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Cite as: P. Brakes et al., Science
10.1126/science.aaw3557 (2019).
POLICY FORUM
First release: 26 February 2019 www.sciencemag.org (Page numbers not final at time of first release) 1
Animal culture, defined as “information or behavior—shared
within a community—which is acquired from conspecifics
through some form of social learning” (1), can have important
consequences for the survival and reproduction of individu-
als, social groups, and potentially, entire populations (1, 2).
Yet, until recently, conservation strategies and policies have
focused primarily on broad demographic responses and the
preservation of genetically defined, evolutionarily significant
units. A burgeoning body of evidence on cultural transmis-
sion and other aspects of sociality (3) is now affording critical
insights into what should be conserved (going beyond the
protection of genetic diversity, to consider adaptive aspects
of phenotypic variation), and why specific conservation pro-
grams succeed (e.g., through facilitating the resilience of cul-
tural diversity) while others fail (e.g., by neglecting key
repositories of socially transmitted knowledge). Here, we
highlight how international legal instruments, such as the
Convention on the Conservation of Migratory Species of Wild
Animals (CMS), can facilitate smart, targeted conservation of
a wide range of taxa, by explicitly considering aspects of their
sociality and cultures.
CONSEQUENCES OF SOCIAL KNOWLEDGE
An important aspect of social learning is the speed with
which new behaviors can potentially spread through popula-
tions, with effects that may be positive (e.g., adaptive exploi-
tation of a new food source) or negative (e.g., increasing
conflict with humans, such as when sperm whales learn to
remove fish from longlines) (2). Transmission can be medi-
ated by an inherent propensity to adopt innovations [e.g.,
“lobtail” feeding in humpback whales (1)], or curbed by cul-
tural conservatism [e.g., southern resident killer whales’ per-
sistent foraging specialization on Chinook salmon (2)].
Social learning can result in the emergence of subpopula-
tions with distinctive behavioral profiles, erecting social bar-
riers, as observed in distinct vocal clans of sperm whales (see
the figure). Culturally mediated population structure has im-
portant implications for conservation efforts (4), as it can
influence species-wide phenotypic diversity and adaptability
to changing conditions (5). In some cases, such as humpback
or blue whale song, cultural variation can reflect demography
and facilitate more efficient, or less invasive, assays of con-
temporary genetic population structure (1, 4). Most pro-
foundly, culture can play a causal role in establishing and
maintaining distinct evolutionary trajectories (6).
Another consequence of social learning can be the in-
creased importance of key individuals as repositories of accu-
mulated knowledge, making their targeted protection
particularly important for the persistence of social units. For
example, the experience of African elephant matriarchs (see
the photo) has been shown to positively influence the fertility
rates of younger females in their social group, through the
transmission of information about the social and ecological
landscape (7). Yet, traditional approaches to species conser-
vation often prioritize younger individuals for their direct re-
productive potential.
Positive conservation outcomes can depend on the resto-
ration of cultural knowledge. For example, because whooping
cranes learn migratory routes socially, human surrogates in
ultralight aircraft can guide naïve, captive-bred birds along
their first migration, potentially boosting the effectiveness of
reintroduction programs (8, 9). Similarly, without the benefit
of socially inherited knowledge, bighorn sheep and moose
translocated to unfamiliar habitats can take generations to
master the skill of tracking the seasonal distribution of high-
quality forage (10). Social learning can also be exploited to
ameliorate human-wildlife conflict, for example, by artifi-
cially “seeding” desirable behavior, such as avoidance of par-
ticular foods or sites (3, 11).
To improve the efficacy of conservation efforts, we there-
fore argue that it is critical to consider the interplay between
social structure and the transmission of social information.
This may be particularly important if different categories of
individuals vary in their propensity to innovate, or are more
likely to be copied by naïve group members (11). In some
cases, populations may be structured into distinct cultural
Animal cultures matter for conservation
Philippa Brakes, Sasha R. X. Dall, Lucy M. Aplin, Stuart Bearhop, Emma L. Carroll, Paolo Ciucci, Vicki
Fishlock, John K. B. Ford, Ellen C. Garland, Sally A. Keith, Peter K. McGregor, Sarah L. Mesnick, Michael J.
Noad, Giuseppe Notarbartolo di Sciara, Martha M. Robbins, Mark P. Simmonds, Fernando Spina, Alex
Thornton, Paul R. Wade, Martin J. Whiting, James Williams, Luke Rendell, Hal Whitehead, Andrew Whiten,
Christian Rutz
Author affiliations are listed in the supplementary materials.
*Corresponding author. E-mail: p.brakes@exeter.ac.uk; christian.rutz@st-andrews.ac.uk
Understanding the rich social lives of animals benefits international conservation efforts
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units with differing resource requirements. For instance, cul-
tural transmission of vocal patterns among sperm whales in
the Eastern tropical Pacific results in distinct vocal clans (1)
(see the figure). Clans vary in their feeding success during El
Niño and La Niña oceanographic cycles (1), meaning that if
these cycles increase, as predicted under climate change, pop-
ulation-level impacts may not be uniform.
CETACEANS, AND BEYOND
Despite mounting evidence that aspects of sociality can have
far-reaching implications for wildlife conservation, interna-
tional policy forums—where most large-scale conservation
strategies are conceived—have so far not engaged substan-
tially with the challenges and opportunities presented by this
new scientific perspective. A notable exception is the CMS
Scientific Council, which has conducted work at the interface
of cutting-edge science and international policy-making.
The CMS signatories work to develop collaboration be-
tween range states for the conservation of species that move
across jurisdictional boundaries. They agree to support re-
search, to endeavor to provide immediate protection for mi-
gratory species listed in CMS Appendix I, and to work toward
developing agreements for the conservation and manage-
ment of migratory species in CMS Appendix II. Although, like
many other international agreements, CMS does not have a
compliance mechanism, its standing committee is in the pro-
cess of critically reviewing the impacts of its decisions, to im-
prove effectiveness (background CMS documents are
available in the supplementary materials).
Impetus for an animal culture initiative was provided by
a growing body of evidence for social learning and culture in
cetaceans that raised important questions about how best to
conserve these animals (12). The CMS Scientific Council’s
preexisting expertise in evaluating threats to aquatic mam-
mals made CMS a natural avenue for examining these issues
further. In 2014, a formal consultation revealed an extensive
range of circumstances in which social structure, social learn-
ing, and cultural variation in whales and dolphins can affect
the planning or outcomes of conservation efforts. This culmi-
nated in the adoption of a ground-breaking resolution,
through which the CMS signatories formally acknowledged
the importance of social learning and culture for the conser-
vation of some highly social species.
Following the 2014 resolution, the CMS Scientific Council
established an expert group to broaden the scope of this ini-
tiative beyond cetaceans. The group determined that social
learning has conservation relevance across a wide range of
vertebrate taxa, including birds, fishes, and many marine and
terrestrial mammals (1, 3). At a 2018 cross-taxa CMS culture
workshop in Parma, Italy, the authors of the present article
reviewed relevant evidence, with a particular focus on species
in which social learning has the potential to strongly
influence migratory behavior, habitat use, foraging, or inter-
action with human activities. On the basis of this work, we
recommend, among other things (for details, see table S1):
augmenting the designation of evolutionarily significant
units; conserving individuals that are critical repositories of
social knowledge; refining the criteria used for identifying
and prioritizing species and populations for assessment; im-
proving reintroduction schemes through strategic manage-
ment of social knowledge; planning effective mitigation
strategies for anthropogenic impacts using aspects of social-
ity; systematically cataloging the dimensions of cultural di-
versity; and raising awareness about the value of conserving
animal cultures.
The overall
aim of this initia-
tive is to maximize
the efficacy of con-
servation efforts
through enhanced
consideration of
sociality in gen-
eral, and social
learning and (both
adaptive, and
seemingly arbi-
trary) cultural pro-
cesses in
particular. Under-
standing the im-
portance of
behavioral diver-
sity will benefit
conservation poli-
cies both when as-
sessing the status
of potentially vul-
nerable popula-
tions (e.g., when
delineating units
to conserve, by ac-
counting for cul-
tural segregation) and when devising effective conservation
strategies (e.g., by identifying key repositories of social
knowledge). Achieving these ambitious goals will require a con-
siderable amount of work. For example, although there is broad
agreement that successful reintroduction programs require in-
dividuals to be behaviorally competent (8), for many species it
will still be necessary to establish the degree to which key be-
haviors are socially learned [e.g., migratory routes in birds (9,
13)]. To facilitate progress, we highlight a few additional oppor-
tunities, both in terms of particular species that may merit fur-
ther consideration and promising research approaches.
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The CMS encourages its signatories to engage in collective
conservation measures through its “concerted action” mech-
anism. This process is particularly relevant when considering
collaboration between range states for gathering and sharing
data on cultural diversity for populations that move predict-
ably across national borders. In 2017, CMS adopted a con-
certed action for Eastern tropical Pacific sperm whales (see
the figure). A variety of species may benefit from similar con-
sideration, to evaluate the importance of aspects of their so-
ciality for their conservation. This includes species as diverse
as cod (not currently listed in the CMS Appendices), which
appear to socially learn migratory routes, and chimpanzees
(recently listed in the CMS Appendices), where a culture of
nut-cracking tool use thrives in a small area of Western Africa
(see fig. S1), yet spans multiple national jurisdictions and may
provide access to an important food source during the dry
season (3).
An important challenge is to identify those populations,
or social units, that would most benefit from our proposed
approach, and to predict how specific biological processes
may influence conservation outcomes (11). Recent studies il-
lustrate how innovative rapid-assessment techniques could
aid the identification of distinct cultural units, which may be
particularly vulnerable (e.g., as a result of socially learned for-
aging strategies). Where socially transmitted traits—such as
foraging tactics (and hence resource requirements) and vocal
behavior—covary (1), it may be possible to document cultural
variation with well-established, inexpensive survey protocols
(4). Another approach is to harness new survey technologies,
such as autonomous motion-triggered cameras, as exempli-
fied by a recent attempt to chart putative cultural variation
in wild chimpanzees (14) in the face of urgent threats from
habitat destruction and poaching (see fig. S1). In addition,
appropriately parameterized formal models may provide re-
liable predictions about the impact of specific conservation
interventions on sociocultural processes (5). The field of ani-
mal social learning is now sufficiently mature to provide key
parameters for robust modeling of some systems, where rele-
vant data are available from long-term field studies and con-
trolled experiments.
MOVING FORWARD
Our growing understanding of the conservation relevance of
cultural variation urges that scientists and policy-makers col-
laborate closely to ensure that policy is informed by the latest
scientific advances. Many cultural systems are highly com-
plex, and the conservation impact of cultural processes is con-
text dependent, necessitating careful case-by-case
consideration. Recommendations from the Parma workshop
will inform discussions at the November 2019 Meeting of the
Sessional Committee of the CMS Scientific Council and the
2020 CMS Conference of the Parties in India. A key challenge
will be to determine if evidence warrants explicitly recogniz-
ing some distinct cultural units listed in the CMS Appendices,
and how insights from this work can be used to inform con-
servation efforts across the entire CMS portfolio of agree-
ments.
Within the broader context of international wildlife law
(15), it is important to consider the relevance of animal cul-
ture for scientific assessments and policy decision-making
across a range of relevant multilateral environmental agree-
ments, such as the Convention on International Trade in En-
dangered Species of Wild Fauna and Flora (CITES) and the
Convention on Biological Diversity (CBD). We see opportuni-
ties to extend our approach beyond species and issues cur-
rently covered by CMS, for example, when assessing the
sustainability of exports and trade through CITES processes.
Such consideration is timely, because 2020 is the final year of
the United Nations Decade on Biodiversity, when govern-
ments will negotiate the Post-2020 Biodiversity Framework.
Given the prevalence of social learning and cultures across a
wide range of taxa, a comprehensive, integrated approach is
essential to maintaining the natural diversity and integrity of
Earth’s rich ecosystems.
REFERENCES AND NOTES
1. H. Whitehead, L. Rendell, The Cultural Lives of Whales and Dolphins (Univ. of Chicago
Press, 2015).
2. H. Whitehead, Learn. Behav. 38, 329 (2010).
3. A. Whiten, Interface Focu s 7, 20160142 (2017).
4. E. C. Garland et al., Conserv. Biol. 29, 1198 (2015).
5. S. A. Keith, J. W. Bull, Ecography 40, 296 (2017).
6. A. D. Foote et al., Nat. Commun. 7, 11693 (2016).
7. K. McComb, C. Moss, S. M. Durant, L. Baker, S. Sayialel, Science 292, 491 (2001).
8. T. Mueller, R. B. O’Hara, S. J. Converse, R. P. Urbanek, W. F. Fagan, Science 341,
999 (2013).
9. C. S. Teitelbaum, S. J. Converse, T. Mueller, Conserv. Lett. 12, e12599 (2019).
10. B. R. Jesm er et al., Science 361, 1023 (2018).
11. A. L. Greggor, A. Thornton, N. S. Clayton, Behav. Ecol. Sociobiol. 71, 16 (2017).
12. H. Whitehead, L. Rendell, R. W. Osborne, B. Würsig, Biol. Conserv. 120, 427 (2004).
13. B.-U. Meyburg et al., J. Exp. Biol. 220, 2765 (2017).
14. H. S. Kühl et al., Sci. Rep. 6, 22219 (2016).
15. A. Trouwborst et al., BioScience 67, 784 (2017).
ACKNOWLEDGMENTS
We are grateful to the CMS executive secretary and the CMS secretariat for
organizing the Parma workshop; the hosts and sponsors of the workshop (the
Appennino Tosco-Emiliano National Park, the Fondazione Monteparma, and the
Principality of Monaco); M. Prideaux for helpful policy discussions; S. Smart for
graphic design; and the experts that have supported this initiative. G.N.d.S. is the
CMS Conference of the Parties Appointed Councillor for Aquatic Mammals. F.S.
is the chair of the CMS Scientific Council and is the councillor for Italy. J.W. is the
UK scientific councillor for CMS and works for the Joint Nature Conservation
Committee, which is a statutory adviser to the UK Government.
SUPPLEMENTARY MATERIALS
www.sciencemag.org/cgi/content/full/science.aaw3557/DC1
Published online 26 February 2019
10.1126/science.aaw3557
on February 26, 2019 http://science.sciencemag.org/Downloaded from
Animal cultures matter for conservation
Hal Whitehead, Andrew Whiten and Christian Rutz
M. Robbins, Mark P. Simmonds, Fernando Spina, Alex Thornton, Paul R. Wade, Martin J. Whiting, James Williams, Luke Rendell,
Ellen C. Garland, Sally A. Keith, Peter K. McGregor, Sarah L. Mesnick, Michael J. Noad, Giuseppe Notarbartolo di Sciara, Martha
Philippa Brakes, Sasha R. X. Dall, Lucy M. Aplin, Stuart Bearhop, Emma L. Carroll, Paolo Ciucci, Vicki Fishlock, John K. B. Ford,
published online February 26, 2019
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