Journal of Comparative Psychology © 2009 American Psychological Association

2009, Vol. 123, No. 4, 385–390 0735-7036/09/$12.00 DOI: 10.1037/a0016275

New Evidence on Imitation in an Enculturated Chimpanzee (Pan troglodytes)

Lara Carrasco, Sandra Posada, and Montserrat Colell

University of Barcelona

Imitation in the great apes continues to be an active field of research and one that is not free of
controversy. Several studies suggest that these species do not tend to match the motor movements of the
model they observe, but try to achieve the same results using their own methods (emulation of results).
In the studies reviewed, gestures have been used very infrequently outside an intraspecific communica-
tive context to evaluate imitation. In fact, the imitation of gestural actions has been tested only in
4 individual great apes. This study assessed a chimpanzee’s (Pan troglodytes) ability to imitate 52 actions
in 4 categories. The levels of accuracy attained by the subject in her imitations exceeded those described
in previous studies. Moreover, contrary to the idea defended in some articles, the subject seemed to find
it easier to imitate gestures than actions with objects.

Keywords: social learning, imitation, emulation, chimpanzees, gestures

Imitation is a form of social learning characterized by its cog-
nitive complexity (Byrne, 1995; Whiten & Ham, 1992) and rele-
vance in the acquisition and transmission of new behaviors
(Whiten, Horner, & Marshall-Pescini, 2003). Traditionally, non-
human primates have been considered to be great imitators (Hag-
gerty, 1913; Yerkes, 1916; Yerkes & Yerkes, 1929, cited in Call &
Carpenter, 2003), but their ability to do so is currently being
questioned, given that some cases first considered to be evidence
of imitation have since been explained by simpler social learning
processes (Zentall, 1996). As indicated by Heyes (1994), Russon
and Galdikas (1995), and Giraldeau (1997), in order to imitate, the
subject must acquire new actions or new ways to perform actions
after observing them being performed by a model. It is also
empirically useful to define imitation by exclusion, that is, by
ruling out other mechanisms (Zentall, 1996). However, the dis-
tinction between imitation and other forms of social learning,
particularly emulation, has also generated considerable contro-
versy in the literature (Horner & Whiten, 2005).
The different approaches to studying imitation in nonhuman
primates have also led to contradictory results. The standard ap-
proach uses the “two-action method” in which individuals are
shown two alternative ways to receive a reward beyond their
Lara Carrasco, Sandra Posada, and Montserrat Colell, Department of
Psychiatry and Clinical Psychobiology, Faculty of Psychology, University
of Barcelona, Barcelona, Spain.
This research project was supported by an FI grant from the Ministry of
Education of the Autonomous Community of Catalonia (2005/08). We are
grateful to the Centro de Conservación de Primates, Rainfer, for permission
to perform the study at the center and to the staff for their close collabo-
ration. We must also thank Raul Cabrera and Tamara de la Montaña, the
independent observers in the evaluation period. Finally, many thanks to
Miguel Calvo for his advice on statistics.
Correspondence concerning this article should be addressed to Lara
Carrasco, Department of Psychiatry and Clinical Psychobiology, Faculty of
Psychology, University of Barcelona, Campus de la Vall d’Hebron, Passeig
de la Vall d’Hebron 171, 08035 Barcelona, Spain. E-mail:
laracarrascopesquera@hotmail.com
reach: Imitation is defined as a statistically significant tendency to
copy the observed method. Thus, Whiten et al. (1996) consider the
results of their study to provide evidence of imitation within an
imitation– emulation continuum. On the other hand, several re-
searchers argue that great apes learn to use tools by emulation, that
is, not by copying the steps of the action but by reproducing the
final result (Call & Tomasello, 1994; Myowa-Yamakoshi & Mat-
suzawa, 2000; Nagell et al., 1993; Tomasello, Davis-Dasilva, Ca-
mak, & Bard, 1987). To be able to differentiate between imitation
and emulation, it is first necessary to effectively separate actions
from their results. The strategies used to do this could be (a)
blocking some sources of information; (b) replacing objects used
in the first demonstrations with slightly different ones; or (c)
presenting actions that do not modify the physical environment in
a permanent or observable way, as occurs when an attempt is made
to reproduce gestures (Bjorklund, Yunger, Bering, & Ragan, 2002;
Call, 2001; Myowa-Yamakoshi & Matsuzawa, 2000). In studies of
juvenile great apes, the imitation of gestural actions has been
evaluated in three chimpanzees (Custance et al., 1995; Hayes &
Hayes, 1952) and one orangutan (Miles et al., 1996). According to
these authors, copying gestures is often detectable but imperfect.
For this study, we accepted the definition of imitation proposed
by the Associative Sequence Learning Theory (Heyes & Ray,
2000), and the design of our study made it possible to differentiate
between actions and their results. Very little convincing evidence
of imitation has been obtained with primates, but when such
evidence has been found, it has involved enculturated primates
(like the subject of our study) that were trained by following
requests to imitate (Custance et al., 1995; Miles et al., 1996). Our
aim was to assess whether or not chimpanzees have imitative
skills, to determine whether this study design makes it possible to
differentiate between imitation and other social learning processes
such as emulation, to evaluate whether it is more difficult to copy
actions in one category or another, and, finally, to determine
whether there is a difference in the imitation when the actions
reproduced involved in-sight body parts and out-of-sight body
parts.

385
386 CARRASCO, POSADA, AND COLELL
Method
The study was carried out on a 5-year-old enculturated female
chimpanzee (Pan troglodytes) named Lili at the Centro de Con-
servación de Primates, Rainfer, Spain. Lili was raised by hand in
a human environment (she lived with people in a house) until she
was 2 years old. When we began our research, she still had
frequent contact with humans but was housed in a cage next to that
of her mother. All sessions were filmed for subsequent analysis.
The procedure was divided into three phases:
Phase 1. Habituation of the Subject
For 4 weeks, we visited the subject 5 times per day, and with
verbal reinforcement and food we encouraged her to play and
interact positively with the model. To ensure that the actions we
were going to evaluate would be new to the subject, we allowed
the subject a session of free play before the training began. In that
session, we gave the subject the objects that we would later use in
the evaluation to check that she did not do any of the actions we
would be evaluating.
Phase 2. Training Period (9 Months)
The subject learned the basic rules of imitation (wait your
turn, match the model’s actions, and direct the action toward
yourself). Sessions were 30 – 40 min long and took place 3 times
a day, 5 days a week. The model stood in front of the subject and
demonstrated a series of actions preceded by the order “Do this”
(Table 1). Next, the objects were placed inside the subject’s cage
(bowls, rope, clothing, etc.); if she matched the sequences cor-
rectly, she was rewarded (with praise, fruit juice, dried fruit and
nuts, other food, etc.). The rewards were kept hidden and used
strategically to exact greater accuracy from the subject in her
performances. At first, the training period included 14 actions.
However, we observed that the demonstration of a limited number
of actions in the training session reduced the subject’s motivation
and made the study run less smoothly. We therefore decided to
increase the number of actions used in this period and, after a
month of study, chose 20 of these actions for systematic training
and a further 6 actions that were slightly modified to be used in the
transition. When the subject was able to copy more than 80% of
the actions in the training session over 3 consecutive days
(Custance et al., 1995), the model demonstrated 6 transition ac-
tions (similar to the training actions but slightly modified; see
Table 1). After 9 months of training, Lili was able to imitate 95%
of the training actions and the 6 transition actions.
Phase 3. Evaluation Period (1.5 Months)
The subject was shown a series of 52 new actions (not previ-
ously shown in training) classified in 4 categories, each containing
13 actions. The categories were (a) gestures (Gestures), (b) actions
in which one object was handled (Object), (c) actions in which
objects were used in combination (Object–Object), and (d) actions
in which an object was used with parts of the body other than the
hands (Object–Subject; see Table 1). Three or 4 new actions were
inserted among the actions already demonstrated in the training
period. Half of the actions demonstrated involved things within the
subject’s field of vision (i.e., actions involving in-sight body parts
such as touching the knees with the hands) and the other half
involved things that were not (i.e., actions involving out-of-sight
body parts such as touching the top of the head with the hand).
Some of them (13) had to be done with a specific extremity. Once
again, the model stood in front of the cage and demonstrated the
actions once, then waited 3–5 min before demonstrating the action
again. Whether or not the subject did the actions after seeing them,
the second demonstration was made. Every new action was pre-
sented 6 times, in three cycles of two demonstrations. After com-
pleting a cycle (52 actions done twice), the order of the actions was
changed and two more cycles were done. Throughout the entire
process, the subject was motivated by praising her and playing
with her. Although she was rewarded with food on a few excep-
tional occasions, this was not linked to any particular response. To
increase the likelihood that the actions were novel during this
phase, we used three strategies. First, we evaluated the information
gathered during the series of actions used in the training period.
Second, we eliminated behavior observed during the sessions of
free play with the objects given to the subject. Last, we used
sequences of several actions to be copied in the correct order and
completely (Whiten, 1998).
The method used to evaluate the chimpanzee’s actions was the
same as that used by Custance et al. (1995). We recorded the
sessions with a camera (SONY model HC 40E), which we placed
on a tripod behind the model. After the recording ended, we
prepared two videos. The first showed the model performing the
actions correctly and the second showed only the actions per-
formed by the chimpanzee. In other words, the second video did
not contain the parts in which the model performed the action and
it was this video that the observers assessed. Therefore, these two
people (not connected to the project) were not aware of the action
the model had performed and saw only the actions done by Lili.
Initially, we provided the observers with a complete list of the
actions included in each study period, and they were shown the
first video so they could become familiar with these actions. They
were then shown the second video and saw the actions performed
by Lili (as many times as necessary). Finally, as was done by
Custance et al. (1995), we asked the observers to indicate which
action they thought the chimpanzee was performing and to give it
a score from 0 to 3 on the basis of how well it was done (where 0
meant they could not recognize the action; 1 meant the action
performed was not complete; 2 meant the action was performed as
a mirror reflection, i.e., not with the limb the model had used; and
3 meant the action was completely correct). The observers recog-
nized 100% of the actions performed by the subject, but, naturally,
not all actions were considered perfect imitations (score of 3). We
calculated how often the same score was given by the two observ-
ers and the main researcher for each action in the second video. We
grouped the six scores of each observer as medians and compared
the scores of each observer. In 86% of the cases, the three observ-
ers gave a specific action the same score; therefore, we consider
the level of agreement to be acceptable. The 2 ⫻ 2 kappa coeffi-
cients were also calculated, and the mean agreement reached was
82.3%, with a range of .75 to .92.
Results
Of the 20 actions presented in the training period, Lili success-
fully imitated 19 (95%), as well as the 6 transition actions. In the
 IMITATION IN AN ENCULTURATED CHIMPANZEE 387

Table 1
Actions Demonstrated During the Training Period, Transition Period, and Evaluation Period

Action imitated
Action perfectly

Training
Gestures (G)
1. Clap hands Yes
2. Slap back of left hand with right hand Yes
3. Slap head with right hand Yes
4. Touch tongue with right index finger Yes
5. Touch nose with right index finger Yes

Object (O)
6. Turn bowl upside down and hit it in the center Yes
7. Turn 2 buckets upside down in sequence and hit with right hand Yes
8. Hit big buoy with both hands Yes
9. Stand brick upright and hit with right hand Yes
10. Put handkerchief over face Yes

Object–Object (O–O)
11. Thread rope through big buoy handle and hold ends Yes
12. Hold big buoy by handles and hit barrel on the ground with it Yes
13. Place bucket upright on brick Yes
14. Put small buoy inside 2 buckets in sequence Yes

Object–Subject (O–S)
16. Put sock on right hand Yes
17. Cover right foot with piece of cloth Yes
18. Put right foot in bucket Yes
19. Place glass on right ear Yes
20. Put piece of corrugated sheet on back and hit with right hand Yes

Transition
1. Hit the inside of left elbow Yes
2. Slap stomach with right hand Yes
3. Put right index finger in small buoy and hit barrel on the ground Yes
4. Thread rope through both handles of big buoy Yes
5. Put piece of corrugated sheet on head and hit it with right hand Yes
6. Slap back repeatedly with right hand Yes

Evaluation

Maximum score Imitated perfectly

Action Type awarded 1st attempt

Gestures (G)
1. Blow raspberries with right index finger and thumb in cheeks OSa 3 Yes
2. Close eyes by lowering eyelids with fingers OS 3 No
3. Make U shape with tongue OS 3 Yes
4. Make circle with right index finger and thumb ISa 3 Yes
5. Bring 2 index fingertips together and interlace them IS 3 Yes
6. Rub hands with sand IS 3 Yes
7. Touch left palm with right index finger ISa 3 Yes
8. Touch left thumb with right index finger ISa 3 Yes
9. Slap knees with both hands IS 3 Yes
10. Touch left ear with right index finger, putting arm around
back of head OSa 1 No
11. Pull out bottom lip with right hand OSa 3 Yes
12. Cover eyes with right hand OSa 3 Yes
13. Hug yourself OS 3 Yes

Object (O)
14. Turn bowl upside down and run right index finger along
edge, describing a circle ISa 3a No
15. Slide bowl upside down along floor with both hands IS 3 Yes
16. Shake bowl lid from side to side above head OS NP NP
(table continues)
388 CARRASCO, POSADA, AND COLELL

Table 1 (continued)

Evaluation

Maximum score Imitated perfectly

Action Type awarded 1st attempt

17. Unscrew container cap, drink water inside, and replace cap IS 3 Yes
18. Hold container in right hand behind back and hit it with left
a a
hand OS 1 No
19. Throw container upwards OS 1 No
20. Pick up red container and clean sand from bottom with both
hands IS 3 Yes
21. Turn 2 different-colored buckets upside down and lift 1 up
and down (chosen by model) IS 3 No
22. Put bucket on head and hit it with both hands OS 3 Yes
23. Spin the buoy like a top IS NP NP
24. Weave cloth between cage bars IS 3 No
25. Wrap scarf around neck OS 3 No
26. Twirl handkerchief above head in big circles OS 3 No

Object–Object (O–O)
27. Place bowl, put straw inside, and put on lid IS 3 No
28. Place big buoy on floor and cover it with bowl OS NP NP
29. Put buoy and stick in bowl and remove buoy with stick OS NP NP
30. Put stones in container and shake it with hands IS 3 No
31. Lay container on floor and clean with straw using both hands IS 3 Yes
32. Put stick in container and swing it above head OS 3 Yes
33. Place container on back and hit it with buoy OS NP NP
34. Turn over 2 containers of 2 chosen colors and hit 1 with
small buoy IS 3 Yes
35. Put bucket on back and put straw inside OS 3 Yes
36. Hit bucket with stick above head OS NP NP
37. Put stick through buoy, raise it to chest height, and twirl it IS 1 No
38. Thread rope through buoy, hold ends, and roll it on floor IS 3 Yes
39. Put buoy behind neck and put hat on it OS NP NP

Object–Subject (O–S)
40. Hold container lid with both hands in front of chest and
strike with feet IS 3 Yes
41. Put straw in bowl and mix it with foot IS NP NP
42. Put lid behind head like crown OS 3 Yes
43. Hold bowl by handles and put on head OS 3 Yes
44. Strike container with right foot ISa 3a No
45. Hold container with both hands and hit head OS 3 Yes
46. Hold bucket under arm and hit with right hand ISa 3a No
47. Put right foot in bucket and drag along floor ISa 1a No
48. Thread rope through big buoy handle and put on back OS 3 No
49. Wrap piece of cloth around right index finger ISa 3a Yes
50. Lay shirt out on floor and put it on IS 1 No
51. Put baseball cap on backwards OS 3 No
52. Put scarf around waist and rub lower back IS 3 Yes

Note. IS ⫽ involving in-sight body parts; OS ⫽ involving out-of-sight body parts; NP ⫽ not performed.
a
Actions performed as a mirror reflection (i.e., with the opposite extremity to the model).

evaluation period, 52 new actions were used and the percentage of
copied actions was calculated at least once for each category (see
Table 2). Only 6 actions were not given a perfect score (score of
3) after one of the attempts. To determine whether it was more or
less difficult for the subject to imitate gestures than actions related
to objects, different points were considered (see Table 2): (a) the
number of actions imitated perfectly in each category, (b) the
number of attempts required to correctly imitate an action (score of
3), (c) actions imitated successfully in each cycle of demonstra-
tions, and (d) the number of perfect imitations after the first
attempt. We assumed that the simplest actions would require fewer
demonstrations to be copied exactly, that is, fewer attempts would
be needed to achieve a perfect score. All the gestures were imitated
by Lili at some stage, and 84% of them were done correctly on the
first attempt (an average of 1.58). In the other categories, success-
ful imitation did not appear to be related to the demonstration cycle
(correct imitations were observed in all three demonstration cy-
cles). In the Object category, 9 actions were given a perfect score
(of the 11 performed), and 4 of those were performed successfully
on the first attempt. However, the mean number of attempts
needed to perform an action successfully was higher (2.55; see
Table 2). In the Object–Object category, the subject imitated 7
 IMITATION IN AN ENCULTURATED CHIMPANZEE
Table 2
Number of Actions Performed in Each Category and Results
Obtained in Different Categories
Category
Variable G O O–O O–S
Actions performed, n 13/13 11/13 8/13 12/13
% 100 84 61 92
Actions imitated perfectly, n 12 9 7 10
Mean attempts to earn a score of 3, n 1.58 2.55 2.1 2.5
Actions imitated perfectly in each cycle, n
Cycle 1–2 12 5 5 8 from the difficulty of distinguishing between imitation and other
Cycle 3–4 11 8 7 7 social learning processes, especially emulation. However, this is
Cycle 5–6 11 8 3 5 not a plausible explanation for Lili’s behavior, given that 46
Actions imitated perfectly in first attempt, n 11 4 5 6
Note. G ⫽ Gestures; O ⫽ Object; O–O ⫽ Object–Object; O–S ⫽
Object–Subject.
actions correctly, 5 of which were performed on the first attempt,
and the mean number of attempts needed was 2.1. Finally, in the
Object–Subject category, the subject imitated 10 actions correctly,
with 6 actions given a perfect score on the first attempt and a mean
of 2.5 attempts needed to perform an action successfully. Accord-
ing to Piaget (1962), actions demonstrated that involve in-sight
body parts are initially easier to imitate. In our case, the subject
correctly imitated the actions after the first or second demonstra-
tion, regardless of whether or not they involved in-sight body
parts. For each category, the proportion of actions involving in-
sight body parts that the subject performed perfectly at the first
attempt were 55% in the Gesture category (45% for actions in-
volving out-of-sight body parts), 75% in the Object category (25%
for actions involving out-of-sight body parts), 60% in the Object–
Object category (40% for actions involving out-of-sight body
parts), and 50% in the Object–Subject category (50% for actions
involving out-of-sight body parts). Finally, mirroring was ana-
lyzed: Of the 13 actions that could be performed as a mirror
reflection, Lili performed 5 with the wrong extremity during one or
more of the attempts (38.4%). However, she corrected the error in
subsequent attempts in all cases.
Discussion
The results reflect the subject’s surprising skill at imitating
actions and clearly exceed those achieved by other great apes
(Call, 2001; Custance et al., 1995; Whiten, 1998; Whiten et al.,
1996). The subject’s best results were achieved in the Gestures
category. Whereas in the study by Custance et al. (1995), the two
chimpanzees successfully imitated 27% and 35%, respectively, of
the gestures presented, Lili imitated 92%. Moreover, the subjects
in the study by Custance et al. performed very few perfect imita-
tions, whereas 38 of Lili’s actions (72%) were awarded a score of
3 by the observers. Myowa-Yamakoshi and Matsuzawa (1999)
also used the “Do this” procedure and three categories of actions
with objects (not including gestures). According to these research-
ers, the Object–Object category was the one the subjects imitated
the best and 6% of the actions were correctly reproduced in the
first test compared with 50% by Lili (26 actions). Piaget (1962)
argued that the imitation of perceptually opaque actions is more
389
difficult and represents the final stage in cognitive development, in
which case, differences should have been detected in our study in
the imitation of actions involving in-sight and out-of-sight body
parts. It was seemingly no harder for our subject to imitate actions
that involved out-of-sight body parts. A possible explanation for
these results is that our 5-year-old subject was sufficiently skilled
at mentally representing the parts of her body that were not visible
to her. Custance et al. also found no differences in 4.5-year-old
subjects regarding to their capacity to imitate actions that involved
in-sight and out-of-sight body parts.
The most intense disagreements in this field of research arise
actions were awarded a score of 3 in at least one of the six
repetitions, that is, she imitated the different movements and action
units. Therefore, in accordance with what other authors have
defended (Call & Tomasello, 1994, 1995; Nagell et al., 1993;
Tomasello et al., 1987; Tomasello, Kruger, & Ratner, 1993), the
conditions in which the subject is raised may encourage the de-
velopment of imitative skills, and enculturated animals (such as
Lili) are better at imitating than those raised by their mothers.
It should be pointed out that Lili performed 5 actions (of 13
possible actions) with the wrong extremity compared with the one
used by the model, but she corrected this error in subsequent
attempts. In studies with children (Povinelli & Deblois, 1992), this
correction of the extremity used is considered to be an indication
of the subjects’ ability to adopt the model’s perspective. The
excellent results achieved by Lili can be explained by factors that
may be of interest in future research: The relationship with the
model could explain why only enculturated animals imitated the
actions presented, which is why the habituation period is so im-
portant. However, Custance et al. (1995) did have a relationship
with the subjects in their study, and yet their results were more
modest. Another important point is the subject’s motivation. It is
possible to maintain the interest of the subjects involved through
the use of parallel games, music in the sessions, and verbal
reinforcement. Furthermore, in the series of actions used in the
training period, actions that exceeded the subject’s ability level or
were too simple were not used (Byrne, 1995). Another key aspect
was implementing the study within a context of play that encour-
aged imitation (Miklósi, 1999). Custance et al. pointed out that
rewards can influence the subject’s behavior in two ways: (a)
Withdrawing the reward can inhibit the subject from imitating the
actions, and (b) the reward itself can distract the subject, who may
pay less attention to the action demonstrated. Thus, the cases of
emulation recorded in previous studies may have been a fast way
for the subject to receive the reward, or the subject may not have
understood that the “Do this” command meant that it had to
reproduce the actions as perfectly as possible. To avoid these
problems, we used the reward system strategically by progres-
sively asking the subject for greater perfection and kept the reward
out of her field of vision. Another way to ensure that the basic rules
of imitation are complied with is to extend the training period as
much as necessary. In our study, the training period lasted 9
months, whereas it lasted 3 months in Custance et al. and 8 months
in Myowa-Yamakoshi and Matsuzawa (1999).
Another aspect to bear in mind is the age of the subjects under
study. According to the bibliography, most of the study subjects
390 CARRASCO, POSADA, AND COLELL
that showed an interest in performing tasks were individuals ages
4 to 8 years (Myowa-Yamakoshi & Matsuzawa, 1999; Tomasello
et al., 1987). However, given that imitation calls for analytical
skills by the subject and that these skills are developed with age,
adult animals were used in most of the laboratory studies (Miklósi,
1999), although researchers overlooked the fact that these individ-
uals are most resistant to changing their behavior on the basis of
what they see. Given that a certain degree of cognitive maturity is
required, the subjects chosen for studies should be at least 4 years
old.
Finally, there has been very little focus in previous studies on
the role of attention and memory. It is generally accepted that
imitation involves “more than one attempt,” a sufficiently long
demonstration time, variable attention depending on the relative
novelty of the action, and sufficient short-term memory to store
the action observed (Miklósi, 1999). Therefore, the more com-
plex the action, the more often it will have to be demonstrated to
be correctly imitated. If the demonstration of the actions is insuf-
ficient, the subject will not remember the details of the tasks and
will register them as cases of emulation. In our study, new actions
were demonstrated 6 times because we felt that limited attention
and memory capacity may be compensated for by an increase in
the number of demonstrations (Custance, Whiten, Sambrook, &
Galdikas, 2001).
In conclusion, it is clear that the imitative behavior displayed by
the subject in this experiment suggests that this animal possesses
flexible, complex visual–motor control and coordination. She also
far exceeded the success of the subjects in previous studies, which
may indicate that one of the determining factors of the results of
those studies is individual differences.
References
Bjorklund, D., Yunger, J., Bering, J., & Ragan, P. (2002). The generali-
zation of deferred imitation in enculturated chimpanzees (Pan troglo-
dytes). Animal Cognition, 5, 49 –58.
Byrne, R. (1995). The thinking ape. The evolution of intelligence. Oxford,
England: Oxford University Press.
Call, J. (2001). Body imitation in an enculturated orangutan (Pongo pyg-
maeus). Cybernetics and Systems, 32, 97–119.
Call, J., & Carpenter, M. (2003). On imitation in apes and children.
Infancia y aprendizaje, 26, 325–349.
Call, J., & Tomasello, M. (1994). The social learning of tool use by
orangutans (Pongo pygmaeus). Human Evolution, 9, 297–313.
Call, J., & Tomasello, M. (1995). Use of social information in the problem
solving of orangutans (Pongo pygmaeus) and human children (Homo
sapiens). Journal of Comparative Psychology, 109, 308 –320.
Custance, D., Whiten, A., & Bard, K. (1995). Can young chimpanzees
(Pan troglodytes) imitate arbitrary actions? Behaviour, 132, 837– 859.
Custance, D., Whiten, A., Sambrook, T., & Galdikas, B. (2001). Testing
for social learning in the “artificial fruit” processing of wildborn oran-
gutans (Pongo pygmaeus), Tanjung Putting, Indonesia. Animal Cogni-
tion, 4, 305–313.
Giraldeau, L. A. (1997). The ecology of information use. In J. R. Krebs &
N. B. Davies (Eds.), Behavioural ecology (4th ed., pp. 42– 68). Oxford,
England: Blackwell.
Hayes, K. J., & Hayes, C. (1952). Imitation in a home-raised chimpanzee.
Journal of Comparative Psychological Physiology, 45, 450 – 459.
Heyes, C. M. (1994). Social learning in animals: Categories and mecha-
nisms. Biological Reviews, 69, 207–231.
Heyes, C. M., & Ray, E. D. (2000). What is the significance of imitation
in animals? Advances in the Study of Behavior, 29, 215–245.
Horner, V., & Whiten, A. (2005). Causal knowledge and imitation/
emulation switching in chimpanzees and children. Animal Cognition, 8,
164 –181.
Miklósi, Á. (1999). The ethological analysis of imitation. Biological Re-
views, 74, 347–374.
Miles, H., Mitchell, R., & Harper, S. (1996). Simon says: The development
of imitation in an enculturated orangutan. In A. E. Russon, K. A. Bard,
& S. T. Parker (Eds.), Reaching into thought: The minds of the great
apes (pp. 371– 403). New York: Cambridge University Press.
Myowa-Yamakoshi, M., & Matsuzawa, T. (1999). Factors influencing
imitation of manipulatory actions in chimpanzees (Pan troglodytes).
Journal of Comparative Psychology, 113, 128 –136.
Myowa-Yamakoshi, M., & Matsuzawa, T. (2000). Imitation of intentional
manipulatory actions in chimpanzees (Pan troglodytes). Journal of Com-
parative Psychology, 114, 381–391.
Nagell, K., Olguin, R., & Tomasello, M. (1993). Processes of social
learning in the tool use of chimpanzees (Pan troglodytes) and human
children (Homo sapiens). Journal of Comparative Psychology, 107,
174 –186.
Piaget, J. (1962). Play, dreams and imitation in childhood. New York:
Norton.
Povinelli, D., & Deblois, S. (1992). Young children’s understanding of
knowledge formation in themselves and others. Journal of Comparative
Psychology, 106, 228 –238.
Russon, A. E., & Galdikas, B. M. F. (1995). Constraints on great apes’
imitation: Model and action selectivity in rehabilitant orangutan (Pongo
pygmaeus) imitation. Journal of Comparative Psychology, 109, 5–17.
Tomasello, M., Davis-Dasilva, M., Camak, L., & Bard, K. (1987). Obser-
vational learning of tool use by young chimpanzees and enculturated
chimpanzees. Human Evolution, 2, 175–183.
Tomasello, M., Kruger, A., & Ratner, H. (1993). Cultural learning. Behav-
ioural and Brain Sciences, 16, 495–511.
Whiten, A. (1998). Imitation of the sequential structure of actions by
chimpanzees (Pan troglodytes). Journal of Comparative Psychology,
112, 270 –281.
Whiten, A., Custance, D., Gomez, J., Teixidor, P., & Bard, K. (1996).
Imitative learning of artificial fruit processing in children (Homo sapi-
ens) and chimpanzees (Pan troglodytes). Journal of Comparative Psy-
chology, 110, 3–14.
Whiten, A., & Ham, R. (1992). On the nature and evolution of imitation in
the animal kingdom: Reappraisal of a century of research. Advances in
the Study of Behaviour, 21, 239 –283.
Whiten, A., Horner, V., & Marshall-Pescini (2003). Cultural anthropology.
Evolutionary Anthropology, 12, 92–115.
Zentall, T., R. (1996). An analysis of imitative learning in animals. In
C. M. Heyes & B. G. Galef (Eds.), Social learning in animals: The roots
of culture (pp. 221–238). San Diego, CA: Academic Press.
Received April 15, 2008
Revision received April 27, 2009
Accepted April 27, 2009 䡲