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The New of , Conservation and Welfare

Credit: Institute of Research, Kyoto THE NEW SCIENCE OF ANIMAL CULTURE, CONSERVATION AND WELFARE

An abundance of evidence converges to suggest that humans are not the only species to create culture. Field and laboratory studies in have demonstrated that social and the transmission of the traditions that make up a culture can be found across many different species including , , fish and even insects, and in many different contexts.Professor Andrew Whiten at the and his extensive network of collaborators are pioneering conceptual and methodological frameworks to promote the conservation and welfare of animals from a scientifically rigorous perspective.

Article written by Dr Catherine Deeprose.

Culture: A Uniquely Human the country. Not long after, Japanese Characteristic? monkeys on Koshima islet were observed slowly but surely Culture has such a profound impact taking up the washing of sand-covered on our lives that it even shapes our sweet potatoes in water before eating, understanding of what it means to adopting the behaviour first observed be human. From foraging techniques by researchers in a young female named and to science and religion, Imo. Finally, evidence in support of our complex reflect the social vocal cultures in animals arose through learning we garner from others and the identification of regional dialects the transmission of traditions through in male sparrows in the wild, acquired generations. But is culture really in around the first 100 days of life. a uniquely human characteristic? Laboratory experiments confirmed that Research over the past 70 years has many need to learn their traditions – such as tool use, grooming accumulated to suggest that, contrary songs from the previous generation. and foraging techniques – being found to long-standing belief, humans are to have distinct features transmitted to not the only animals to create cultures These early but compelling others over time. that spread across populations and observations were followed by further generations. evidence that culture achieved through Social Learning: From Mothers and social learning and the transmission Others As early as in the 1940s, UK blue tits of traditions can be found across and great tits were observed tearing many different species including In a review article published in 2018 by open aluminium seals on milk bottles primates, birds, fish and insects in Professor Andrew Whiten (University to syphon off the cream inside. This both naturalistic (wild) and laboratory of St Andrews) and Professor Erica van behaviour, first spotted in only a very settings. Research with de Waal (University of Lausanne), three small number of birds, eventually has been particularly influential, with main phases of social learning were spread to entire populations across demonstrations of multiple cultural identified in the lives of many primates.

WWW.SCIENTIA.GLOBAL At the simpler end of the spectrum, there is associative learning, in which a new response becomes associated with a particular stimulus. For example, bumblebees who observe either trained or artificial bees foraging from either orange or green flowers later demonstrate the same preference. This type of learning may be the most common across many animal species.

The copying of complex behavioural routines occurs at the opposite end of the spectrum. Termite fishing by chimpanzees is a relatively complicated behaviour, requiring a number of steps including the selection of a suitable tool (e.g., a stick or blade) to extract the insects from the termite mound or subterranean chambers. Although The of swamp sparrow song in a 30-year study. From Annual Review of , a behaviour found in several different Evolution, and Systematics, 2019, with permission. communities, researchers have identified many distinct local The first describes the learning acquired monkeys with the greatest expertise, actions used for termite fishing in by infant primates from their primary enhancing their social learning of this different communities, indicating caregiver, usually their mother. In all-important skill. . Similarly, recent the main, primate mothers carry and research has shown that groups of breastfeed their infants for the first Finally, the third stage, dispersal, occurs and have specific months (in , years) of life, and when a new group member engages and unique vocal repertoires, food these maternal interactions provide in social learning of the dynamics and preferences and other behaviours. many opportunities for social learning. knowledge of the group they have The researchers cite the experimental joined. Returning to the study of wild Implications for Conservation example of groups of wild vervet vervet monkeys and the dyed corn monkeys fed corn dyed either pink or described above, the same researchers The protection of biodiversity – the rich blue, with one colour having an additive observed that new males to the group biological variety and variability of life making it too bitter to eat. Although were quick to adopt the food preference on Earth – is a key aim of conservation. young suckling monkeys were too (i.e., the colour of corn) of the new As such, efforts are typically young to eat the corn at this time, when group, even when this was contrary to geared towards sustaining natural presented the dyed corn four months the preference in the previous group populations of wild species, taking later when they were weaning – with from which they came. into account the genetic, ecological neither colour now containing the bitter and demographic risk factors for their additive – all the young monkeys ate The Cognitive Underpinnings of demise. Until recently, this involved little only the colour chosen by their mother, Social Learning consideration of social learning and indicating social learning of food culture. preference. Social learning and cultural transmission require sufficient cognitive Supported by the wealth of scientific In the second stage, young primates capacity – that is, the capability to evidence illustrating social learning extend their repertoire of social undertake the mental actions and and culture across a wide and diverse learning beyond that provided by their processes that lead to knowledge range of species, leaders in the study mother to others within their widening and understanding. Social learning of animal behaviour are now calling social circle, further expanding their takes differing forms across different for this knowledge to be applied to knowledge and skills. In a decade-long species, and these vary significantly the development of conservation study of brown capuchin monkeys and in complexity and thus, cognitive priorities and practice. In a significant their use of stones to crack open hard- underpinnings. step forward led by Dr Philippa Brakes, shelled nuts, younger capuchins were the importance of animal culture was found to preferentially observe older recently brought to the fore by the

WWW.SCIENTIA.GLOBAL Credit: Kokolopori Research Project

Convention on the Conservation of Migratory Species of Wild Among many uses, social learning can be employed to Animals, an environmental treaty which operates under the enhance socialisation with others. One study in chimpanzees aegis of the United Nations Environment Programme. demonstrated that their tendency towards food sharing could be increased by partnering one chimpanzee with another Researchers point to the distinct role that social learning plays that shared food with them. Afterwards, the chimpanzee who in the transmission of behaviours – and also note the rapid had been the recipient of the shared food was more likely to speed at which this can be observed (intra-generationally) share their own food with others. More harmonious social in contrast to behaviours that are genetically inherited environments within groups of marmoset monkeys have (between generations). Furthermore, the breadth of behaviours also been created by allowing them to ‘overhear’ positive associated with social learning, including foraging, migration vocalisations made by individuals in neighbouring enclosures, routes and mate choice, can all be critical to conservation thus fostering positive cultural norms. efforts. Finally, when social learning gives rise to the transmission of information through groups, this can lead to In addition to socialisation, enrichment of the environments culturally specific behaviours that are sustained both over of captive animals is fundamental to promoting good welfare, time and through generations. These are also of conservation and here, social learning again provides a valuable tool. For , not least because the transmission of locally specific example, when part of a social group, many animals are more knowledge and behaviours increases the likelihood of survival likely to explore novel spaces, eat novel food and engage with for populations at risk. novel stimuli (e.g., unfamiliar veterinary equipment) meaning that presentation to social groups rather than individuals is We are still some way off from truly embedding our likely to be more successful in social species. Relatedly, social understanding of animal social learning and culture into learning in this form has the potential to maximise training conservation policy and practice. To achieve this, much efforts, allowing animals to participate in procedures such as closer collaborations are required between scientists those required for husbandry or research with minimal stress and conservation policy-makers and practitioners, driven and intervention. forward within the context of appropriate conceptual and methodological frameworks. Such applications are not limited to animals living in captive settings. Researchers have proposed ways in which Implications for studying what and how animals socially learn can strengthen our understanding of wild animals’ welfare, promote the Learning, including social learning, allows animals to maximise maintenance of key behaviours, and mitigate negative their health, longevity, and reproductive success. While the human-animal interactions. Like captive animals, many wild benefits of this are readily apparent for animals in the wild, animals are exposed to anthropogenic stressors. Seeking a what about those kept in captivity? Dr Lydia Hopper (Johns comprehensive picture of the various social influences that Hopkins University) takes a particular interest in utilising animals experience will be key to enhancing the ways we care recent advances in our understanding of social learning and for and conserve animals across settings. culture in animals to improve the care and welfare of those in captivity. For captive animals, opportunities for learning can be Building on Scientific Foundations cognitively enriching, conferring welfare benefits. Additionally, captive animals are typically exposed to a vast array of social Over the last 70 years, our knowledge and insight into the social information from different sources, including their cage mates, learning and cultures of animals have expanded exponentially. other animals housed nearby, and also humans, providing We can no longer consider culture to be a uniquely human multifaceted opportunities for social learning. characteristic. But equally importantly, these scientific advances can now be used to propel and expand conservation efforts of animals in the wild, and to improve the management and welfare of animals, both in captivity and in the wild.

WWW.SCIENTIA.GLOBAL Meet the researcher Professor Andrew Whiten School of & Neuroscience University of St Andrews St Andrews UK Andrew Whiten has played a leading role in developing the field of animal culture, particularly in . He collaborated with Phillipa Brakes and colleagues in advancing policies linking animal culture and conservation and Lydia Hopper completed her PhD under his supervision. Professor Whiten completed his PhD in animal behaviour at the in 1972 followed by a Research Fellowship at the Alex Mesoudi (University of Essex) University of Oxford. He was appointed Lecturer in Psychology Mark Nielsen (University of Queensland) at the University of St Andrews in 1975 and later a Reader Luke Rendell (University of St Andrews) and Professor. He was promoted to a distinguished Wardlaw Christian Rutz (University of St Andrews) Professorship in 2001 and is now Emeritus Wardlaw Professor. Erica van de Waal (University of Lausanne) Over the course of his career, Professor Whiten has been (University of Zurich) instrumental in establishing the Centre for Social Learning and Gillian Vale (University of Texas) Cognitive Evolution (Founding Director 2003–2016) and the Stuart Watson (University of Zurich) ‘Living Links to ’ Primate Research Centre Hal Whitehead (Dalhousie University) (Founding Director 2008–2015). An elected Fellow of the Royal Society of Edinburgh, British Academy and international FUNDING Cognitive Science Society, he has been the recipient of a multitude of awards, including the Delwart International Biotechnology and Biological Sciences Research Council Scientific Prize (2001), Rivers Memorial Medal (2007), Osman British Academy Hill Medal (2010) and Sir James Black Medal (2014)., Professor Economic and Social Research Council Whiten is the first scientist to be awarded both the Royal Swiss National Science Foundation Society of Edinburgh Senior Prize and Medal in Life Sciences John Templeton Foundation and Senior Prize and Medal for Public Engagement. Royal Society Royal Zoological Society of Scotland CONTACT Leverhulme Trust Wellcome Trust E: [email protected] W: https://risweb.st-andrews.ac.uk/portal/en/persons/andrew- FURTHER READING AND RESOURCES whiten(00983056-5f70-4a76-ad39-5820ce066bf3).html A Whiten, The burgeoning reach of animal culture, Science, KEY COLLABORATORS 2021, 372(6537), eabe6514. LM Hopper, Leveraging Social Learning to Enhance Captive Christophe Boesch (Max Planck Institute for Evolutionary Animal Care and Welfare, Journal of Zoological and Botanical ) Gardens, 2021, 2(1), 21–40. Philippa Brakes (Exeter University) Emily Burdett (University of Nottingham) P Brakes, et al., A deepening understanding of animal culture Christine Caldwell () suggests lessons for conservation, Proceedings, Biological Nicolas Claidiere (Centre National de la Recherche Scientifique) Sciences, 2020, 288(1949), 2718. (Emory University) P Brakes, SRX Dall, LM Aplin, et al., Animal cultures matter for Sarah Davis (University of St Andrews) conservation, Science, 2019, 363, 1032–1034. David Erdal (University of St Andrews) Emma Flynn (University of Belfast) H Whitehead, et al., The reach of gene-culture in (Jane Goodall Institute) animals, , 2019, 10, 2405. Rachel Harrison (University of Lausanne) A Whiten, Social learning and culture in child and chimpanzee, Lydia Hopper (Johns Hopkins University) Annual Review of Psychology, 2017, 68, 129–154. Kevin Laland (University of St Andrews) ‘Animal Cultures: Core findings and New Horizons’, an online Nicola McGuigan (University of West of Scotland) lecture series: https://learn. Bill McGrew (University of St Andrews) culturalevolutionsociety.org/ Sarah Marshall-Pescini (University of Vienna)

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