American J o u r n a l of Primatology 37:65-71 (1995)

Influence of a Skilled Model on the Behavior of

Conspecific Observers in Tufted Capuchin Monkeys
(Cebus apella)
LEAH ADAMS-CURTIS' AND DOROTHY M. FRAGASZY2
'Department of Basic Sciences, Uniuersity of Illinois College of Medicine at Peoria, Box 1649,
Peoria Illinois; 'Department of Psychology, University of Georgia, Athens, Georgia

We analyze how the presence of a skilled juvenile capuchin monkey inter-

acting with a mechanical puzzle requiring sequential actions affected the
behavior of group-mates towards the puzzle. Using this study a s a n exam-
ple, we suggest a methodological approach to the evaluation of social en-
hancement of activity and imitation. We suggest that this design could be
useful in determining if social and demographic factors influence the oc-
currence of these phenomena. o 1995 Wiley-Liss, Inc.

Key words: social learning, social facilitation, local enhancement,

imitation, problem solving, age differences

INTRODUCTION
Field data suggesting social influences on acquisition of unusual, locale-spe-
cific behavior among many species of animals are widely known, and enhancement
of interest in objects and places following observation of the behavior of conspe-
cifics toward that object or in that place (local enhancement) has been experimen-
tally documented in many species [Galef, 1988, 1990, 19921. It is less widely ap-
preciated, however, that experimental confirmation in animals of facilitative
social influences on performance or acquisition of specific behaviors is generally
weak and historically plagued by methodological constraints [Galef, 1988, 1992;
Visalberghi & Fragaszy, 1990a; Heyes, 19931.
New experimental approaches have appeared in studies with animals
[Palameta 1989; Palameta & Lefebvre 1985; Giraldeau & LeFebvre 1986, 1987;
Lefebvre 1986; Laland & Plotkin 1990, 1992; Heyes & Dawson, 19901 and with
human infants [Meltzoff 1988a,bl which do permit strong conclusions about obser-
vational influence on behavior. These studies include such improvements as
planned control groups and statistical comparisons between spontaneous levels of
the criterion behavior and the levels observed following modeling. However, of this
list, only Giraldeau and Lefebvre [1986, 19871 and Lefebvre [1986] studied indi-
viduals in groups. As pointed out by Laland et al. [19931, attention to the social
context of social learning has been minimal, and experimental methods for study-

Received for publication May 3, 1994; revision accepted January 3, 1995.

Address reprint requests to Dr. Dorothy M. Fragaszy, Department of Psychology, University of Georgia,
Athens, GA 30602.

0 1995 Wiley-Liss, Inc.

66 I Adams-Curtis and Fragaszy
ing the transmission of learned behaviors in groups of animals are still only poorly
developed.
The small sample size of this study limits the strength of our conclusions.
However, the study does illustrate the usefulness of using a puzzle to elicit behav-
ior, and the advantages of visual and relative physical isolation of the testing site.
The puzzle produced a n easily coded, finite sequence of common behaviors to com-
pare between model and observer, in distinction from time near the apparatus
which was also easily scored. Thus we could discriminate between local enhance-
ment of interest in the apparatus, and the production of specific sequences of acts
matching those of a skilled demonstrator. We also considered two independent
variables which might be responsible for inter-individual variance within the so-
cial group in the degree of observational influence on behavior directed toward the
experimental apparatus: age of the observer and availability of food.
METHODS
Subjects
The subjects were 12 juvenile and older members of a captive group of tufted
capuchin monkeys (Cebus apella; one adult male, five adult females, four 2-3-
year-old females, and two 1-2-year-old males). The group was housed indoors in
two adjoining rooms (6 x 2.5 x 2.5 m, and 3 x 2.5 x 2.5 m).
Apparatus
Two puzzle boards, 23 x 23 x 1cm, one painted black and one painted white,
were the test materials. One side of each board contained a series of three latches
and hasps that had to be opened in sequence. First the eye bolt (part 1)had to be
removed from the hasp. Next, the hasp (part 2) had to be opened, and last, the
hinge (part 3) had to be lifted. The other side of each board was painted, but
otherwise plain. A raisin was hidden under part 3 during the presentation of the
white puzzle. Presentations of the white puzzle are referred to hereafter as the F
condition (food available); and presentation of the black board a s the NF condition
(no food available). The animals had no previous experience with this type of
puzzle.
Procedure
A single puzzle board was presented by attaching it to a ledge outside of a test
area (33 x 58 x 53 cm) constructed of wire mesh. To manipulate the puzzle,
subjects reached through bars (.6 cm diameter; 5 cm apart) forming the outer wall
of the test area. The test area was accessible a t all times from the main portion of
the home cage through a doorway (20 x 30 cm). Three juveniles or two to three
adults could be present in the test area without excessive crowding. Subjects could
not clearly observe another animal manipulating the puzzle unless present in the
test area.
Each puzzle (F and NF) was presented for twelve 20-min sessions. A five-
minute presentation of the blank side of the puzzle board (the control condition)
occurred alternately before or after each test session. Presentation of both puzzles
(F and NF) occurred daily (separated by at least three hours) in balanced order
across time periods. All presentations of the puzzle and the control board were
videotaped. The experimenter remained outside the observation area, and watched
the proceedings on a video monitor. Each time the puzzle was solved, the experi-
menter entered the observation area, lowered a n opaque barrier, and reset the
puzzle (this took approximately 30 sec). Time spent by the experimenter in reset-
ting the puzzle was not considered part of the 20-min presentation time.
 Influence of a Skilled Model in Capuchins I 67
TABLE I. Percentage of Time Spent in Test Area*
Acquisition period for Q (%) Post-acquisition for Q (%)
Adults Juveniles Adults Juveniles
First 3 F with Q sessions 1 23 Last 3 F with Q sessions 2 9
(55) (29)
First 2 NF sessions 4 20 Last 10 NF sessions 2 24
(38) (52)
*Values in parentheses represent percentage of time that Q was present in which another juvenile was present
with Q.

The solver, Q, a juvenile female, was present during the first six F sessions (F
with Q). In order to determine the influence of Qs presence on the behavior of her
conspecifics, she was removed from the group during the last six F sessions (F
minus Q). Q was present for all N F sessions.

Analysis
Social influences. The following variables were derived per subject and con-
dition from videotapes: (1) time in the test area, (2) percent of time in the test area
in which Q was also present, (3) percent of time in the test area spent contacting
the puzzle.
Immediate effects of observation. Contacts with the puzzle by Q’syounger
male sibling, U, during the post-acquisition “F with Q” sessions were analyzed. U
was the only individual to contact the reset puzzle immediately several times after
having observed the previous solution and also several times when he had not been
present in the test area during the previous solution. Thus, only data from U were
available for these analyses. Each presentation in which U contacted the puzzle
outside of his attempting to grab the raisin as Q solved the puzzle (theft attempts)
was categorized by whether it immediately followed observation of solution (N =
11)or whether the prior solution had not been observed (N = 8).
Two-part sequences of contacts were chosen as the unit of analysis. This rule
eliminated scoring repeated rapid contacts to the same part of the puzzle as sep-
arate contacts. Contact was considered to have ended when the hand was moved
completely away from the puzzle board.

RESULTS
Level of Interest
The test area was popular during presentation of the puzzle, even when no food
was available. At least one animal was present in the test area for 70%(NF and the
“F minus Q ’ sessions) and 87% f“Fwith Q ’ sessions) of the presentation time. Each
individual spent about 12% of its time in the test area, and about 20% of its time
in the test area contacting the puzzle. Overall, the juveniles spent more time in the
test area and more time contacting the puzzle than did the adults (see Table I).
However, the two age-groups did not differ significantly across sessions for these
measures.
It appeared that the puzzle, by itself, was attractive. For example, with all
individuals considered, the percentage of time individuals contacted the test object
was greater when the puzzle was available (‘‘Fwith Q ’ sessions) than in associated
control sessions, in which the reverse side of the board was presented (M = 24%“F
with Q vs. M = 11%control, Wilcoxon Matched Pairs Rank Test T = 0, P < .01).
68 I Adams-Curtisand Fragaszy
Comparisons of other conditions (NF, F minus Q) with their associated control
conditions yielded a similar (nonsignificant) trend.

Local Enhancement
With all subjects considered, no influence of the model on the behavior of her
groupmates was evident: Comparisons of “F with &” sessions to “F minus Q” or to
“NF” sessions yielded no significance. However, even though there was no statis-
tically significant difference between age groups in the total amount of time at the
test site during “F with Q” sessions, juveniles tended to share the test area with Q
during a greater proportion of their time there in these sessions than did adults (M
= lo%, adults vs. M = 31%, juveniles, Mann-Whitney Test, U = 6, P = .062).
Moreover, one or more juveniles was present with Q during much of her time in the
test box in all conditions.
Effects of Observing Solution
There was no evidence that U learned, from watching Qs solutions, the se-
quences of contacts necessary to solve the puzzle, nor even that he associated a
particular part of the puzzle with the solution. The distribution of sequences of
contacts did not differ significantly depending on whether they followed observa-
tion of solution or not (Wilcoxon Matched Pairs Rank Test). Moreover, U did not
contact Part 3 more frequently following observation of the solution than in other
cases (31%of contact sequences following observation included part 3, vs. 52% of
sequences not following observation).
However, two aspects of U’s behavior are consistent with the occurrence of
local enhancement. First, 20 of U’s 26 theft attempts followed observation of the
solution on the previous presentation (x2 = 7.54 df = 1, P < .01). Second, U
contacted all parts of the puzzle at a higher rate following observation of solution
(14 contacts with any part of the puzzle during the next presentation following
observation of solution vs. 6 contacts with any part of the puzzle not following
observation. This difference approaches significance, x2 = 3.2, df = 1,P < .1.)

DISCUSSION
We focus our remarks upon two aspects of social learning which could poten-
tially have been expected to occur in this situation, in which one member of a group
repeatedly solved a mechanical puzzle, and consequently obtained a food reward.
These two aspects are enhancement (of interest in a place, or of contact with a n
object), and imitation (in this case, of a sequence of actions). Both of these are
aspects of social learning holding general interest, and the study of both is ham-
pered by methodological inconsistencies [Galef, 1988; Lefebvre & Palameta, 1988;
Tomasello et al., 19931.

Enhancement
With the caveat that our sample size was limited, and thus our results are
preliminary, it appeared that enhancement of interest in the test apparatus asso-
ciated with the immediate presence of the skilled individual was present in juve-
nile tufted capuchins but not in adults of the same group. Enhancement was
measured in this instance as percentage of time a t the test area and contacting the
puzzle. The findings suggest that age affects the expression of social enhancement
in species-normal situations in this species [as apparently in Japanese macaques;
Menzel 19661. Presenting the puzzle in a portion of the home cage with limited
access made determination of these patterns straightforward. Had we presented
 Influence of a Skilled Model in Capuchins / 69
the puzzle in the main portion of the home cage, these aspects of the group’s
behavior would have been much more difficult to ascertain.
It is often suggested that juveniles are more likely than adults to display
interest in others’ activities or possessions, and if this is true, they have better
opportunities than adults to learn from other group members [e.g., Cambefort,
1981; Kummer and Goodall, 19851. Three (non-exclusive) hypotheses can be pro-
posed to explain the greater propensity of juveniles to display enhancement. First,
juveniles may be generally more attentive to the activities of others than are
adults-an intrinsic attentional difference across ages. Second, they may be better
able to act upon their inclinations t o approach another individual, as a conse-
quence of the social dynamics in the group involving tolerance towards younger
individuals. Finally, juveniles may be a more cohesive subgroup than adults under
all circumstances (leading to greater probability of enhancement with any juvenile
demonstrator, as Q was, all other things being equal). Cambefort [1981]posited the
third explanation for the rapid dissemination of a novel food habit in juvenile
baboons. The first two seem at least as likely to be responsible for the variations in
enhancement seen in capuchins. Juvenile capuchins looked more reliably at an-
other animal than did adults in a recent study in our laboratory (Johnson, unpub-
lished data), in line with the first hypothesis. With regard to the second hypothesis,
it has been found that infants less than a year old are allowed virtually uncon-
tested access to objects, even desirable food objects, handled by adults [Mitra et al.,
19933. On the other hand, enhancement of activity a t specific sites associated with
the activity of adults is differentially evident across juvenile capuchins as a func-
tion of the age and identity of the juvenile [Fragaszy et al., 19941. The differenti-
ation is apparently related to the increasing role that specific social status rela-
tionships play in behavior as juveniles mature.
Patterns of social spacing and regulation of social relationships undoubtedly
also play a role in the extent to which enhancement occurs. For example, if kin are
more likely to be near one another, then this is an obvious route for enhancement.
This may in part account for the enhancement observed in U (as the skilled indi-
vidual was his sister). The power or attractiveness of others near the skilled indi-
vidual may also block or facilitate enhancement, depending upon the particular
relationships among all the individuals. Again, restricted access to the test appa-
ratus makes these components of social enhancement easier to detect. Overall,
enhancement of activity is more likely among some members of a group than
others. Clearly more enhancement, and more frequent social transmission of in-
formation in general, can be expected in species characterized by relaxed and
tolerant social interactions among group members [Coussi-Korbel, 19931.
Immediate Effects of Observing Solution
Analysis of one individual’s contacts with the puzzle revealed no effect of
observation of solutions on the topography of his behaviors with the puzzle. The
negative finding is in accord with other recent experimental studies seeking to
document imitation in capuchins [Fragaszy & Visalberghi 1989,1990; Visalberghi
& Fragaszy, 1990a,b; Fragaszy et al., 19941. Capuchin monkeys do not learn how
to use a tool from watching each other, nor how to wash sand from food or grain.
However, where social relations support this (i.e., among matrilineal kin), they
may display great tolerance, co-action, and “scrounging.” Scrounging may inter-
fere with social transmission of food finding, as has been shown in studies with
pigeons [Giraldeau & LeFebvre, 19871. Scrounging was observed in this study, by
Q’s brother and mother.
We adopted a task that we know to be within the capacity of all individuals to
70 I Adams-Curtis and Fragaszy
perform-lifting and pulling are part of the manipulative repertoire of all capu-
chins. However, the specific sequences of actions to particular targets are, in their
totality, novel. Thus we could be sure that the lack of imitation in our study was
not a function of the difficulty of the motor components of the task.
The negative finding on imitation of specific acts by nonhuman primates in
this study and in others should not undermine appreciation of the potentially
significant role of social learning in these animals in natural settings, and of the
significance of such learning to adaptive behavioral change at the population level
[Laland et al., 19933. Laboratory studies should focus on social influences on the
transmission of learned behaviors in ways that inform us about the contributions
of social learning to behavior of group-living animals [Laland et al., 1993; Coussi-
Korbel and Fragaszy, in press].
Methodological Issues
We have suggested that limited access to the test site was a n advantage in this
study, because we could note accurately what actions each individual could have
seen done and did itself with the puzzle. Limited access is a n intermediate condi-
tion between the traditional test paradigm of separated demonstratorlobserver
dyads and the more naturalistic setting of simultaneous access to a test site by a n
entire group. Second, limited access ensured tha t we could note which individuals
were present with which others. Records of specific pairings at the site may be used
to address the contribution of social dynamics to opportunities for social learning
in groups, particularly if the data are coupled with independent assessments of
social relationships outside the test situation.

CONCLUSIONS
1. Age is potentially a n important factor in the occurrence of local enhance-
ment in capuchin monkeys, because of its probable relation to attention andlor
access to others and the sites of others’ activities.
2. Juveniles, but not adults, exhibited behaviors consistent with the occur-
rence of local enhancement.
3. A sequential task provides a clear basis for detecting increased matching of
sequences of actions following observation vs. in other conditions. The increase is
one means of detecting imitation.
4. There was no evidence of imitation of action sequences.

ACKNOWLEDGMENTS
This work was supported by NSF grant BNS85-3603 and grant MH41543 and
a Research Scientist Development Award from the National Institute of Mental
Health to D. Fragaszy. We thank Rena Carlson-Lammers for her assistance with
data collection and Jim Whipple for his assistance with data analysis. J im Whip-
ple, Roger Davis, Sabine Coussi-Korbel, and a n anonymous reviewer provided
useful comments on a n earlier version of this manuscript. The data collection was
completed a t Washington State University, Pullman, WA.
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