre-adaptationfor the productionand use of stone tools
are present in extant nonhumanprimates,and that capuchinscan be used to model processes
associatedwiththe evolutionof technologyandsymboliccommunicationin humans.

Keywords

Capuchin;Cebus;evolution;hominid;language;tool use.

Introduction

Hominids first produced stone tools up to 2.5 million years ago, during the Oldowan
technological stage (Schick and Toth 1993). Oldowan tool assemblages include flakes,
flaked cores and battered stones. Although stone-tool manufacture shares structural
features with the syntax of language (Holloway 1969), it is widely believed that symbolic
communication is not a prerequisite for the action sequencing of flint knapping (Davidson
1991; Wynn and McGrew 1989).
The cognitive skills which facilitate the production of stone tools have been examined by
Toth (1985a). Toth found that novice stone workers produce most of the forms in Leakey's
(1971) system of classification, including choppers, polyhedrons and discoids. He
concluded that early hominids could have produced similar forms as unintentional

World Archaeology Vol. 27 (1): 1-9 Symbolic Aspects of Early Technologies

? Routledge 1995 0043-8243
2 Gregory Charles Westergaard

by-products of flake production. In this view the earliest known flaked artifacts represent
technological 'paths of least resistance' rather than preconceived 'stylistic norms'.
Wynn and McGrew (1989) have compared the material culture of Oldowan hominids
with that of wild chimpanzees. The most striking difference between the two technologies
was that Oldowan hominids produced flaked stone tools whereas chimpanzees primarily
produce tools from vegetation (but see Boesch and Boesch [1990] for a description of
chimpanzees using unmodified stones to crack open nuts). The authors described several
similarities between the two material cultures, including modes of artifact production and
gradation of manufactured forms, and concluded that all behavior patterns which can be
inferred from Oldowan artifacts fall within the adaptive grade of modern great apes.
The production of stone tools has been examined experimentally in two great ape
species. In a pioneering experiment Wright (1972) taught an orangutan to produce stone
flakes which the animal used to cut through a cord that secured a food box. Toth et al.
(1993) demonstrated stone flaking to a pygmy chimpanzee who later learned to produce
his own flakes for cutting. These results indicate that nonhuman primates can be used to
model processes associated with hominid stone-tool technology.
Capuchins are arboreal New World monkeys that produce and use tools in several
contexts that have been hypothesized for Oldowan hominids (Homo habilis and perhaps
broadly contemporaneous Australopithecines; see Sussman 1994). Wild capuchins have
been observed to use a stick to club a snake and shells to crack open oysters (Boinski 1988;
Fernandes 1991). The tool-using and tool-making traditions of capuchins are much richer
in captivity, where these monkeys spontaneously (without training or human demon-
stration) use and modify sticks as probes, and stones and bone fragments as cutting and
nut-cracking implements (Westergaard and Fragaszy 1987; Westergaard and Suomi 1993,
1994a, 1994b). Similarities in tool using and tool making among capuchins, great apes and
early hominids are consistent with the idea that omnivorous, extractive foraging is
associated with complex sensorimotor propensities in primates (Parker and Gibson 1977;
Westergaard and Fragaszy 1987).
This report examines the stone-tool technology of capuchin monkeys and its possible
implications for the evolution of symbolic communication in hominids. It is organized into
three main sections. The first reviews methods of stone artifact production, the second the
use of stones as cutting tools and the third aspects of neurobehavioral laterality and their
relationship to the evolution of language. A summarizing conclusion follows.

The production of stone artifacts

Stone artifacts provide striking evidence of technological innovation which allows us to

infer cognitive functioning in the individuals that produced them (Wynn 1979). Schick and
Toth (1993) have identified four stone-flaking techniques that were available to Oldowan
hominids. The method used most widely, hard-hammer percussion, involves striking a
hand-held hammer-stone against a hand-held stone core. A second method, the bipolar
technique, consists of placing a core against a surface and then striking it with another
stone. When modern humans use this technique they tend to produce smaller flakes than
when they use hard-hammer percussion. A third method, the anvil technique, involves
 The stone-tool technology of capuchin monkeys 3

striking a stone against a stationary surface. This technique is simpler than the percussion
and bipolar techniques because it involves selection and control of only one stone. The
fourth method, the throwing technique, consists of projecting a stone against a hard
surface such as a large rock. Throwing can split a stone into two or more pieces with sharp
edges.
Capuchins produce stone flakes by striking together stones held in each hand
(hard-hammer percussion), striking stones against objects that they position in front of
them (bipolar technique), striking stones against stationary surfaces (anvil technique) and
by throwing stones down from perches (throwing technique; see Plate 1). They use the
anvil technique more frequently than any other stone-reduction method, and tend to
produce larger flakes when they use the anvil and throwing techniques than when they use
the hard-hammer percussion and bipolar techniques (similar findings have been noted for
a pygmy chimpanzee, see Toth et al. 1993). Capuchins sometimes produce large (>4 cm.
in length) flakes which make effective tools for cutting (Westergaard and Suomi, 1994b,
under review a).
Although capuchins exhibit impressive artifact production skills, their stone-flaking
behavior is primitive when compared to that of Oldowan hominids as manifested in the
archaeological record. Capuchins do not produce large flakes through hard-hammer
percussion, use precise, highly controlled striking actions, actively search for acute core
angles, use flake scars as striking platforms or consistently produce invasive flake scars.
Similar limitations have been noted in the stone-flaking behavior of a pygmy chimpanzee
(Toth et al. 1993). These findings indicate that nonhuman primate stone flaking is simpler
than that of Homo habilis. It seems likely that the first hominid or proto-horninid tool
makers were similarly constrained, and that their artifacts would probably resemble those
produced by great apes and capuchins. Subsequent refinement of these relatively simple
stone-reduction techniques would have led to production of larger, more complex forms,
such as those which characterize Acheulean technology (Isaac 1982).

The use of stones as cutting tools

There is considerable evidence that Oldowan hominids used stone artifacts to butcher
animals. Processing fresh carcasses (obtained through scavenging or hunting) would have
provided Homo habilis with valuable nutrients, and would have been greatly facilitated by
the use of stone tools. Fossilized animal bones provide important clues to prehistoric
animal butchery, including the tell-tale marks of cutting tools. Since stone-tool cut marks
on Plio-Pleistocene bone were first described systematically (Bunn 1981; Bunn and Kroll
1986; Potts and Shipman 1981), bone surface modification has played an important role in
reconstructing the development of hominid lithic technology (Schick and Toth 1993).
Capuchins use the sharp edges of stones and stone flakes as cutting tools (Westergaard
and Suomi 1994a, 1994b). They typically place the stone against the material to be cut, and
then push down on the tool in a back-and-forth motion. Some animals combine stones as a
human would combine a hammer with a chisel. This technique can reduce stone and
produce additional sharp-edged flakes. Capuchins use stone tools to cut acetate and to
deflesh bone fragments. When defleshing bone fragments, capuchins produce surface
4 Gregory Charles Westergaard

(a) (b)
Plate 1 A sample of stone
artifacts produced by capuchin
monkeys: a) a core from which
a single flake has been re-
moved; b) a core which has
been flaked along two opposite
edges; c) a core which has been
flaked along three adjoining
edges; and d) a core (right) and
four flakes. The ruler in each
photograph is marked in centi-
metres.

(c)

(d)
 The stone-tool technology of capuchin monkeys 5

Plate2 A modifiedswine rib bone fragment.The modificationswere producedby the pounding

andcuttingactionsof an adultmalecapuchinmonkeyas he defleshedthe fragmentwitha stonetool.
The dark vertical lines on either side of the abrasion were drawn by the author. Original
magnificationapproximately6X. ReprintedfromWestergaardandSuomi(1994a).

modifications (cut marks and percussion marks) analogous to those which have been found
at early stone age archaeological sites (see Plate 2).
Fossilized primates have been found at several locations which contained bones that had
been modified with stone tools (Potts 1988). However, little consideration has been given
to the idea that tool-using non-hominid primates could have modified at least some of
these specimens (Wynn and McGrew 1989). This is particularly surprising because
chimpanzees and Oldowan hominids share similar environments (McGrew, Baldwin and
Tutin 1981), and propensities for hunting (Riss and Busse 1977) and tool-aided
bone-marrow processing (Boesch and Boesch 1989). Capuchins use tools in nature
(Boinski 1988; Fernandes 1991) and I speculate that they produce and use stone artifacts
within their New World habitat.

Neurobehavioral laterality and the evolution of symbolic communication

For many years researchers have examined prehistoric artifacts to determine if they were
produced by predominantly right- or left-handed populations (e.g., Brinton 1896;
Mortillet 1890). In this context Toth (1985b) has provided convincing evidence that
right-handedness evolved in hominids by 1.4 to 1.9 million years ago. Toth based his
argument on an analysis of lithic artifacts which revealed disproportionate clockwise
rotation of stone cores during flaking (the typical pattern for right-handed individuals).
Recent experiments indicate that capuchins exhibit population-level right hand biases
for stone artifact production and the use of stones as cutting tools (Westergaard and Suomi
under review b). In one experiment fourteen capuchins struck stones against hard
surfaces, including cage structures and other stones. Ten of these animals exhibited a
6 Gregory Charles Westergaard

lateral bias for these actions, and nine of them were biased towards use of their right hand.
These findings were not related to the subject variables age or sex class. In a second
experiment six monkeys used stones as cutting tools to penetrate acetate and obtain prized
food. Each of these animals exhibited a right-handed bias for this activity.
MacNeilage, Studdert-Kennedy and Lindblom (1987) have hypothesized that primate
handedness evolved as a result of selection pressures that favored lateral bias for feeding
and postural support. These authors believe that primates first evolved a left-hand
specialization for visually guided reaching and then, with the emergence of the opposable
thumb and decreased postural demands on the right upper limb, a right-hand specializ-
ation for manipulation and bimanual co-ordination. Chimpanzees are more strongly
right-handed for throwing from a bipedal stance than from a quadrupedal one (Hopkins
et al. 1993), a finding which is consistent with this idea. Hominids are believed to have
stood bipedally prior to evolving intensive tool-using skills (Johanson, Johanson and
Edgar 1994), and I suggest that the first bipedal hominids were predisposed toward
right-handed stone manipulation and flaking.
Several authors have hypothesized a relationship between the evolution of language and
complex object manipulation. Frost (1980) has argued that hemisphere specialization for
language and manipulation evolved in hominids as a result of selection pressures which
favored asymmetric use of the limbs during tool making. Corballis (1991) believes that
hominid gestural communication competed with the use of tools. The transition to vocal
communication is said to have freed the hands and facilitated a generative relationship
between manipulation and language. Greenfield (1991) has argued for parallel develop-
ment of manipulative and communicative skills in human children (Greenfield, Nelson
and Saltzman 1972). Greenfield hypothesizes that during the first two years of human life a
common neural substrate (roughly Broca's area) underlies the hierarchical structure of
both speech and the use of tools. An evolutionary homologue for Broca's area is said to
have provided a neural foundation for language prior to the divergence of great apes and
hominids. This hypothesis is supported by evidence of a Broca's area homologue in Old
World monkeys (Greenfield 1991), and parallel constraints in symbolic communication
and the use of tools in apes (Greenfield and Savage-Rumbaugh 1990).
The hypothetical relationship between language and object-manipulation suggests that
research on lateralized use of tools may help to define neurobehavioral continuity among
primates. Connolly and Dalgleish (1989) examined spoon use in human infants and found
an increasing right-hand bias between the ages of one and two years. Infant chimpanzees
exhibit a high degree of right-handedness when they first use stones to crack open nuts.
This pattern reverses during an intermediate period before stabilizing at the adult level of
no population-level bias towards either hand (Boesch 1990). I suggest that differences
between human and nonhuman primates in the development of hand preference for the
use of tools reflect a language-based discontinuity in the evolution of primate neuro-
behavioral laterality (Westergaard 1994).
The propensities of capuchins to produce and use tools are anomalous among New
World primates (Parker and Gibson 1977; Visalberghi 1990; Westergaard and Fragaszy
1987), but primitive when compared to those of modern humans. Capuchin combinatorial
behavior is dominated by a simple object-pairing strategy whereas human combinatorial
behavior is dominated by a sequential strategy which involves assembling structures and
 The stone-tool technology of capuchin monkeys 7

combining them with specific targets (Greenfield 1991; Westergaard and Suomi 1994c).
Capuchins combine vocalizations in a manner that satisfies at least some of the conditions
for lexical syntax (Robinson 1984), although the extent to which these monkeys organize
their communicative behavior is uncertain (Westergaard and Suomi 1994c). Pifion and
Greenfield (1994) have suggested that the neural substrate which mediates both language
and the use of tools could have begun its evolution at least 45 million years ago when New
World monkeys first diverged from the main primate lineage. If this hypothesis is correct,
then the evolutionary origins of symbolic communication may be much more ancient than
we could possibly determine from comparing humans with our nearest living relative, the
chimpanzee (Piion and Greenfield 1994).

Conclusions

This report describes how a simple stone-tool technology in monkeys can help us to
understand psychological processes which facilitate stone-tool production in hominids.
Evidence presented here and elsewhere indicates that the cognitive and biomechanical
conditions of pre-adaptation for the production and use of simple stone tools are present in
great apes and capuchin monkeys (Toth et al. 1993; Westergaard and Suomi 1994b; Wynn
and McGrew 1989). It is hypothesized that an ancient neural substrate underlies the
development of both language and the use of tools (Greenfield 1991), and that research on
the lithic capabilities of extant monkeys and apes will facilitate greater understanding of
the mental life of Homo habilis and other prehistoric tool-making hominid species.

Acknowledgements

The author was supported by an Intramural Research Training Award from the National
Institute of Child Health and Human Development. The capuchins used in this research
were housed in Laboratory of Comparative Ethology facilities at the National Institutes of
Health Animal Center in Poolesville, MD. I thank Clara Menuhin-Hauser and L. V. H.
M. H. Westergaard for commenting on an earlier draft of this manuscript.

Laboratory of Comparative Ethology

National Instituteof Child Health and Human Development
Poolesville, MD

Note

Correspondence should be addressed to Gregory Charles Westergaard, National Institute

of Health Animal Centre, PO Box 529, Poolesville, MD 20837, USA (phone: (301)496-
0444; fax: (301)496-0630).
8 Gregory Charles Westergaard

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