Behavioural Processes 65 (2004) 8793

Tool-use and tool-making by captive, group-living orangutans

(Pongo pygmaeus abelii) at an artificial termite mound
Masayuki Nakamichi
Laboratory of Ethological Studies, Faculty of Human Sciences, Osaka University, Suita 565-0871, Japan

Accepted 16 July 2003

Abstract
The present study examined the use and making of tools to obtain foodstuffs in artificial-mound holes by five captive,
group-living Sumatran orangutans (Pongo pygmaeus abelii). Three adult orangutans frequently stripped leaves and twigs from
a branch provided (tool-making), and then inserted the tool into a hole to obtain foodstuffs (tool-using). A 5-year-old female
juvenile usually used the tools that adult orangutans had previously used, but rarely made tools herself. A 2-year-old male infant
did not use any tools. The adult orangutans tend to leave one to several leaves at the top of the branch than to leave many leaves
on the branch or to strip all leaves. It seemed likely that tools with appropriate leaves are easier to insert into holes and obtain
more foodstuffs, compared with branches with many leaves or sticks without any leaves. When the orangutans were unable to
insert a tool into a hole, they usually modified the tool and/or changed their tool-using technique, such as changing how they
grasped the tool. These findings are discussed from the perspectives of the orangutans behavioral flexibility regarding tool-use
skills and hierarchical organization in food-processing techniques.
2003 Elsevier B.V. All rights reserved.
Keywords: Pongo pygmaeus abelii; Orangutan; Tool use; Tool making; Tool modification; Tool learning

1. Introduction in captivity have been reported to intentionally throw

Many observations have shown that wild chim-
panzees (Pan troglodytes) are consistent and habitual
tool-users and tool-makers (see review by McGrew,
1992). On the other hand, there have been no pre-
vious reports on tool-using behavior in wild gorillas
(Gorilla gorilla) or bonobos (Pan paniscus), despite
long-term field studies (Fossey, 1983; Kano, 1992).
There have been many reports that captive gorillas in
zoos use objects as tools, such as by throwing missiles,
or sponging and probing with sticks (Boysen et al.,
1999; Parker et al., 1999). Moreover, some gorillas
Tel.: +81-6-6879-8129; fax: +81-6-6879-8010.
E-mail address: naka@hus.osaka-u.ac.jp (M. Nakamichi).
sticks into the foliage of trees, which the gorillas
could not climb due to electric wire, to knock down
leaves and seeds, and to pull tree branches toward
themselves using sticks (Nakamichi, 1999). These
tool-using behaviors might reflect a hitherto unknown
aspect of the cognitive ability of this species. Thus,
observations of tool-using and tool-making among
great apes, both in the wild and in captivity, are in-
dispensable for understanding cognitive ability from
an evolutionary perspective.
There have been only a few observations of tool
use among wild orangutans (Pongo pygmaeus) (Van
Schaik et al., 1996; Fox et al., 1999; Peter, 2002),
while free-ranging rehabilitant orangutans have been
reported not only to use objects as tools in a variety

0376-6357/$ see front matter 2003 Elsevier B.V. All rights reserved.
doi:10.1016/j.beproc.2003.07.002
88 M. Nakamichi / Behavioural Processes 65 (2004) 8793
of contexts, but also to perform human-like behav-
iors with human artifacts frequently (Galdikas, 1982;
Russon and Galdicas, 1995; Russon, 1999). Observa-
tional and experimental studies on captive orangutans
have also demonstrated that their capacity for tool use
is as high as that in chimpanzees (Lethmate, 1982;
Visalberghi et al., 1995). Compared with chimpanzees,
however, there is little information available on the
spontaneous tool-using and tool-making behaviors of
orangutans. In particular, few studies have dealt with
how orangutans modify tools or change their behav-
ior after failing to use tools and little information is
available about the development of tool-use skills in
immature orangutans.
To enhance the psychological well-being of captive
chimpanzees, some zoological parks have provided
artificial mounds, designed to simulate the earth
mounds made by African termites for which wild
chimpanzees fish by inserting grass or twig tools into
the holes (Nash, 1982). Such a man-made mound
has been provided for the group-living Sumatran
orangutans (Pongo pygmaeus abelii) at the San Diego
Zoo. Adult orangutans in this group stripped leaves
and small twigs from branches (i.e. tool-making) and
inserted them into holes to obtain foodstuffs in the
mound holes (i.e. tool-using). These tool-using and
tool-making behaviors by orangutans, which could
provide important opportunities for understanding the
cognitive abilities of this species, should be analyzed
with precision.
The purpose of the present study was to investi-
gate tool-using and tool-making behaviors to obtain
foodstuffs in artificial-mound holes by the orangutans
ranging in age from 2 to 37 years at the San Diego
Zoo. I answered the following questions by ana-
lyzing video-recorded data: How and how often do
orangutans use and make tools? Do orangutans mod-
ify tools or change their behavior after failing to use
tools? Can immature orangutans make and use tools?
2. Materials and methods
Five Sumatran orangutans living at the San Diego
Zoo were observed from late August to mid-December
1997: one full-adult male (Clyde, captive-born, 21-
year-old), one full-adult female (Josephine, wild-born,
37-year-old, estimated), one young-adult female
(Indah, captive-born, 11-year-old), a juvenile female
(Karen, captive-born, 5-year-old), and one infant
male (Satu, captive-born, 2-year-old). The four older
orangutans had lived together in a group for at least
three years before the start of the present study. The
remaining orangutan (Satu) was born in the group.
The five orangutans usually came out in a grassy
outdoor enclosure (approximately 200 m2 ) between
8:30 and 9:00 h, and stayed there until they returned
to their indoor cages at around 17:00 h. There was the
artificial mound (180 cm high and 500 cm round) in
the center of the enclosure. The mound, constructed of
concrete, had five holes, each 38 mm in diameter and
5681 cm from the ground. These gave access to plas-
tic pipes of varying length from 43 to 76 cm. Some
of the holes were filled with foodstuffs such as mus-
tard, BBQ sauce, applesauce, peanut butter slurry etc.
up to 10 to 20 cm from the bottom, every morning be-
fore the orangutans came out. The mound was set up
in the enclosure at least three years before the start of
the present study.
Many fresh branches (approximately 50100 cm
long and 0.52 cm in diameter at the thick end) with
small twigs and leaves were provided every morning.
These branches consisted of figs of several species
(Ficus spp.), eugenia (Syzygium spp.), mulberry
(Morus alba), and hibiscus (Hibiscus rosa-sinensia).
The orangutans ate the leaves and seeds from these
branches, and also used them as tools to obtain the
foodstuffs in the mound.
Observations were usually conducted for one to
three hours immediately after the orangutans came out
in the enclosure, since they were more likely to show
tool-using and tool-making behaviors during this time
of the day. The orangutans were observed from the
public space for a total of 44.5 h (19 days of observa-
tions).
Tool-using and tool-making behaviors by orangutans
were recorded with an 8-mm videotape recorder.
Based on this videotaped data, I recorded those
branches which they selected to obtain the food-
stuffs in the mound, i.e. new branches or sticks. New
branches had not previously been used as a tool or
eaten by orangutans, and thus still had at least several
leaves and/or small twigs. Sticks had few or no leaves
since orangutans had already used them to obtain
foodstuffs or had eaten the leaves. I also recorded
whether or not orangutans stripped off leaves or twigs
 M. Nakamichi / Behavioural Processes 65 (2004) 8793 89

when they used a new branch as a tool. When an
orangutan was not able to insert the tool into the hole,
for example, because there were too many leaves on
the branch, this was regarded as an unsuccessful act.
On the other hand, when the tool was inserted into
the hole, this was regarded as a successful act. I also
recorded the behaviors of tool-users after unsuccess-
ful acts.
3. Results
3.1. General patterns of tool-using and tool-making
behavior
Table 1 summarizes tool-using and tool-making
behaviors by the orangutans. The three adult and
one juvenile orangutans showed both tool-using and
tool-making behaviors to obtain foodstuffs in the
mound, whereas the remaining 2-year-old male infant
did not, although he did insert his index finger into
the hole, wiped the inside of the hole and then put it
in his mouth to lick.
To obtain foodstuffs from within the mound, the
5-year-old juvenile female (Karen) was recorded to
pick up sticks 27 times (93% of the total acts of
picking up materials to obtain foodstuffs), while she
selected new branches only twice (P < 0.01 on a bi-
nomial test). Thus, she customarily used tools that had
been previously used by other orangutans, while she
rarely selected new branches from which the leaves
and twigs needed to be stripped. The adult female
(Josephine) and young-adult female (Indah) picked
up new branches as often as sticks (Ps > 0.10 on

Table 1
Numbers (%) of various acts performed by orangutans at the San Diego Zoo, CA, USAa
Clyde (21, m)b Josephine (37, f) Indah (11, f) Karen (5, f)

Picking up materials to obtain foodstuffs

New branches 57 (64.0) 43 (41.0) 13 (43.3) 2 (6.9)
Sticks 32 (36.0) 62 (59.0) 17 (56.7) 27 (93.1)
Stripping leaves/twigs from new branches
Yes 39 (68.4) 37 (86.0) 11 (84.6) 2
No 18 (31.6) 6 (14.0) 2 (15.4) 0
Making the tool
Stripping all leaves/twigs 5 (12.8) 1 (2.7) 2 (18.2) 1
Leaving leaves at the top of the branch 15 (38.5) 31 (83.8) 7 (63.6) 0
Stripping leaves/twigs inappropriately 19 (48.7) 5 (13.5) 2 (18.2) 1
Which end of the tool was inserted first?
Distal (thin) end 658 (95.8) 583 (99.1) 85 (96.6) 238 (100)
Proximal (thick) end 29 (4.2) 5 (0.9) 3 (3.4) 0 (0)
Inserting the tool into a hole
Successful 563 (82.0) 553 (94.0) 86 (97.7) 221 (92.9)
Unsuccessful 124 (18.0) 35 (6.0) 2 (2.3) 17 (7.1)
Behavior after unsuccessful attempts
Modify tool/change technique for inserting the tool 103 (90.4) 29 (82.9) 2 14 (82.4)
Perform again in the same manner 7 (6.1) 3 (8.6) 0 2 (11.8)
Leave the mound 4 (3.5) 3 (8.6) 0 1 (5.9)
Details of modify tool/change technique for inserting the tool
Modify the tool 27 (26.2) 6 (20.7) 0 0 (0)
Change technique for inserting the tool into a hole 66 (64.1) 14 (48.3) 0 11 (78.6)
Change tool 7 (6.8) 7 (24.1) 2 1 (7.1)
Change hole 4 (3.9) 3 (10.3) 0 3 (21.4)
a The percentages in parenthesis are shown as the ratio of the number of acts in a sub-category to the number of total acts in the main

category. For example, Clyde picked up new braches at 64.0% of the total acts of picking up materials to obtain foodstuffs.
b Age in years, m: male, f: female.
90 M. Nakamichi / Behavioural Processes 65 (2004) 8793
binomial tests), while the adult male (Clyde) picked
up new branches more often than sticks (P < 0.05 on
a binomial test).
When picking up new branches, all of the three
adults stripped some leaves and twigs from the branch
at least two-thirds of the time before inserting them
into the holes. When stripping leaves and twigs from
new branches, Josephine and, to a lesser degree, Indah
were more likely to leave one to several leaves at the
top of the branch (i.e. good tool) than to leave many
leaves on the branch or to strip all leaves (bad tool).
It seems likely that good tools are easier to insert into
holes and obtain more foodstuffs, compared with bad
tools. None of the orangutans were observed to chew
or bite the leaves from the tip of the tool when they
put it into their mouth to obtain foodstuffs. Josephine
made good tools significantly more often than Clyde
(2 = 16.36, P < 0.01).
Clyde, Josephine and Indah striped leaves and twigs
from branches with their fingers or picked them off
from the distal (thin) to proximal (thick) end of the
branch, while grabbing the distal end of the branch
in the fist, in 29 of 39 trials (74.4%), in 34 of 37
trails (91.9%) and in all 11 observed trials (100%),
respectively. It seemed likely that leaves and twigs
were more easily detached from the stem when they
were stripped from the distal to proximate end than in
the opposite direction.
All four orangutans inserted tools into holes with the
distal end first in at least 95% of acts. They sometimes
tried to insert the stick with the distal end first, even
though this end seemed to be too limp to be inserted
compared to the opposite end.
The rate of successful acts of inserting the tool into
the hole for Clyde was significantly lower than those
for each of the three females (Table 1): Josephine
(2 = 42.48, P < 0.01), Indah (2 = 14.26,
P < 0.01), Karen (2 = 16.28, P < 0.01). There
were no significant differences among the three fe-
males: Josephine and Indah (2 = 2.00, P > 0.10),
Josephine and Karen (2 = 0.41, P > 0.10), Indah
and Karen (2 = 2.78, P > 0.10).
After more than 80% of unsuccessful acts, they
again tried to insert the tool into the hole after modi-
fying the tool and/or changing their technique. Their
tools were modified by stripping obstructive leaves
and twigs from the branch so that it could enter the
hole smoothly. They also changed their technique by
changing how they grasped the tool (i.e. grasping the
more distal end of the branch), by changing the hand
in which the tool was grasped (i.e. right to left hand
or vice versa), or by changing one-hand to two-hand
grasping (i.e. grasping the distal end of the tool in
one hand and the proximal end in the other). Only
Josephine pushed the leaves at the top of the branch
into the hole with the extended index finger of her
free hand: these leaves made it more difficult to insert
the branch but also made the tool more effective for
obtaining more foodstuffs from the hole. Clyde and
Josephine were also recorded to change branches and
holes after unsuccessful acts. Karen was recorded to
modify the tool only once (7%) after unsuccessful acts,
while she changed her technique, such as by changing
how she grasped the tool, 11 times (79%).
3.2. Josephines special tool
Josephine was once observed to make a special tool:
she chewed a 120-cm long (estimated) stick for ap-
proximately 40 cm from the distal end for 30 s, and
crumpled it, which effectively made a sponge at the
end of an 80 cm handle. She pushed the sponge into the
hole, by grasping the middle of the stick and pushing
it into the hole, then withdrew the sponge by extract-
ing the stick, and kept it in her mouth for 1025 s to
suck out the liquid. While she sucked the sponge the
fifth time, her 2-year-old son (Satu) pulled the stick.
As a result, the stick detached from the sponge, which
still remained in Josephines mouth. She continued
sucking the sponge for another 18 s, and then pushed
it from her mouth. Thereafter, she inserted the stick
into the same hole seven more times.
4. Discussion
The results of the present study demonstrate that
adult orangutans habitually made and used tools to
obtain foodstuffs, a 5-year-old juvenile female was
also a habitual tool-user but not a tool-maker, and a
2-year-old infant male did not use tools. These results
are largely consistent with a finding in the wild. Ac-
cording to Fox et al. (1999), a wild 3- to 4-year-old
juvenile female orangutan frequently observed her
mother break off a branch and strip it to fish for ants
in tree holes, but did not do so herself. However, she
 M. Nakamichi / Behavioural Processes 65 (2004) 8793
made her own tool and probed tree holes at the age
of 5 to 6 years. Although new branches as well as
used sticks were usually available near the mound
in the morning, the 5-year-old female in the present
study preferred using tools that had been previously
used by adults rather than making tools from new
branches, which were less elaborate than those made
by adult orangutans. These findings indicate that the
use of tools that have been previously used by adults
may be an important step for juvenile orangutans to
acquire appropriate tool-using and tool-making skills.
The special tool (i.e. stick with a sponge) made by
the adult female in this study (Josephine) consisted of
two parts and each had its own function; i.e. the stick
was used to push the sponge into the hole and to ex-
tract it again, while the sponge, which could not be
used without the stick, was used to soak up more food.
Moreover, the adult orangutans including Josephine
stripped leaves from new branches, but tended to leave
one to several leaves at the top of the branch to ob-
tain more foodstuffs. Such a tool with leaves can also
be considered a single tool consisting of two parts,
each with its own function. Therefore, we may say
that Josephines special tool and the tools with ap-
propriate leaves at the top resemble man-made tools,
such as spoon and fork, which have functionally dis-
tinct parts. Wild chimpanzees habitually use tools for
ant- and termite-fishing (McGrew, 1992). Each tool for
fishing seems to consist of only one part, and thus has
only one function. Since it is well known that chim-
panzees use a variety of objects as tools (McGrew,
1992), the question of whether or not chimpanzee tools
have functionally distinct parts should be addressed
systematically. However, this is beyond the scope of
the present study.
More interestingly, the orangutans did not bite off
the leaves left at the top of the tool when they put the
tools in their mouth to obtain the adherent foodstuffs.
Instead, they just wiped the leaves between their lips
or pressed them between their tongue and the roof
of their mouth. Josephine also sucked the sponge at
the top of her special tool, but did not bite off it.
These observations suggest that the orangutans may
understand the functional value of the leaves to obtain
more foodstuffs. Thus, they may understand causality
between the use of the tool with leaves and its out-
come. On the other hand, it is also probable that the
orangutans learned of an association between them
91
through trial and error. To evaluate comprehension of
a causal relation, it can be useful to examine perfor-
mance in novel tasks in which the potential tools can
be used to perform different acts (Visalberghi et al.,
1995; Vizalberghi and Tomasello, 1998). In the present
study, adult orangutans who had been presented with
the artificial mound for at least three years were ob-
served to use and modify probing tools, but no in-
formation was available on the early stages of when
they acquired this tool-using and tool-making behav-
ior. Moreover, although it was difficult to insert new
branches into the holes, in 31, 14 and 15% of the time
that the adult male, full-adult female and young-adult
female in this study picked up a new branch, respec-
tively, they inserted the new branch into a hole with-
out stripping any leaves or twigs (see Table 1). Based
on these facts, the idea that the orangutans learned an
association between the use of a tool with leaves and
its outcome may be more parsimonious than the idea
that they understand the functional value of the leaves
for obtaining more foodstuffs.
Russon (1998) observed food-processing tech-
niques of ex-captive juvenile orangutans that had been
reintroduced to the wild in Kalimantan, Indonesia,
and concluded that the orangutans showed hierarchi-
cal organization when obtaining difficult foods. She
suggested that hierarchization generates complex,
higher level cognition by re-using existing cogni-
tive units to build new cognition structures (Russon,
1998). Byrne and Byrne (1991) also pointed out dex-
terous food-processing techniques in wild mountain
gorillas, whose diet is dominated by a few herbaceous
species. Since these herbs are typically defended by
stingers, hooks, spines or fibrous outer cases, multiple
processing steps are needed, where each step must be
completed before the next can begin, and sequences
of 4 to 6 steps are normal. In the present study, the
orangutans sometimes failed to insert the tool into
the hole. However, in more than 80% of these unsuc-
cessful trials, they modified the tool or changed their
technique to successfully insert the tool into the hole,
such as by stripping obstructive leaves and twigs from
the branch or grasping the more distal end of the
branch. These acts may reflect hierarchical organiza-
tion in orangutan food-processing techniques. Russon
(1998) also offered evidence that in orangutans, the
cognition that governs motor actions was hierarchi-
cally organized to correct errors.
92 M. Nakamichi / Behavioural Processes 65 (2004) 8793
On the other hand, the orangutans in the present
study inserted their tools into holes with the distal
end first in almost all cases. Even immediately after
acts in which they could not insert the tool into the
hole due to fraying of the distal end, they did not try
to insert the tool from the opposite end (i.e. the prox-
imal end). As discussed above, orangutans commonly
show hierarchically organized food-processing tech-
niques. However, the orangutans in the present study
persevered in inserting the tool with the distal end
first, and the reason for this perseverance is unknown.
These results suggest that the orangutans behavioral
flexibility regarding tool-use skills or hierarchical
organization in food-processing techniques may be
context-dependent.
Wild Borneo orangutans Pongo pygmaeus abelii
have rarely been observed to display tool-using skills
despite long-term field studies (Galdikas, 1989), while
Sumatran orangutans (P. p. abelii) in a restricted area
are habitual tool-users: they use sticks to fish for ants
from tree holes and to extract seeds from hard-husked
fruit (Fox et al., 1999; Van Schaik et al., 1996). Based
on their finding that the wild orangutans that use tools
are the most gregarious and socially tolerant among
the known study populations, Van Schaik et al. (1999)
proposed that some level of gregariousness and some
level of tolerance during foraging are required for so-
cial transmission. In the wild, orangutans lead rather
solitary lives, while in captivity they can live in social
groups, like the orangutans at the San Diego Zoo in the
present study. Such captive, group-living orangutans
can offer important opportunities for conducting both
uncontrolled observations and experimental manipu-
lations, and thus promote a better understanding of
their cognitive abilities. In particular, by examining the
performance of group-living orangutans in novel tasks
including tool-using and tool-making, we can evalu-
ate the importance of gregariousness and tolerance for
social transmission, as proposed by Van Schaik et al.
(1999), and thus gain more information about how
much sociality influences the process of building new
cognition structures in orangutans.
Acknowledgements
I wish to express my sincere thanks to D. Lindburg,
the Center for Reproduction of Endangered Species
(CRES), for his kind permission to perform this re-
search at the San Diego Zoo and to H. Fitch-Snyder,
behavior specialist at CRES, for making the neces-
sary arrangements for my research. I also thank M.
Bate and the other keepers at the San Diego Zoo for
their kind help during the present observations. A.
Toth taught me the names of the trees in the enclo-
sure and those of branches given to the orangutans.
Two anonymous referees made helpful comments. I
am also grateful to J. Moore, D. Tuzin, A. Mitsuta
and the staff of the Laboratory of Ethological Stud-
ies and Department of Ethology, Faculty of Human
Sciences, Osaka University, for their help and encour-
agement. This work was supported by a grant from
the Ministry of Education, Sciences, and Culture of
Japan.
References
Boysen, S.T., Kuhlmeier, K., Halliday, P., Halliday, Y.M., 1999.
Tool use in captive gorillas. In: Parker, S.T., Mitchell, R.W.,
Miles, H.L. (Eds.), The Mentality of Gorillas and Orangutans.
Cambridge University Press, Cambridge, pp. 179187.
Byrne, R.W., Byrne, J.M., 1991. Hand preferences in the skilled
gathering tasks of mountain gorillas (Gorilla g. berengei).
Cortex 27, 521546.
Fossey, D. 1983. Gorillas in the Mist. Houghton Mifflin Company,
Boston.
Fox, E.A., Sitompul, A.F., van Schaik, C.O., 1999. Intelligent
tool use in wild Sumatran orangutans. In: Parker, S.T.,
Mitchell, R.W., Miles, H.L., (Eds.), The Mentality of Gorillas
and Orangutans. Cambridge University Press, Cambridge,
pp. 99116.
Galdikas, B.M.F., 1982. Orang-utan tool use at Tanjung Putting
Reserve, Central Indonesian Borneo (Kalimantan Tengah). J.
Hum. Evol. 10, 1933.
Galdikas, B., 1989. Orangutan tool use. Science 243, 152.
Kano, T., 1992. The Last Ape: Behavior and Ecology of Pygmy
Chimpanzees. Stanford University Press, Stanford.
Lethmate, J., 1982. Tool-using skills of orangutans. J. Hum. Evol.
11, 4964.
McGrew, W.C., 1992. Chimpanzee Material Culture. Cambridge
University Press, Cambridge, p. 277.
Nakamichi, M., 1999. Spontaneous use of sticks as tools by captive
gorillas (Gorilla gorilla gorilla). Primates 40, 487498.
Nash, V.J., 1982. Tool use by captive chimpanzees at an artificial
termite mound. Zoo Biol. 1, 211221.
Parker, S.T., Kerr, M., Markowitz, H., Gould, J., 1999. A survey
of tool use in zoo gorillas. In: Parker, S.T., Mitchell, R.W.,
Miles, H.L., (Eds.), The Mentality of Gorillas and Orangutans.
Cambridge University Press, Cambridge, pp. 188193.
Peter, H.H., 2002. Tool use to modify calls by wild orang-utans.
Folia Primatol. 72, 242244.
 M. Nakamichi / Behavioural Processes 65 (2004) 8793
Russon, A.E., 1998. The nature and evolution of intelligence in
orangutans (Pongo pygmaeus). Primates 39, 485503.
Russon, A.E., 1999. Orangutans imitation of tool use: a cognitive
interpretation. In: Parker, S.T., Mitchell, R.W., Miles, H.L.,
(Eds), The Mentality of Gorillas and Orangutans. Cambridge
University Press, Cambridge, pp. 117146.
Russon, A.E., Galdicas, B., 1995. Constraints on great apes
imitation: model and action selectivity in rehabilitant orangutans
(Pongo pygmaeus) imitation. J. Comp. Psychol. 109, 517.
Van Schaik, C.P., Fox, E.A., Sitompul, A.F., 1996. Manufacture and
use of tools in wild Sumatran orangutans. Naturwissenschaften
83, 186188.
93
Van Schaik, C.P., Deaner, R.O., Merrill, M.Y., 1999. The conditions
for tool use in primates: implications for the evolution of
material culture. J. Hum. Evol. 36, 719741.
Visalberghi, E., Fragaszy, D.M., Savage-Rumbaugh, S., 1995.
Performance in a tool-using by common chimpanzees (Pan
troglodytes), bonobos (Pan paniscus), an orangutan (Pongo
pygmaeus), and capuchin monkeys (Cebus apella). J. Com.
Psychol. 109, 5260.
Vizalberghi, E., Tomasello, M., 1998. Primate causal understanding
in the physical and psychological domains. Behav. Processes
42, 189203.