Cultural Variation in the Use of Overimitation by the Aka and Ngandu of the Congo Basin
Cultural Variation in the Use of Overimitation by the Aka and Ngandu of the Congo Basin
Richard E. W. Berl 0 1
Barry S. Hewlett 0 1
0 1 School of Biological Sciences, Washington State University , Pullman, Washington , United States of America, 2 Department of Anthropology, Washington State University Vancouver , Vancouver, Washington , United States of America
1 Academic Editor: Kenny Smith, University of Edinburgh, School of Philosophy, UNITED STATES
Studies in Western cultures have observed that both children and adults tend to overimitate, copying causally irrelevant actions in the presence of clear causal information. Investigation of this feature in non-Western groups has found little difference cross-culturally in the frequency or manner with which individuals overimitate. However, each of the non-Western populations studied thus far has a history of close interaction with Western cultures, such that they are now far removed from life in a hunter-gatherer or other small-scale culture. To investigate overimitation in a context of limited Western cultural influences, we conducted a study with the Aka hunter-gatherers and neighboring Ngandu horticulturalists of the Congo Basin rainforest in the southern Central African Republic. Aka children, Ngandu children, and Aka adults were presented with a reward retrieval task similar to those performed in previous studies, involving a demonstrated sequence of causally relevant and irrelevant actions. Aka children were found not to overimitate as expected, instead displaying one of the lowest rates of overimitation seen under similar conditions. Aka children copied fewer irrelevant actions than Aka adults, used a lower proportion of irrelevant actions than Ngandu children and Aka adults, and had less copying fidelity than Aka adults. Measures from Ngandu children were intermediate between the two Aka groups. Of the participants that succeeded in retrieving the reward, 60% of Aka children used emulation rather than imitation, compared to 15% of Ngandu children, 11% of Aka adults, and 0% of Western children of similar age. From these results, we conclude that cross-cultural variation exists in the use of overimitation during childhood. Further study is needed under a more diverse representation of cultural and socioeconomic groups in order to investigate the cognitive underpinnings of overimitation and its possible influences on social learning and the biological and cultural evolution of our species.
Funding: Work by REWB was funded through a
fellowship with the Washington State University NSF
IGERT Program in Evolutionary Modeling (http://
ipem.anth.wsu.edu/). Work by BSH was funded by
the Japan Society for the Promotion of Science (PI:
Hideaki Terashima, Kobe Gakuin University; https://
www.jsps.go.jp/english/). Funding for publication was
provided by the College of Arts and Sciences at
Washington State University, Vancouver (http://cas.
vancouver.wsu.edu/). The funders had no role in
Human behavior is the result of a complex coevolutionary history written by the network of
interactions between our biology, our culture, and our environment [1, 2]. From this rich history
study design, data collection and analysis, decision to
publish, or preparation of the manuscript.
Competing Interests: The authors have declared
that no competing interests exist.
has arisen a great deal of behavioral variability both within and between human groups .
That many aspects of behavior in humans and other animal species are not innate but socially
learned from parents, other adults, or peers means that the acquisition of behavior is highly
contingent upon an individuals sociocultural environment. Further, enculturation begins in
infancy and early childhood , possibly even in utero , suggesting that there is no such
thing as a culture-less child. Hence, it is unlikely that any socially learned behavior or the
cognitive mechanisms underlying its transmission and acquisition are ever unaffected by an
individuals particular culture and ontogeny.
These concerns underscore the importance of cross-cultural comparative research in
psychological, anthropological, and biological lines of inquiry that address human behavior and
cognition, particularly those concentrating on social learning, cultural transmission, and
associated mechanisms that fall under the study of cultural evolution . This need is especially
crucial given that often the primary focus of these studies is to search for core mechanisms and
processes that are universal to modern humans. It is surprising, then, that very few studies
have included non-Western samples in their analyses. Undergraduate university students
enrolled in psychology courses constitute the sole sample for 67% of psychological studies based
in the United States and 80% of studies elsewhere . Only 18% of samples from leading
psychology journals in the mid-2000s were from non-English speaking countries, with the
majority of those non-English samples coming from highly industrialized countries in Western
Europe. This has left 95% of the research effort concentrated on a heavily biased subsample
drawn from just 12% of the worlds population . It is thus inappropriate to attempt to
generalize findings from this restricted sample of Western people to any non-Western groupor,
indeed, to other Western cultures or to sub-cultures with different socioeconomic conditions
without additional sampling sufficient to explore the breadth of cross-cultural variation .
Much recent interdisciplinary effort in social learning research has been focused on by what
means and under which conditions humans and other animal species employ a number of
possible social learning mechanisms and strategies , with the most basic of these being
imitation and emulation. Following Whiten et al. , we refer here to the broad definitions of
imitation and emulation, with imitation as a focus by the observer on the reproduction of the
form of modeled actions rather than the result or goal of those actions and, conversely, with
emulation as a focus on reaching the modeled outcome rather than on the details of the actions
that accomplish that outcome. The phenomenon of overimitationthough the term was
coined by Lyons et al. was first observed by Horner and Whiten  in a study
investigating whether children (from two nursery schools in the United Kingdom) and captive
chimpanzees could switch between imitative and emulative mechanisms given the availability of
visual information about causal relationships. In the experiment, a model demonstrated a
series of actions by which a reward could be retrieved from an artificial apparatus. Some of the
demonstrated actions were causally relevant and necessary to obtain the end reward, but others
were shown to be irrelevant and could be omitted by using emulation, thereby increasing the
efficiency of the procedure and reaching the reward more quickly. They found that the
chimpanzees did so but that the childreneven in the presence of visual information about the
causality of their actionscontinued to imitate the irrelevant demonstrated actions rather than
switching to an emulative approach. Overimitation is thus defined here as the high-fidelity
copying of causally irrelevant actions in the presence of clear causal information.
Subsequent studies have observed overimitation in Western children under a variety of
experimental conditions. It appears to develop between the ages of 18 months and three years
, to increase through ages three to five [20, 21, 2527], and is observed at even higher
rates in adults [28, 29]. Whether it persists through adolescence or re-emerges in early
adulthood is presently unknown due to the lack of studies in this developmental period. Curiously,
overimitation appears to be largely absent in children up to at least 18 months [23, 30, 31] and
different social or contextual cues can trigger selective copying at the same ages in which
overimitation is observed (for a review, see ). Many of these studies have attempted to tie
overimitation into current theory on the cultural and biological evolution of our species, as a
nonselective copying strategy has the potential to facilitate rapid adoption of the vast amount of
causally opaque cultural knowledge available during childhood [27, 28, 3340]. Nielsen and
colleagues have suggested that overimitation is a universal human trait ( (p. 729), 
(p. 156),  (p. 171)).
To date, there have been only two studies of overimitation conducted with non-Western
groups. Nielsen and Tomaselli  investigated overimitation in the !X, Khomani, !Xun,
and Khwe San peoples of Botswana (!X) and South Africa (Khomani, !Xun, and Khwe),
while Nielsen et al.  extended this research to two groups of Aboriginal Australians, the
Yanyuwa and the Garrwa in the Northern Territory, and performed additional work with the
San groups from the previous study. Both studies found little difference between the
indigenous populations and a group of Western children in Brisbane, Australia in the frequency or
manner with which children overimitate.
However, there is good reason to believe that the San and Aboriginal Australians involved in
these studies are no longer actively engaged in a way of life typical of hunter-gatherers or other
small-scale cultures. Robins  describes the present-day Khomani San as a group of
superexploited and hypermarginalized ex-farm workers. The San people are a loose affiliation of
indigenous groups that have had continuous exposure to Western ideas, values, and technology for
many decades, and this life beside and within Western society has fundamentally altered their
respective cultures . In all cases, the San have been made to adapt to political, social,
economic, and religious pressures exerted by Western influences (see S1 Appendix). Aboriginal
Australian groups have been subjected to similarly intense pressures throughout their history of
contact and conflict with Western cultures .
In considering these two non-Western studies and a third incorporating a sample of
children from Colombo, the urban, industrialized de facto capital of Sri Lanka , each additional
cultural group provides valuable data toward the examination of overimitation and other social
learning processes and broadens cultural and socioeconomic representation in this line of
research. With this being said, we agree with the assessment of Whiten  that it is
inappropriate at present to claim human universality of a trait that has only been observed in such a
narrow span of the worlds cultures. Speculation on the significance of overimitation in the
evolution of human cumulative culture has been similarly overextended beyond the limits of
our knowledge of its underlying causal mechanisms.
In an effort to investigate overimitation in a context with limited Western cultural
influences, we conducted a study with the Aka hunter-gatherers and the neighboring Ngandu
horticulturalists of the Congo Basin rainforest in the southern Central African Republic. The Aka
(also referred to as the Biaka or Bayaka) are a group of foragers that express the set of pygmy
phenotypic traits presumed to be adaptive for life in the tropical rainforest . About
40,000 Aka inhabit the equatorial rainforests of the northern Republic of the Congo and
southern Central African Republic, with about 2,000 living in and around the study area. The Aka
live in highly mobile groups of 2535 people, typically made up of families related by kinship
and marriage, which move camp several times per year. They actively rely upon a wide variety
of hunting and gathering techniques for their day-to-day subsistence.
The Ngandu are a group of Bantu-speaking farmers that live in villages of 50200
individuals and cultivate manioc, corn, plantains, and peanuts. They exchange some of their
domesticated crops for meat and other forest products from the Aka. Women plant, maintain, and
harvest the fields and provide the majority of dietary calories while men fish, hunt, and trade.
About 15,000 Ngandu live mostly within the Central African Republic, with about 4,000 living
in and around the study area.
The Aka in this study have complex economic, ritual, and kinship relationships with the
Ngandu [4, 52] similar to those seen in many central African hunter-gatherer groups associated
with farmers [53, 54]. The Aka and Ngandu speak closely-related Bantu languages (Ethnologue
identifiers axk and ngd, respectively) , though a portion of the Aka language in certain
domains is shared with other foragers in the Congo Basin . The Aka are in close contact with
the Ngandu but, because the area as a whole is relatively isolated due to the lack of local
infrastructure, the Aka themselves have limited exposure to Western influences, do not live under
socio-economic poverty, receive no government assistance or subsidy, and maintain their
foraging culture and social norms with relatively few outside pressures ([46, 50, 53] and see
Given that overimitation has been observed in children from every culture examined thus far,
it was predicted that Aka and Ngandu children also would overimitate. However, few studies
have focused on children from cultures that are not Western, educated, industrialized, rich, and
democratic (WEIRD ). This makes definitive predictions difficult, as there are a number of
foundational differences between WEIRD cultures and small-scale non-Western ones that
structure their perceptions and values and lead to variation in behavior ( and see S2 Appendix on
the Aka and Ngandu). Additionally, though the Ngandu are largely non-Western and have daily
contact with the Aka, the Ngandu receive formal education through schooling (see S1 Appendix)
and emphasize respect for elders and those of higher status (see S2 Appendix), which could be
expected to generate differences between Aka children and Ngandu children in their approaches
to social learning.
All participants provided verbal informed consent to take part in the study after having been
read and acknowledging that they understood an approved consent script. Adults provided
consent to the researcher in the presence of a witness, as did participating children and their
parents. Written consent could not be obtained because most of the participants were not
literate. Research protocols, the use of verbal consent, and verbal consent scripts were approved by
the Washington State University Institutional Review Board (protocol #12344).
Three groups were recruited for the study, consisting of 28 Aka children, 29 Ngandu children,
and 14 Aka adults. Ages ranged from 4 to 7 years in Aka children (M = 5.4, SD = 1.2), from 4
to 7 years in Ngandu children (M = 5.3, SD = 0.9) and from 20 to 38 years in Aka adults (M =
28.4, SD = 6.0). Sex representation was equal or approximately equal in all groups (Aka
children: 14M, 14F; Ngandu children: 14M, 15F; Aka adults: 7M, 7F). Ngandu adults could not be
sampled due to field research constraints and ongoing political unrest in the Central African
The study apparatus was a transparent polycarbonate box (see Fig. 1), measuring roughly
15 cm x 15 cm x 15 cm and similar to those used by Horner and Whiten , McGuigan et al.
, McGuigan and Whiten , and McGuigan et al. , differing only in the top door
mechanism. Two holes (each 2 cm x 2 cm) allowed entry into the box, one on the top side and
Fig 1. Experimental apparatus and tool. Used for all participants and tests in this study. Relevant features
include: (A) sliding top door; (B) transparent barrier (below the plane of the top side of the apparatus); (C)
opaque reward chamber; (D) sliding/lifting front door; and (E) manipulating tool (shown to scale with
apparatus). See Methods for details of construction and use.
one on the front side. The front hole was covered by a door with a small knob and allowed
entry by sliding the door to either side or by lifting it upward and outward. The top hole was
covered by a sliding door with an open notch and could be slid only to the left. A 22-cm-long
aluminum tool with two short knobs protruding from one end and a long flat section on the
opposite end was used to interact with the box and retrieve the rewarda small piece of bubble
gumwhich was stored in an opaque black rectangular prism connected to the inside of the
front hole. The top hole led to a chamber with a transparent barrier separating it from the rest
of the box and the reward. Thus, demonstrated actions performed on the top or sides of the
box were causally irrelevant to obtaining the reward. Opening the front door and obtaining the
reward were judged to be causally relevant.
Participants in each group were randomly assigned to receive a demonstration or to receive no
demonstration, such that roughly one third of the available participants in each group were
assigned to the no demonstration condition (Aka children: N = 9; Ngandu children: N = 9; Aka
adults: N = 4). In the demonstration condition, participants each observed an ingroup adult
male model as he performed a sequence of actions on the box which resulted in the retrieval of
the reward. Participants in the no demonstration condition had the box and tool placed in
front of them without any actions demonstrated by the model. The Aka and Ngandu models
were both members of the local community, in their late thirties, and had been prior research
assistants for the second author. They were known to the participants but not familiar to them
and were not close relatives of any participant in the study. Neither held a position of particular
status within the community.
Tests were conducted in locations familiar to the participants. Aka children and adults were
tested in a camp house or on a nearby forest trail and Ngandu children were tested in a village
home. The participant sat next to the model, with the apparatus and tool placed in front of and
midway between the model and participant. In the demonstration condition, the model said
Watch me, and demonstrated the complete sequence three times, reloading the box out of
view of the participant after each demonstration. He then placed the box and tool in front of
the participant, said Your turn, and did not interact with the participant for the remainder of
The demonstrated sequence consisted of 6 actions, with the first 4 being causally irrelevant
and the final 2 being causally relevant to the goal of obtaining the reward: tap the right side of
the box (TR, irrelevant), tap the left side of the box (TL, irrelevant), slide the top door
(ST, irrelevant), tap the barrier inside the top door (TT, irrelevant), slide the front door
(SF, relevant), and retrieve the reward (RR, relevant). Thus, the complete demonstrated
sequence in order was: TR, TL, ST, TT, SF, RR. Participants were given up to one minute to
interact with the apparatus and, if they did so, two minutes to retrieve the reward.
In the no demonstration condition, after the box and tool were placed in front of and
between the model and participant, the model said, Can you find the treat? and did not interact
with the box while feigning disinterest for the rest of the test. Participants in the no
demonstration condition were prompted with this phrase in order to provide context for the task as a
puzzle, which would otherwise be absent given that the highly artificial nature of the experimental
apparatus was incongruent with that of a naturalistic foraging problem. Circumstances were
otherwise identical to the demonstration condition.
All tests were recorded on video for coding and review. Trials were coded in randomized
order. Coding of tests included only the six demonstrated actions; other actions were noted for
the purpose of determining copying fidelity but were not coded as relevant or irrelevant due to
their inherent ambiguity of purpose.
From the coded datain addition to the number of irrelevant actions performedthree other
measures were calculated for each test. First, an irrelevant imitation score [21, 22, 26, 28, 57]
was generated by dividing the number of tool or hand insertions into the top irrelevant hole (TT)
by the total number of insertions into both the top irrelevant and front relevant holes (TT + RR).
This measurement ranges from 0, meaning that all insertions were relevant, to 1, meaning that
all insertions were irrelevant. Trials where no insertions took place and, by consequence, the
reward was never retrieved could not be given an irrelevant imitation score.
Second, an irrelevancy quotient was determined by dividing the total number of irrelevant
actions (TR + TL + ST + TT) by the total number of irrelevant and relevant actions (TR + TL +
ST + TT + SF + RR), and thus is simply the proportion of actions performed by the participant
that were irrelevant. Participants that did not engage with the task and therefore did not
perform any actions on the box were not given an irrelevancy quotient. This measure was
calculated in order to represent the full proportion of irrelevant actions performed by the
participant rather than limiting observations to insertions into either hole on the box, which was
originally done due to methodological constraints 
A proportional measure such as the irrelevant imitation score or irrelevancy quotient is
necessary because strictly examining the number of irrelevant actions does not account for
variance in the total number of actions performed. An equal number of irrelevant actions could be
obtained from a situation in which a few irrelevant actions were performed along with a few
relevant ones, or from one in which a great number of actions were performed and a small
proportion of them were irrelevant. The irrelevant imitation score and irrelevancy quotient take
this issue into account by providing measures of the relative irrelevancy of the full complement
of actions performed.
The irrelevancy quotient also allows for comparison with studies that may not have one
primary irrelevant action suitable to be paired with one relevant reward retrieval action, as in the
irrelevant imitation score. For a more rigorous comparison between studies, an adjustment
should be made to standardize irrelevancy quotients by the proportion of irrelevant actions in
the demonstrationwhich in this case is 4 out of 6, or two thirdsbut obtaining the raw data
and calculating this for each presently available study is beyond the scope of this paper.
Third, a fidelity quotient was calculated by first determining a fidelity score, defined as the
longest string of actions performed in the same order as the demonstration. For example, a
trial consisting of the actions TR, TL, ST, SF, RR would have a fidelity score of 3, while the
actions TR, TL, SF, RR would have a fidelity score of 2 for the particular demonstrated sequence
of this study (TR, TL, ST, TT, SF, RR; as above). Higher fidelity scores indicate greater attention
to the imitation of sequential structure, which is useful in distinguishing imitation from other
transmission mechanisms [58, 59]. The fidelity score is then divided by the maximum possible
fidelity score, i.e. the length of the demonstrated sequence (6 for this study), to obtain the
fidelity quotient. The calculation of a fidelity quotient yields an easily-determined quantitative
measure of the level of copying fidelity given a short sequence of demonstrated actions. For
experiments in which the demonstrated sequence contains irrelevant actions, the fidelity
quotient complements the irrelevancy quotient in assessing overimitation, since overimitation (as
defined for this study) requires both the reproduction of irrelevant actions and high copying
fidelity. As with the irrelevancy quotient, a fidelity score or quotient could not be assigned for
trials in which no actions were performed on the box.
The social learning mechanism employed by each participant was determined based upon
the actions exhibited following the demonstration. As in previous studies, the use of one or
more irrelevant actions that matched those demonstrated constituted imitation, whereas the
exclusion of these irrelevant actions qualified as emulation [21, 22, 26]. Participants that
ignored the apparatus or did not complete the test could not be assigned a social learning
mechanism unless an irrelevant action matching the demonstration had been used.
As an additional measure of interest, the duration of each test was timed from the first
action performed on the box by the participant (after any demonstrations) to the time when the
reward was successfully retrieved. No duration was recorded for participants that did not
participate or failed to retrieve the reward. This measure was recorded, as in McGuigan et al. 
and McGuigan and Whiten , to investigate if either social learning mechanism led to a
higher task efficiency.
An independent observer unfamiliar with the experimental hypotheses coded 15 trials,
selected at random from the total 58 in which participants engaged with the test, representing
26% of the total sample. The observer and experimenter agreed on the specific coding of
observed actions to a high degree (81%, Cohens = .778).
Univariate factorial ANOVA tests were used to examine the main effects and interactions of
demonstration condition (demonstration or no demonstration), group (Aka children, Ngandu
children, or Aka adults), and sex (male or female) on each dependent variable: the number of
irrelevant actions, irrelevant imitation scores, irrelevancy quotients, and fidelity quotients.
Participants in the demonstration condition were found to exhibit a higher number of
irrelevant actions than participants in the no demonstration condition (F[1, 45] = 4.138, p = .048,
p2 = .084, 95% CI [.275, 2.387]; see Fig. 2). Multiple comparisons were unable to make
distinctions between conditions within groups due to the low number of participants in the no
demonstration condition that engaged with the experiment (Aka children: N = 4; Ngandu
children: N = 6; Aka adults: N = 3). Irrelevant imitation scores and irrelevancy quotients did not
vary by condition (95% CIs [.120, .233] and [-.411, .110], respectively) while fidelity quotients
were lower in the no demonstration condition (F[1, 51] = 18.822, p << .001, p2 = .270, 95%
CI [.198, .539]). Sex was not a significant predictor for any measure and no significant
interaction effects were found.
Fig 2. Number of irrelevant actions by group and demonstration condition. Bars indicate sample means
and 95% confidence intervals, computed through basic nonparametric bootstrapping. Bubbles show the
distribution of values within each sample; the size of each bubble represents the relative proportion at
For methodological comparison, Welchs t-tests and Pearsons chi-squared tests were
conducted as in Nielsen and Tomaselli  and Lyons et al. , respectively, on the number of
irrelevant actions, irrelevant imitation scores, irrelevancy quotients, and fidelity quotients
collapsed across groups. Results of these tests agreed qualitatively with findings from the
ANOVA F-tests for all t-tests (number: t[19.538] = 3.226, p = .004; fidelity: t[52.932] = 6.258,
p << .001) and chi-squared tests (fidelity: 2 [1, N = 56] = 5.628, p = .018). The chi-squared
test of the number of irrelevant actions became equivalent to that of the irrelevancy quotient
due to both being converted to count data.
Subsequent analyses below pertain solely to participants within the demonstration
condition (Aka children: N = 19; Ngandu children: N = 20; Aka adults: N = 10). Full ANOVA tables
for all analyses, including confidence intervals and effect sizes for significant and nonsignificant
effects, are available in the supporting information (see S1 Table; S2 Table; S3 Table).
Reproduction of Irrelevant Actions
Following the control analyses on demonstration condition, univariate factorial ANOVA tests
were used to examine the main effects and interactions of group (Aka children, Ngandu
children, or Aka adults) and sex (male or female) on each measure of irrelevancy: the number of
irrelevant actions, irrelevant imitation scores, and irrelevancy quotients. ANCOVA was used to
test for differences by group (Aka children or Ngandu children) due to the effects of participant
age (continuous) on these same measures, excluding Aka adults to avoid confounding group
The number of irrelevant actions performed by each participant was found to vary by group
(F[2, 37] = 3.706, p = .034, p2 = .167; see Fig. 2), with post-hoc Tukeys HSD confirming that
Aka children performed fewer irrelevant actions than Aka adults (p = .026, 95% CI [.243,
4.382]), while other groupwise comparisons were nonsignificant (Aka children and Ngandu
children: p = .402, 95% CI [-.819, 2.694]; Ngandu children and Aka adults: p = .249, 95% CI
[-.695, 3.445]). There was no difference by sex in the number of irrelevant actions performed
and no difference was attributable to the age of the participant within or between the two
groups of children. All interaction terms were found to be nonsignificant.
Irrelevant imitation scores differed between groups (F[2, 35] = 4.501, p = .018, p2 = .205;
see Fig. 3), with Aka children having lower scores than both Ngandu children (Tukeys HSD,
p = 0.034, 95% CI [.017, .516]) and Aka adults (Tukeys HSD, p = 0.035, 95% CI [.019, .596]),
but with no difference between Ngandu children and Aka adults (Tukeys HSD, p = .936,
95% CI [.248, .329]). No differences were found by participant sex or age and all interaction
terms were found to be nonsignificant.
Irrelevancy quotients were also found to differ between groups (F[2, 37] = 3.332, p = .036,
p2 = .153; see Fig. 3), but post-hoc tests were not conclusive (Tukeys HSD, Aka children and
Ngandu children: p = .077, 95% CI [.022, .528]; Aka children and Aka adults: p = .078, 95%
CI [.027, .622]; Ngandu children and Aka adults: p = .940, 95% CI [.280, 0.369]). No
differences were found by participant sex or age and all interaction terms were found to
To then examine copying fidelity, a univariate between-groups factorial ANOVA was used to
examine the main effects and interactions of group (Aka children, Ngandu children, or Aka
adults) and sex (male or female) on fidelity quotients. ANCOVA was used to test for
differences by group (Aka children or Ngandu children) due to the effects of participant age
(continuous) on fidelity quotients, again excluding Aka adults.
Fig 3. Measures of irrelevancy and fidelity by group. Irrelevant imitation scores and irrelevancy quotients represent proportions of irrelevancy and fidelity
quotients represent copying fidelity (see Analysis for descriptions). Bars indicate sample means and 95% confidence intervals, computed through basic
nonparametric bootstrapping. Bubbles show the distribution of values within each sample; the size of each bubble represents the relative proportion at
Fidelity quotients varied by group (F[2, 39] = 6.268, p = .004, p2 = .243; see Fig. 3), such
that Aka children had lower scores than Aka adults (Tukeys HSD, p = .003, 95% CI [.124,
.690]) and Ngandu children had lower scores than Aka adults (Tukeys HSD, p = .038, 95% CI
[.014, .592]) while there was no difference seen between Aka children and Ngandu children
(Tukeys HSD, p = .541, 95% CI [.134, .342]). No differences were found by participant sex or
age and all interaction terms were found to be nonsignificant.
Social Learning Mechanism
Only participants that succeeded in obtaining the reward (Aka children: N = 15; Ngandu
children: N = 13; Aka adults: N = 10) were analyzed in terms of the social learning
mechanism used. The proportion of individuals that succeeded by use of emulation rather than
imitation differed by group (Fishers Exact, p = .017; see Fig. 4), with Aka children using
emulation more often than Ngandu children (Fishers Exact, p = .024) and more often than
Aka adults (Fishers Exact, p = .033) with no difference seen between Ngandu children and
Aka adults (Fishers Exact, p = 1.000). The use of emulation was a significant predictor of
the number of irrelevant actions ( = 3.720, t(35) = 7.435, p << .001), of irrelevant
imitation scores ( = .347, t(35) = 4.867, p << .001), of irrelevancy quotients ( = .614,
t(35) = 15.870, p << .001), and of fidelity quotients ( = .377, t(35) = 4.268, p < .001). A
significant proportion of the variance in the number of irrelevant actions (r2 = .601, F(1, 35) =
55.28, p << .001), irrelevant imitation scores (r2 = .387, F(1, 35) = 23.68, p << .001),
irrelevancy quotients (r2 = .874, F(1, 35) = 251.7, p << .001), and fidelity quotients (r2 = .324,
F(1, 35) = 18.22, p < .001) was explained by the use of emulation. No differences were
found by participant sex or age and all interaction terms were found to be nonsignificant.
These results were compared with data from overimitation studies by McGuigan et al. 
and McGuigan and Whiten , which report the social learning mechanisms used by toddlers
Fig 4. Use of emulation by group. Bars indicate the proportion of those participants that succeeded at
obtaining the reward that did so using an emulative social learning mechanism, wherein the copying of one or
more irrelevant actions matching the demonstration constituted imitation and the copying of zero matching
irrelevant actions constituted emulation (see Introduction and Analysis). Error bars indicate 95% confidence
intervals, computed using the Clopper-Pearson (exact) method. Frequencies of the use of emulation by
Western toddlers and children from the United Kingdom are included for comparison. Data for these groups
are from overimitation studies by McGuigan et al.  and McGuigan and Whiten  and are summarized in
Table 1. The two age classes within each Western group are collapsed here for clarity.
and children from nursery schools and parent-toddler groups in the United Kingdom (see
Fig. 4; Table 1). Aka children displayed levels of emulation comparable with 23-month-old
and 30-month-old Western toddlers, but emulated at much higher frequencies than
42distinguished from any of these Western groups. At a mean age of 5.4 years, or roughly 65
months, the Aka children that succeeded at the task were much older than the comparably
emulating toddler groups (see Table 1). Though a direct comparison cannot be made
between the Aka and Ngandu participants and the San studied by Nielsen and colleagues [27,
40], the San participants were of ages that ranged both younger and older than the Aka
chilSD = 3.1, range = 213; : M = not reported, SD = not reported, range = 36).
Confidence limits of the proportions of groups using emulation were calculated using the
Clopper-Pearson (exact) method. For statistical tests, Fishers exact test and the
Holm-Bonferroni method were used to obtain p-values corrected for multiple comparisons. Ages are given
in years as precision beyond half a year was not known for Aka and Ngandu participants. Data
from Western toddlers and children in the United Kingdom were obtained from McGuigan
et al.  and McGuigan and Whiten . Western toddlers and children were classified by
their dominant approach, as in the source texts.
The length of time each participant spent interacting with the apparatus was significantly
predicted by the number of irrelevant actions performed ( = 3.984, t(34) = 3.805, p < .001), by
irrelevancy quotients ( = 27.654, t(34) = 3.564, p = .001), and by decreasing use of an emulative
social learning mechanism ( = 16.625, t(34) = 3.186, p = .003), but not by irrelevant
imitation scores or fidelity quotients. The number of irrelevant actions, irrelevancy quotients, and
social learning mechanism were found to explain a significant proportion of the variance in
test duration (number: r2 = .278, F(1, 34) = 14.48, p < .001; irrelevancy: r2 = .251, F(1, 34) =
12.70, p = .001; mechanism: r2 = .207, F(1, 34) = 10.15, p = .003), while irrelevant imitation
scores and fidelity quotients did not. The participants sex was found to have no effect on test
duration and no significant interactions were found. Participant age was a significant predictor
of test duration among Aka children and Ngandu children, with older ages associated with
longer times ( = 7.443, t(25) = 2.176, p = .039) and with age explaining a significant proportion of
the variance in test duration (r2 = .126, F(1, 25) = 4.736, p = .039), but no difference was found
between these groups and there was no significant interaction between age and group.
In order to infer a difference between groups in the tendency to overimitate, it must first be
determined at what rates the demonstrated irrelevant actions would be produced in the absence
of a demonstration and whether these rates differ. Bias could exist if the actions were somehow
intuitive to the task or were themselves affected by cultural, developmental, or environmental
factors. Our results suggest that this is not the case (see Fig. 1). Participants that received a
demonstration displayed a greater number of irrelevant actions than those that did not receive
a demonstration, indicating that some degree of social learning took place. That there were no
significant differences between groups within the no demonstration condition suggests that
such differences either do not exist or were undetectable due to the small sample of individuals
that chose to engage with the box in the no demonstration condition. Notably, few of the
irrelevant actions observed in the no demonstration condition matched those modeled in the
demonstration. This suggests that participants that did not receive a demonstration were merely
trying to determine the function of the box through exploration or play. These results indicate
that the irrelevant actions exhibited by participants exposed to a demonstration were copied
from that demonstration by means of imitation rather than the result of individual learning or
In comparing the reproduction of irrelevant actions between Aka children and Aka adults,
children copied fewer irrelevant actions (see Fig. 2), and had lower irrelevant imitation scores
(see Fig. 3). Though Ngandu children were intermediate between Aka children and Aka adults
in all three measures of irrelevancy, they were only statistically distinguishable from Aka
children, and only in irrelevant imitation scores rather than the number of irrelevant actions. This
may suggest that a proportional measure is more informative than the raw count data.
Although the irrelevancy quotient has the advantage of being comparable across studies with
differing methodologies and was seen to be a significant predictor of the time to reward retrieval,
the small sample sizes of this study restricted its statistical power. Considering all three
measures, Aka children were found to copy the demonstrated irrelevant actions at lower rates than
both Aka adults and Ngandu children.
In accordance with the results found for irrelevant actions performed, Aka children were
seen to have lower fidelity to the demonstrated sequence than Aka adults (see Fig. 3). Ngandu
children were again intermediate to both groups but only statistically distinguishable from Aka
adults. As overimitation involves a high degree of fidelity to the demonstrated sequence as well
as the production of irrelevant actions, these two sources of data together suggest that Aka
children were less likely to engage in overimitation than both Aka adults and Ngandu children.
When examined strictly on the basis of the social learning mechanism used (see Fig. 4), this
trend becomes much more pronounced: 60% of Aka children that succeeded in the task did so
by using emulation rather than imitation, compared to 15% of Ngandu children and 11% of
Aka adults. Therefore, not only were Aka children engaging in overimitation much less
frequently than the closely associated but culturally distinct Ngandu, they were also overimitating
less than the adults in their own community. Comparisons with toddlers and children from the
United Kingdom (see Fig. 4; Table 1) reveal that Aka children also displayed patterns of social
learning distinct from those being used by their developmental peers in a typical Western
culture. In previous Western studies focusing on early to middle childhood, the frequency of the
use of overimitation has been seen to increase with age while the use of emulation decreases
[20, 21, 2527], such that Western children of comparable age to the Aka tested here
overimitate almost exclusively, thereby utilizing a non-selective copying strategy which has been
shown to be less efficient at accomplishing complex tasks in the short term ([22, 26] and
present study). Aka children, by contrast, seem to readily filter out this costly irrelevant
information, much as Western infants and toddlers do [22, 23].
A number of explanations could ultimately be responsible for the observed cross-cultural
differences in childhood overimitation. Firstly, there are fundamental differences between
hunter-gatherer societies such as the Aka and more Westernized populations. These
differences are exemplified by three foundational schemas among the Aka, which are also common
among other hunter-gatherer groups: egalitarianism, or equal respect for all individuals;
autonomy, or the ability of each individual to do as he or she wishes; and sharing, that resources and
responsibilities are spread between members of the group (see S2 Appendix and  for more
information and for foundational schemas among the Ngandu). The cultural beliefs
represented by foundational schemas shape peoples perspectives on their social relationships,
interactions, and values, as well as their perception of and interactions with the world around them.
Culture can thus dictate the ways in which people learn by influencing whom they pay
attention to, which aspects of a models behavior are most salient to them, and how they perceive
and approach a novel problem. These features of individual and social learning are commonly
investigated through the action of cognitive biases, including frequency-dependent biases such
as conformity and model-based biases of individual aspects like prestige [9, 34, 60].
In any experimental study, one must consider the importance of the testing environment
and its relevance to the participants culturally normative experience. In this experiment,
participants were tested in naturalistic settings and demonstrations were performed by an ingroup
demonstrator. Experiments conducted in an artificial laboratory space, with video
demonstrations rather than live ones, or by demonstrators of a different gender, age, or ethnic or
socioeconomic group relative to the observer could find results that contrast with those of studies
that take these factors into account [60, 61]. These effects could be even more pronounced in
small-scale cultures where these kinds of experiences and interactions would be less familiar.
Nielsen and Tomaselli  did use ingroup models for some tests among the !Xun and Khwe
San and failed to find a significant difference between Western and local models, though this
shared community has a complex intercultural history with significant Western influences (see
S1 Appendix). An important observation is that rates of overimitation are lower in Western
studies in which the demonstrator is the same age or younger than the observer [28, 6264].
This is likely due to the perception of an older model as an individual of greater status and
experience that expects deference and respect, a cultural value common in Western groups but
rare in small-scale hunter-gatherers. The egalitarian nature of Aka society leads to the
perception of other Aka as equal regardless of age or status but does call for respect for members of
outgroups with greater influence such as the Ngandu. For this study, initially an adult Ngandu
man was trained to model for the Aka participants, but many Aka childrenparticularly
girlsdeclined to participate out of deference to him. Thus, the identity of the demonstrator
and the participants cultural views on equality and status can be very important components
of an experiment. The testing environment should also be carefully considered in research on
non-human primates and other animals, particularly when experiments are performed under
conditions that differ starkly from the natural ecology of the species or when actions are
demonstrated by humans rather than an intraspecific model.
Differences in levels of overimitation could also be driven by behavioral responses to
culturally constructed environmental niches . Aka children may typically be presented with
objects that are causally transparent (or at least translucent) rather than causally opaque. This
contrasts with the Western case in which so many of our modern technological artifacts,
including the toys and other objects that our children are exposed to from an early age, have
visible effects that are connected only in a very loose sense to the actions performed upon them.
This scenario could lead to blanket copying via overimitation being an advantageous strategy
for young children in a Western context but not in a small-scale non-Western one in which
emulation could be the more efficient approach. Overimitation could still be expected to
increase with age in both settings as the complexity of skills being acquired increases. In
smallscale cultures, the number of particular tools and social rituals with causally opaque functions
tends to increase with age, so the transmission and acquisition of these skills could be aided by
overimitation at older ages. However, this explanation is unlikely to apply uniformly to the
wide diversity of practices among non-Western cultures.
The finaland potentially most powerfulexplanation presented for the cultural
differences observed here is that Western attitudes on and approaches to education fundamentally
structure how children learn to learn from an early age. Even before children are enrolled in
formal schooling, parents begin the pedagogical process by applying the same cultural ideas
and practices regarding teaching that were used in their own education as, in a modern
Western context, the act of parenting is itself equated with teaching . Though so-called
Western schooling has a high degree of variation and has been introduced to the worlds
nonWestern societies at different times by different educational traditions , it is in all cases
markedly different from the environmental context of social learning in hunter-gatherers and
other small-scale groups. Aka children learn the majority of their skills through observation,
imitation, and participation. Some teachingdefined as the modification of ones behavior to
facilitate learning of information, knowledge, or skills in anotherdoes occur among the Aka,
but formal teaching as experienced in a classroom by Western children is absent [4, 68]. The
premise that learning is itself culturally learned is supported by a recent study by Mesoudi et al.
, which found cross-cultural variation in the use of social learning strategies between a
group of mainland Chinese students and other dispersed groups with varying levels of
In non-Western societies, the introduction of Western schooling has marked effects on the
value placed on traditional skills, knowledge, and attitudes and on the transmission of these
cultural variants, and often serves to integrate peripheral groups into dependent and
less-privileged roles in the dominant culture . Nielsen et al.  note that the groups of San and
Aboriginal Australians tested in the only non-Western studies of overimitation all had access
to formal schooling, albeit at different levels and with opportunities most limited among the
Khomani San. The Ngandu of Bagandu have had access to primary school for at least 40
years, meaning that most Ngandu adults have had some level of formal education and that
younger children have older siblings and peers that regularly attend school. The Aka in the
study area have had only limited access to local services and, consequently, none of the Aka
children or Aka adults that participated in this study have ever attended school (see S1
Appendix). The potential influence of these ontogenetic and cultural effects on the dynamic processes
of social learning and cognition should not be understated.
This study has demonstrated that cross-cultural variation exists in the degree to which
overimitation is expressed at particular ages in human children. Our data indicate that pronounced
overimitation does not universally emerge in early childhood; specifically, overimitation may
emerge later in life within small-scale, non-Western cultures. In contrast to Aka children, who
were found to be distinctly emulative, Aka adults displayed a high degree of overimitation
comparable to levels seen in Western populations. Ngandu children, with greater access to Western
influences and formal education, as well as cultural views and values that contrast sharply with
the Aka, also more closely resembled Western groups in their tendency to overimitate. As a
feature of social learning with possible implications for biological and cultural evolution,
overimitation merits further investigation across cultural, socioeconomic, and non-human groups to
gain a better understanding of its cognitive underpinnings and influences from cultural,
ontogenetic, and ecological factors.
There are notable limitations to the present study that point toward future directions in
research. The ongoing political crisis in the Central African Republic prevented the enlargement
of our sample size and the inclusion of Ngandu adults, which would allow for improved
statistical power and a more balanced assessment of overimitation across age groups in both
cultures. A shortfall of this study and others is the lack of attention on middle to late childhood
and adolescence which could shed light on changes in social learning strategies between
developmental periods across the lifespan . Future study on infants and toddlers in small-scale
groups could also track the development of overimitation at earlier ages than those examined
here. This study did compare overimitation in both children and adults among the Aka, which
has not been done for any other non-Western population and could provide insight into the
developmental components of overimitation.
The primary intended significance of this work is to echo recent introspection in the
behavioral, cognitive, social, and cultural sciences by Arnett , Henrich et al. , and others urging
the importance of avoiding sampling biases, even when cross-cultural sampling efforts can be
difficult or expensive. Cultures effects on human behavior are varied and profound and not
easily discerned from a subsample of ones own cultural group. This paper describes one of the
few contributions toward the cross-cultural study of childhood social learning and
overimitation. Though we believe the Aka at this field site are more representative of small-scale,
nonWestern societies than those previously studied due to their limited Western influences and
continued dependence upon hunting and gathering for their daily subsistence, it is also clear
that no two cultures are identical, whether they be Western or non-Western in nature. For
perspective, one must consider the range of cultural differences among the many groups of Aka
and other forest foragers of the Congo Basin, which number over 350,000 people and live in a
variety of conditions [53, 75]. The Aka studied here are but one example of the diversity of
ancient and modern small-scale cultures that have inhabited our planet.
S1 Appendix. Contrasting Western influences on the San, Aka, and Ngandu. Additional
comments on the Western cultural influences on the San populations studied by Nielsen and
Tomaselli  and Nielsen et al.  and on the Aka and Ngandu of the present study.
S2 Appendix. Foundational schemas of the Aka and Ngandu. Detailed descriptions of the
culturally distinct ways of thinking that structure many aspects of life for the Aka and Ngandu.
S1 Table. Type II ANOVA tables from Control analyses. Includes ANOVA tables, partial eta
squared values, confidence intervals, and results of Tukeys HSD post-hoc tests.
S2 Table. Type II ANOVA tables from Reproduction of Irrelevant Actions analyses.
Includes ANOVA tables, ANCOVA test result for age effects, partial eta squared values,
confidence intervals, and results of Tukeys HSD post-hoc tests.
S1 Dataset. Overimitation in the Aka and Ngandu. Full dataset used in analyses and the
generation of figures, in CSV format. Contains information on participant group, age, sex,
experimental setting, demonstrations, success, tool and door use, irrelevant actions, calculated
measures of irrelevancy and fidelity, test duration, and social learning mechanism used. Age is
in years and test duration is in seconds. See text for description of variables.
We would like to express our deep gratitude to the Aka and Ngandu families that agreed to
participate in this research. We also wish to extend our thanks to Andrew Whiten for his
assistance and the use of the experimental apparatus and to Kenny Smith, Mark Nielsen, and two
anonymous reviewers for their valued feedback on this manuscript.
Conceived and designed the experiments: BSH. Performed the experiments: BSH. Analyzed
the data: REWB. Contributed reagents/materials/analysis tools: REWB BSH. Wrote the paper:
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