Animal Cultures at the Edge of Extinction

Abstract

Over the course of the latter part of the 20th century the notion that some animals might partake in a cultural form of life has gained growing support in the natural sciences. Iconic examples of tool using chimpanzees, sweet potato washing macaques, and milk bottle opening birds have captured scientific and popular interest alike. But at the same time that this effort to describe, define, and study animal cultures was developing, the global ecological crisis was deepening. This article explores this strange juxtaposition. It offers a critical overview of the concept of animal culture as it has taken shape in the natural sciences. Building on this foundation, the article explores how and why animal cultures matter at the edge of extinction, exploring possible roles for the environmental humanities, extinction studies, and philosophical ethology, in developing new approaches to the question of animal culture in a time of escalating biodiversity loss.

Introduction

Over the course of the latter half of the 20th century, the cultural lives of animals have slowly emerged as a legitimate topic of scientific inquiry. At first, they were carefully documented and analysed by only a small number of scientists, and with a focus on only a few nonhuman species, especially among the primates, cetaceans, and birds. These early researchers were captivated by a few animals up to very interesting things, particularly because the emergence of these behaviours was difficult to explain: blue tits who learned to pierce the foil lids and drink from the milk bottles left on the doorstep of UK residents; macaques of Koshima Island who found that sweet potatoes were better enjoyed when they went to the trouble of dipping them in salt water; chimpanzees of Gombe using wood and stone tools in ways that show social complexity, cultural variation, and individual personality; and song sparrows of North America who composed songs with regional variation or dialects that they were open to changing over time. From a handful of examples, interest in the question of animal culture has steadily grown to the present day.

Precisely what holds all of these examples together – in other words, what it is about each of them that constitutes what might be called ‘culture’ – is a topic of considerable contention. This makes sense, of course. Culture has proven to be notoriously difficult to define wherever it occurs, including amongst humans. The extensive study of human cultures, undertaken by a range of disciplines, has most often been founded upon a fundamental distinction of humanity from animality. Importantly, the connections between this vast body of literature and the study of animal cultures remain largely underexplored (Lestel, 2003). For the most part, however, scientists studying animal cultures have understood it as a specific domain in which certain animal behaviours are shared amongst a group via social interaction – that is, they are ‘socially learned’ – rather than being determined in some way by genetics or environmental factors.

There is something fascinating and promising about this recognition of culture in animals within the mainstream Western sciences of biology, ethology, psychology, and related fields. And yet, it comes with a range of complex compromises and challenges. Not least amongst them today is the fact that we – and the many animals and other species we share this planet with – now find ourselves living in the midst of what other scientists are referring to as the planet’s ‘sixth mass extinction event’ (Barnosky et al., 2011; Cowie et al., 2022). Over roughly the past 60 years, as efforts to describe, define, and study animal cultures multiplied, relatively little attention was paid to the fact that this research was taking place in the midst of an escalating ecological crisis. In recent years this has begun to change, with a small but growing group of biologists calling for greater attention to how and why ‘animal cultures matter for conservation’ (Brakes et al., 2019). In the humanities and social sciences, too – perhaps most notably in work in ‘extinction studies’ and the broader environmental humanities – some scholars have been exploring the transformation and loss of animal cultures in this time of extinctions and why and how this matters (Crist, 2013; De Vos and Chrulew, 2018; van Dooren, 2014).

This article explores this strange and confronting juxtaposition of a growing scientific and popular awareness of the complexity of animal life and of the incredible rates of species loss. The first part of the article lays the foundation, offering an overview of the concept of animal culture as it has taken shape in the sciences over the past several decades. Alongside highlighting some of the key contours of scientific thought in this area, this part also draws out some of the major limitations in how this domain is defined and suggests how more interdisciplinary approaches might help to overcome them, drawing in particular on scholarship in the environmental humanities, extinction studies, and philosophical ethology.

The second part of the article turns to how and why animal cultures matter at the edge of extinction. This discussion is aimed primarily at scholars working in the environmental humanities and the related fields of extinction studies and multispecies studies. It explores what taking the cultural lives of animals seriously might demand of those of us working in these areas, how it might expand and complicate our sense of what is lost in extinction and why this matters, what the implications might be for practical responses to endangerment, and what kinds of collaboration might be possible in this space.

I What Counts as (Animal) Culture?

The profoundly anthropocentric nature of Western thought about other species for at least the past several centuries has been well documented. For the most part the rich and complex cognitive, behavioural, social, and emotional lives of animals have been ignored or minimised in a way that has positioned humans – some more than others – as the sole locus of rationality, language, and much more, including, of course, culture (Crist, 1999; Lestel, 2003; Plumwood, 2002). Even within the post-Darwinian framing of evolutionary continuity between humans and animals, which overturned many aspects of this human exceptionalism, nonhuman species (and their cultural traits) remained positioned as mere precursors to properly human culture and were thus studied, when they were studied at all, as models for the insights they could provide into the origins of our exceptional cultural abilities.

This legacy matters deeply. As animal cultures began to be documented by a range of behavioural scientists in the 20th century, the ways in which these phenomena were framed and understood, as well as the methods by which they were studied, were significantly shaped by this history. Following the documentation of cultural innovations in some infamous cases, field and behavioural scientists working largely but not exclusively with birds, primates and cetaceans sought ways to identify and isolate culturally transmitted behaviours (Fisher, 1949; Laland and Galef, 2009; Nishida, 1987; Whitehead and Rendell, 2021; Whiten and Boesch, 2001). The resulting definitions of animal culture have focused on a range of criteria: culture should be socially learned or transmitted, be shared amongst a group, be persistent (but open to change) over time, be picked up through cognitively complex forms of learning (i.e. perhaps not ‘mere facilitation’) and be passed down through generations. Some definitions are far more restrictive, for example arguing that in order for a behaviour to be cultural the form of social learning must be that of imitation or instruction (Galef, 1992). ¹ Others have argued that to be properly cultural a behaviour must constantly be undergoing iterative improvement (often discussed in terms of ‘ratcheting’ and ‘cumulative culture’) (Tennie et al., 2009). While definitions of animal culture remain contentious around the edges, today many scientists seem to be satisfied working with a broad and open definition of a cultural behaviour as one that is ‘socially learnt and acquired, shared by members of a group or community and persistent over time’ (Aplin, 2019: 181; Laland and Janik, 2006).

While there is a great diversity within the scientific research on animal cultures – in terms of disciplinary affiliations, methods, and approaches – it is nonetheless possible to identify two broad empirical approaches. Importantly, as Dominique Lestel notes, these two approaches have maintained largely distinct trajectories and are, by and large, ‘not practiced by the same researchers’ (Lestel, 2014: 98).

The first of these approaches dominates amongst lab-based scientists and is more experimental (than observational) in nature. In the main experimental technique used within this paradigm, a novel behaviour is taught to a given individual so that its spread can then be studied and manipulated to better understand the social mechanisms of its transmission. The neurobiologist Robert M. Sapolsky helpfully sums up this approach, writing: ‘the gold standard for culture is one that can only occur in an experimental setting. This is when an individual(s) is removed from a group, is taught a new behavior that is genetically and ecologically arbitrary, is returned to the group, and where that novel behavior is then shown to spread’ (Sapolsky, 2006: 219). While such controlled conditions might rarely be achievable, a range of experimental interventions have enabled different forms of social learning to be taxonomised, and the patterns of transmission chains to be mapped, whether by using tape recordings to tutor songbirds, by introducing stimuli to modify bumble bee flower colour preferences, or by testing how well chimpanzees copy the behavioural sequences modelled to open an artificial fruit box.

It was the second major approach, more common amongst field biologists and primatologists, which provided the bulk of the early evidence for animal cultures (such as the potato washing and milk bottle opening mentioned above) through longitudinal studies of wild-living populations. Here, observational or ethnographic methods are used to study the presence and/or dissemination of unique, naturally-occurring behaviours amongst a wider group or groups. In most cases, candidate behaviours are identified by virtue of the fact that they are specific to a population, rather than being shared by all members of a species. As population specific behaviours can also arise as a result of unique ecological or genetic differences, scientists also generally try to rule these things out as causes for the candidate behaviour. This approach is referred to as the ‘method of exclusion’ (MoE). While it was originally ‘intended as a tool to detect the presence of culture’, it has today come to function as a central part of how animal culture is understood: ‘the criteria of the MoE are now so ingrained that this operationalization has, to all practical purposes, reached the status of the definition of animal culture’ (Schuppli and van Schaik, 2019: 3).

Although these scientific approaches differ significantly, it is important to note that they are both products of particular disciplinary contexts and histories, and they have both, in turn, come to shape in significant ways what has counted as culture. This point is made explicitly by Schuppli and van Schaik (2019) in relation to the way in which the exclusion of genetic and ecological factors has come to be incorporated into definitions of animal culture. We see this, for example, in Nishida’s classic definition: cultural behaviour is defined as ‘behaviour that is (a) transmitted socially rather than genetically, (b) shared by many members within a group, (c) persistent over generations, and (d) not simply the result of adaptation to different local conditions’ (Nishida, 1987). As is discussed further below, this focus on culture as distinct from genetics and environment, rather than on the ways they intertwine, has a range of conceptual and practical implications for how animal culture is researched.

But the impact of these research methods is also present in less visible ways. For the most part, scientists working on animal culture have understood it as an exceptional element of animal life, one limited to particular behaviours within the larger behavioural repertoire of certain species. As a result, in many of these discussions animal culture is still often framed as something of an ‘add on’ to the fundamentally biologically determined character of animal life. Humans remain the cultural creatures par excellence – beings who live out their lives fully immersed within a cultural domain – while other animals seem to be framed as at best occasional visitors, dipping into such a space from time to time. This understanding has no doubt influenced the kinds of animals amongst whom culture was initially sought, i.e. beginning with those ‘most like us’. At the same time, it is important to recognise that elements of this exceptionalism have been straining and perhaps even breaking down in recent years as examples of animal culture have multiplied. Discoveries of nonhuman cognitive ingenuity and social complexity have spread from the pages of scientific journals to popular science publications and policy documents, contributing to a widespread recognition of and wonder at the marvels of ‘how animals learn to be animals’ (Bonner, 1980; De Waal, 2008; Safina, 2020) and transforming the legal frameworks that regulate their treatment (Cavalieri and Singer, 1993; CMS, 2017). Even invertebrate animals are now commonly described by some scientists as potentially having ‘cultures’ – something that was unthinkable to most in these fields a few decades ago (Bridges et al., 2023; Mery et al., 2009). For example, one study showed that bumblebees can readily learn new behaviours (in this case, how to pull a string to access a food reward) by observing knowledgeable individuals, suggesting that these animals possess ‘the basic cognitive elements required for culture, i.e., group-specific behaviors that spread from “innovators” to others in the group via social learning’ (Alem et al., 2016: 1).

But while the taxa that might be included in this space have expanded, the focus on specific behaviours has remained largely unchallenged. The vast majority of the scientific research on animal cultures is actually research on discrete cultural behaviours that become separated out from the larger sphere of animal life: the opening of milk bottles, the dipping/ washing of sweet potatoes, etc. This focus on specific behaviours is itself a legacy of particular regimes of fact making that focus on the observable and repeatable, and as such tend to break complex, interconnected phenomena down into isolated, measurable units (Dewey, 1929). In the context of animal cultures, exceptional cultural behaviours have taken on that role as entities that are, in theory, more amenable to experimental testing, and thus more readily constitutable as empirical data. In many ways continuing earlier behaviourist legacies, this focus centres in on one element of the broad and interconnected array of attitudes, ideas, habits, practices, and more, that might be thought of as a ‘culture’.

Having said this, however, it is worth noting that even these singular behaviours have proven to be remarkably resistant to forms of scientific testing and validation, whether in terms of challenges in constructing the experimental conditions required to achieve the ‘gold standard’ described above, or in isolating the influence of social learning from other factors contributing to the emergence and dissemination of a given behaviour. Indeed, there is an important question regarding the appropriateness of such methods for phenomena deemed cultural. Any successful approach to animal culture will have to move beyond the inherited dualism and anthropocentrism that has divided the disciplines and made questions of anthropomorphism so vexed (Daston and Mittman, 2005) to contend with the interplay of epistemology and ontology in a domain where the objects of investigation are themselves active, changing, meaning-making subjects (Stengers, 2000). The exceptional, variable and metamorphic nature of the traits being investigated – precisely what makes them worthy candidates for research – tends also to make them difficult to pin down. Schuppli and van Schaik (2019), for example, have argued that there is a major flaw in the exclusion method in that there is no good reason to expect geographic variability in socially learned behaviours. As such, the widespread reliance on the existence of this variability to identify candidate behaviours has led to a variety of socially learned skills being ignored, effectively only allowing ‘the tip of the iceberg’ of animal culture to come into view (a history of underestimation which, as the field of animal studies has more widely noted, has significantly impacted the way animals are viewed and treated). Schuppli and van Schaik support instead a more anthropological (and epistemically more generous) approach of directly counting socially transmitted skills that potentially reveals a much wider distribution of animal culture than typically assumed.

This is an example of a growing effort in recent years to identify multiple cultural behaviours in a single species. Chimpanzees are perhaps the poster children for this effort, with each population that has been closely studied having a distinctive combination of approaches to grooming, tool use, foraging and more (Whiten et al., 1999). On this basis, some primatologists have argued that the term ‘culture’ should be reserved for populations that have a variety of distinctive, socially learned practices (rather than those that possess a single documented ‘cultural behaviour’). But there are other examples, too. While avian song cultures had long been intensely studied, songbirds were assumed by many to be ‘one-trick cultural ponies’ until subsequent research revealed evidence of foraging, nesting, and migratory cultures (Aplin, 2019). In one sense, this effort to identify a variety of cultural behaviours in a single species produces a more complex notion of culture, unsettling its reduction to individual behaviours. But from another perspective, it might be argued that, in remaining with a behavioural definition of culture, this kind of approach only sidesteps the core problem.

We want to suggest that there is an important sense in which the question of animal culture opens the door for, and indeed demands, a more wide-ranging and adventurous transdisciplinarity. The debates among ethologists, behavioural ecologists, ornithologists, primatologists and evolutionary anthropologists (among many others) over the best concepts and methods for animal culture research open up a range of epistemological, ontological, ethical and empirical questions to which fields like the environmental humanities are now poised to make a distinct contribution – and not only in the mode of ‘supplementation’ to which they have often been relegated. For philosopher Dominique Lestel, taking the question of animal culture seriously ‘disqualifies the university organization of knowledges that is explicitly founded on the dichotomy between human and animal’ (Lestel, 2014: 106). The recent upsurge of interdisciplinary cross-pollination in which the environmental humanities and extinction studies participate offers a range of conceptual innovations and experimental approaches less beholden to ‘intractable discipline-specific burdens of proof’ (Taylor, 2021: 1) and potentially more appropriate to the variability and complexity of animal culture (Parathian et al., 2018).

This work of ‘disciplinary rapprochement and boundary crossing’ (Nimmo, 2012: 189) draws fruitfully on earlier science and technology studies scholarship on the way in which animal culture emerged as a sphere of concern, and how the particular histories and practices of different scientific communities (the ‘culture’ of the scientists) influenced the ways in which animal ‘cultures’ came to be revealed and disregarded, defined and dissected, problematised and articulated (the encounter between science studies and primatology being the most well-known example; see, to begin with, Haraway, 1989; Strum and Fedigan, 2000; Strum and Latour, 1987). While the shape of the transdisciplinary scholarship needed to respond to contemporary environmental problems is still being contested, some of its distinctive trajectories can already be discerned, such as a renewed exploration of collaborative teamwork, and a move beyond familiar humanistic gestures of critique, contextualisation and reflexivity towards not only forging better connections (with both scientists and their objects of investigation) but also assuming a more active role in the contestation over legitimate knowledge and practice regarding animal culture itself.

Importantly, a fuller understanding of animal culture must recognise how processes of social learning and interaction shape and pervade the way of being of a group and its members. Culture cannot be cordoned off into one or several behaviours – at least not without significant consequences. And while social interaction and learning are integral to any meaningfully cultural form of life, this thing called culture is not merely a discrete element added onto a biological substrate and cannot be neatly separated from the various ontogenetic and environmental factors that also (and together) shape the specific ways in which animals go about their lives. As such, we are interested less in ‘cultural behaviours’ than in broader ‘ways of being’ or ‘forms of life’ – which always emerge out of and take new shape within complex developmental and social processes. These ways of life are not fixed or static but involve active and participatory intra-actions and transactions with a diversely populated world (van Dooren and Rose, 2016).

The work of Eytan Avital and Eva Jablonka is instructive here:

Think about foxes that move into towns: they acquire a new way of life that is socially transmitted to their offspring. They learn how, when and where to forage in the town’s rubbish bins and gardens, how to avoid humans, where to dig a den and so on. . . . Existing patterns of behaviour become modified or are abandoned, and gradually a new life style develops. . . . It is the total life style that constitutes the culture, and this includes those socially transmitted elements that are partially dependent for their expression upon the contingent conditions. The same kind of argument can be used to justify calling the socially learnt and transmissible life style of red foxes that have colonised urban areas ‘urban-fox culture’. (Avital and Jablonka, 2000: 22–3)

This approach refuses a reduction of culture to cultural behaviours, whether one, two or thirty. It insists instead on a thicker, more multifaceted and integrated, notion of cultural life – what the authors refer to as a life style, and which might also be indicated by terms such as form or way of life. Importantly, for them too, ‘culture’ is not an element of an animal life that can be sensibly isolated from the biology of the species, or from the environment in which that life is lived; moreover, the fact that it is subject to change, including in response to anthropogenic conditions, does not disqualify it from being meaningfully considered as cultural. Furthermore, this framing diverges from the orthodox search for causal explanations in which a behaviour is either genetically, environmentally or culturally determined, and instead recognises not only that these factors are always intertwined and interdependent, but that one of the key elements (and values) of culture lies precisely in the capacity for social learning to enable a variety of inventive and strategic adaptations to local environmental conditions.

From such a perspective, ‘culture’ is not an entity that can be pinned down and exhaustively described. It is not some-thing that an animal, or a population, or a species, has. Rather, it is a part of the doing of a life. To be a cultural being is to be a creature whose way of being is inherited in some part through processes of social learning. But it is also to participate in this form of being in a creative, meaningful, embodied manner – to actively inherit, enact, interpret, transform, and pass on.

Paying attention to the diverse ways in which living beings embody and enact their specific modes of life requires the realisation that there is a multiplicity of ways of being cultural. Animal cultures are not the same as human cultures, nor merely diminished or more primitive forms that teach us about human evolution. Yet they do nonetheless intertwine with various phylogenetic, cognitive and behavioural differences – a ‘heterogeneous multiplicity’ (Derrida, 2008) of embodied and enacted traits that not only challenge humanistic dualisms in their diversity, but that also invite and demand more curious attention in their specificity (Chrulew and Lestel, 2014). Indeed, one of the more profound implications of the cultural revolution in ethology is that of ‘the plurality of cultures’ (Lestel, 2002), which are nonetheless at least partially knowable through well-designed etho-ethnological procedures (Despret, 2016; Lestel et al., 2006). As a result, the exploration of culture will be undertaken most productively when we combine concepts and methods from the biological and social sciences and the humanities and when we work back and forth between multiple cases and definitions that allow us to see a broad range of both similarities and differences between the ways living beings have developed to explore forms of life that are at least partly socially shared.

Importantly, this approach also demands attention to the always multispecies nature of all forms of cultural life. There is a tendency in much of the scientific work on animal cultures to view them through an exclusively single species lens, focused tightly on the transmission of behaviours through social learning amongst conspecifics. In fact, the assumed single species nature of culture is frequently built into definitions. In contrast, the approach that we are suggesting is one that pays attention to the ecological and intercultural agency and impact of the many other species interacted with (human and not), as well as the broader landscape, as active parts of the milieu in which all developmental and reproductive processes – including social learning – take place. Indeed, when thought in Batesonian terms (Bateson, 1972), it is clear that the very form and content of animal cultures will necessarily be shaped precisely by this multispecies ecology composed of various relationships – procreative, predatory, mutualistic, or otherwise – that are at once biologically functional and biosemiotically meaningful.

What are we to take from these critical challenges to dominant modes of understanding animal cultures in the scientific literatures? Is the concept unworkable? We are convinced that something important is being described here, or at least gestured towards. At its core, this scientific attention to animal cultures represents a long overdue recognition that other animals are not mindless automata playing out a genetic or evolutionary script but intelligent and communicative subjects interpreting and acting in their worlds. Even more than this, the recognition of something like ‘culture’ goes a step beyond the appreciation that individual animals might think, learn and invent; that their ways of life might be shaped by personal experiences and life histories. In addition, it asks us to see how those individuals have and might come together to collectively craft a shared way of being that is at once relatively integral and open – and indeed one that can be shared across multiple species. Or, at least, this is the promise of what animal culture research might offer if it can be reimagined in more interdisciplinary and multispecies terms.

II Animal Cultures at the Edge of Extinction

The final half of this article takes up this complex terrain of ideas and research in relation to animal cultures to explore what it might mean for ongoing efforts within the environmental humanities to engage thoughtfully and productively with our present time of extinctions. What questions might animal cultures help to open up? What new alliances and practices (Stengers, 2017) might the growing scientific acceptance of something called ‘animal culture’ enable – even if those alliances and practices require quite a bit of work, at a slower pace more congenial to exchange, not least in navigating what will count as culture? Scholars working in the environmental humanities, especially the field of extinction studies, ought to see this terrain as an opportunity to extend and test their transdisciplinary labours, and set themselves the challenge of contributing meaningfully to the very form and structure of the questions posed and interventions staged in response to the endangerment of animal species and cultures.

Storying the Texture of the Emergence and Decline of Animal Life

One of the core commitments of environmental humanities research on extinction is an effort to move away from a consideration of the phenomenon of extinction in abstract terms and to instead focus on particular extinction events in all their specificity. Scholars in this field have often sought to give an account of disappearing species as distinctive ‘ways of life’, not reducible to biogenetic diversity that can be readily banked as a captive population, a DNA sample, or in some other form (see the essays by Rose, van Dooren, Chrulew, and Kirksey, all in Radin and Kowal, 2017). Rather, species are seen as embodied, intergenerational, relational processes that are sustained in the world through the work of living and reproducing, as well as, increasingly, through the care work of others (van Dooren, 2014). As such, the way of life of a species is not a static and unchanging one. It is a collective, evolving, creative achievement. It has a distinctive texture in and through which its inherently cultural animality weaves its world, and weaves its inter-textural convergences with other species, including humans (Lestel, 2015: 62). And just as species evolve and are sustained in relationship with others, so too is their decline and disappearance a relational process (Reinert, 2013). In this context, work in this field has often understood extinction as a process of unravelling in which diverse, multispecies, biocultural relationships are transformed or undone (Chrulew and De Vos, 2018; De Vos and Chrulew, 2018; van Dooren, 2014).

Through this process, species emerge as something much more than another Latin name on a long list of threatened creatures. In seeking to give an account of disappearing species in this way, work in extinction studies has aimed to more fully articulate what a potential extinction means, why it matters, and to whom – including both the animals themselves and the wider ecologies and communities with which they are intertwined. Doing so is a work of ‘storying extinction’ that aims to summon up the dead and dying, to give them presence on the page, in a way that draws others into a fuller sense of both appreciation and responsibility for a pending loss (Rose et al., 2017; van Dooren and Rose, 2016).

In this context, attending to animal cultures can be an important part of understanding the way of life of a given species, how it holds itself in the world, and how its decline ripples out to impact on a diversity of others. Consider, for example, the social lives of African elephants. Scientists now believe that elephant matriarchs carry the memory of their herds in important ways. As they guide the herd, they know about dangers that might be encountered at particular times and places, as well as where reliable sources of water, food and other important resources might be located. Of course, the matriarchs’ knowledge comes from their own experiences in a particular environment, but it is also something that is socially learned ‘in community’, drawing on their experiences with older elephants earlier in their life, elephants who first benefited from the accumulated knowledge of the generations before them (McComb et al., 2001). This knowledge has important conservation implications: its maintenance and continuity is thought to contribute to higher group survival rates and reproductive success (for younger elephants in the herd), while its disruption has been claimed to contribute to social unravelling and ‘problematic’ behaviour. As a result, some conservationists are calling for management plans that recognise the importance of these individuals, inheritances, and social dynamics (Brakes et al., 2021). But at a more fundamental level, a detailed knowledge of these social processes also transforms our sense of who these creatures are, of their subjectivity and agency, their ethos and character, of what their way of life consists in, of the incredible, intergenerational gift of elephant life, carried along on ‘waves of ancestral power’ (Rose, 2022: 128) by the endeavour, knowledge and skill of countless generations.

In this example we also see how cultural complexity can enable living beings to adapt to a changing world, but at the same time how it might produce critical new forms of fragility. Matriarchal elephant knowledge ‘works’ in the way that it does precisely because it is flexible and adaptable, learning and shifting with the seasons and landscape transformations. But there are limits. The speed and scale of change matters and at some point, for all a herd’s knowledge, they will struggle to adjust to habitat and climate transformations. This malleability brings with it a susceptibility to excessive interruptions and challenges. As with other modes of inheritance, the processes of social reproduction that animate this cultural practice have their own particular forms of fragility: they can be disrupted, wounded, transformed, and ultimately lost through a variety of forms of social disruption. What this case suggests, among other examples of animal cultural knowledge, ² is how attending to animal cultures might significantly enhance the capacity of work in extinction studies and the broader environmental humanities to story the texture of the emergence and decline of animal life.

Operationalising Culture

While scientific research on animal cultures slowly gained pace over the latter half of the 20th century, it is fair to say that it was largely ignored by conservation biologists. Indeed, until quite recently, the entire field of animal behaviour had been largely set aside by conservationists, as scholars in both the environmental humanities and the emerging field of conservation behaviour have argued (Berger-Tal et al., 2011; Greggor et al., 2016; van Dooren, 2023; van Dooren et al., 2023). In just the past few years this has begun to change, with a small group of cognitive and behavioural scientists working to articulate how research on animal cultures might be of relevance to conservation efforts (Brakes et al., 2019). At the core of these arguments is a sense that animal cultures might either pose a risk or offer an opportunity for the ongoing survival of a species; in short, that animal cultures might be operationalised, put to work, for the sake of conservation.

Perhaps unsurprisingly, given what has already been said, these efforts to engage with animal cultures for conservation ends have tended to do so in a way that is focused on individual cultural behaviours, rather than more fully textured ways of life. We might think about some of the many efforts to conserve or reintroduce behaviours lost in captive breeding programs, from foraging knowledge to predator avoidance and migration. Instructive examples include: endeavours to catalogue the knowledge that rehabilitant orangutans need to survive in the forest – and teach it to them (Grundmann et al., 2001); debates over the need for, and the welfare implications of, using various forms of training and simulation to reintroduce fear of predators into captive enclosures, in order to improve the chances of successful reintroduction of captive animals (such as black-footed ferrets) to the wild (Miller et al., 1993); and efforts to teach a few generations of captive-bred whooping cranes when and where to migrate with the use of an ultralight aircraft so that they might pass the behaviour on to others after release to establish a new, safer migratory route (van Dooren, 2014).

In these cases it should be noted that the loss of animal culture being recognised and remediated often occurred, at least in part, as an unforeseen result of conservation interventions themselves. This fact ought to pose a number of difficult questions as to the state of the knowledge and practices involved. Moreover, the solutions proposed are often still largely in keeping with the reductive approaches that have contributed to, if not produced, these disconnections. The rush to intervene within a genuine ‘state of emergency’ that has characterised conservation biology as a ‘crisis discipline’ (Soulé, 1985), as well as the disciplinary norms and boundaries that obstruct meaningful collaboration, have impeded proper attention to the history of these ‘ethopolitical’ experiments in existence (Chrulew, 2021). This is a history that holds important yet all too often unheeded lessons – and it is a history that can only be told by finding ways to combine both scientific assessment of experimental successes and failures, and humanistic critique of the concepts and practices in ethical and epistemological terms (forms of reflection which, for all their disciplinary differences, often converge meaningfully in their aims and methods in unexpected ways; Stengers, 2017).

Importantly, these efforts to operationalise cultural behaviours are not always aimed at protecting or reinstating a behaviour. Sometimes, intimate knowledge of social learning is put to work in conservation contexts in order to ‘stamp out’ behaviours deemed problematic. For example, recent research has identified the emergence of a behaviour among a group of California sea lions whereby they travel upriver to eat large numbers of endangered salmon. This behaviour appears to be spreading, as new individuals observe and learn from others. As a result, some conservationists have proposed a targeted ‘cull’ of the offending sea lions to prevent any further social uptake of the problematic behaviour (Schakner et al., 2016). While arguments for animal perception, behaviour, cognition, subjectivity, personality and culture are often deployed to challenge anthropocentric ideas and practices, this example provides an important reminder that such recognition can be a poisoned chalice: more precise knowledge has regularly been wielded to refine and intensify forms of management and control in the interests of power and capital (Chrulew, 2016). In short, in contemporary conservation worlds, cultural behaviours are by and large either ignored or assessed on the basis of their impacts on relevant threatened species. In this way they become ‘resources for survival’ that can be slotted in or removed to achieve the goals of conservation.

While these kinds of connections between animal cultures and species survival matter profoundly, it is far from clear that this is the best way to understand or intervene in these contexts. The danger to heed is a situation where advances in knowledge of animal cultures only carve out and justify a further domain of intervention in animal lives, an expansion and intensification of biopower (and its psycho-, etho-, enviro- and onto-political extensions) in ever more programmable ecologies. There is an important role for scholars in extinction studies and the environmental humanities in interrogating these life and death making regimes of knowledge and power to ask how and with what consequences particular understandings of animal cultural life emerge and are put to work. In addition, there are also important and productive opportunities here for both collaboration and provocation, working with relevant scientists to utilise and transform these knowledges and practices and challenging them to critically explore the possibilities in this space, including systematically engaging with the histories of their own disciplines and past efforts to grapple with related questions (as well as heeding the lessons of earlier border skirmishes between disciplines, as in the science wars; see e.g. Langlitz and Strum, 2017). There are some promising early signs that this kind of cross-disciplinary conversation and collaboration might be productive (van Dooren et al., 2023).

Theorising Processes of Cultural Loss

What is the relationship between the loss of an animal species that the term ‘extinction’ marks, and the loss of an animal culture? They are clearly related phenomena, but when are they equivalent? The loss of the species necessarily means the loss of its whole way of life, of course, but in an important sense, we must grapple with what it means that an animal’s cultural life can be lost or profoundly diminished while the species biologically endures. It might be objected that this framing only makes sense when we understand ‘animal cultures’ as isolatable behavioural units, but this is precisely the point: recent calls for more holistic, interconnected approaches to conservation have learned from a troubled history that made evident the errors of disconnection. The ironic lesson of numerous experiments in captivity and conservation has been that treating animals in reductively biological terms, pulling apart the complex web of their way of being and its embedded biocultural and ecological relationships, only has the effect of producing ‘bare’ (Agamben, 2004) or ‘wounded’ (Chrulew, 2011) life, divided from itself, the very impoverishment of which is then taken to demonstrate that the conservation of the organism without culture – just like that of the organism without environment – can hardly be called conservation at all.

As noted above, work in extinction studies has often understood biodiversity loss as a process of unravelling: an unravelling of ecological and multispecies relationships, but also of distinctive ways of being. Clearly, socially learned traditions can be an important part of this unravelling process, contributing to or preventing it in various ways, as well as themselves succumbing to it. Whooping crane migration again offers an instructive example. If this behaviour (and the cultural knowledge it requires) disappears, and subsequent generations of birds no longer know how, when, or where to migrate, there is an important sense in which this should be understood as a kind of diminishment of the world that matters profoundly, even if living, breathing whooping cranes themselves endure. There are, of course, ecological and conservation arguments that might be made in favour of migration: that the behaviour enhances the reproduction or survival of the species, or that birds play an ecologically significant role in their summer habitat. But setting these kinds of instrumental arguments aside (which may or may not be compelling in different cases), important questions remain as to the intrinsic value of migration traditions themselves, as well as the broader ways of life they enable. Is the existence of a migratory way of life for this species itself something that should be safeguarded? In what way, and at what cost?

A reductive, biogenetic notion of what a species is might see animals living in a zoo as a successful form of conservation. A more ecological understanding might insist that they be out in the world, taking up their vital relationships with other species. The position advanced in extinction studies has generally built on that ecological argument but taken it further to insist that the significant relationships that species have are also inherently cultural, social, and experiential: including both the important relationships they have with other species, including human communities, and with their own kind across generations, as well as their own thicker modes of meaningfully being in the world. This understanding of species life sets a much higher bar for conservation success. Indeed, it might shift the register beyond the regular bounds of conservation biology to encompass questions and approaches usually found more in other domains such as heritage studies (Harrison, 2015; Harrison and Rose, 2010) and political and ethical theory (Celermajer et al., 2022; Donaldson and Kymlicka, 2011; Palmer, 2010) – fields which will themselves be equally challenged by these new phenomena and processes demanding consideration.

This is an understanding in which animal cultures are definitively more than resources for survival. Instead, they are intricately entangled parts of how a species or population endures in the world as a way of life, not merely as a population of biological organisms. We need to hold onto this fuller sense of animal cultures. Doing so means holding on to two ideas seemingly in tension: to insist on culture as something that is both an integrated part of the larger process that is a species and something that is valuable in and of itself. Importantly, such an approach must also refuse to buy into static notions of animal and species life. We live in a dynamic and changing world (one that has always been that way). As such, not all change is loss, and not all loss claims the same sorrow. This is a point that has been made strongly in discussions of the cultural lives of human communities, and the need to respect their ongoing dynamism (Clifford, 1988; Mitchell, 1995; Nelson et al., 2013). As James Clifford famously put it, ‘cultures do not hold still for their portraits’ (Clifford, 1986: 10). In this context, the fact that a species like the whooping crane that once migrated no longer does so does not necessarily constitute a loss. Instead, for it to be recognised as such, we must be able to make the further step of articulating why this particular change constitutes a diminishment of that way of life, on what terms, and for whom (a task that must always be undertaken with the humility that arises from the fact that many of these dynamics remain beyond the limits of our perception and understanding).

Multispecies Cultures and Biocultural Loss

As we’ve seen, research in extinction studies has emphasised the need to understand extinction as a process of unravelling that draws in a diverse range of living beings. As one species declines, the many others that live alongside it are impacted, for good and ill. In contrast to much of the research in ecology, which focuses exclusively on the loss of ‘nature’ (or the nonhuman world) considered in terms of resources, biodiversity, or otherwise, work in extinction studies emphasises that this is a process of biocultural loss, that impacts profoundly on entangled communities of humans and nonhumans in diverse and unequal ways.

Taking animal cultures seriously expands this conversation about the other living beings caught up in extinction events in fascinating ways. To date, a lot of work in extinction studies has documented the impacts of the loss of plant and animal species on local human communities, including their livelihoods and cultural practices (Chao, 2022; De Vos, 2017; Hatley, 2017; Rose, 2011; van Dooren, 2011, 2022). Some of this work has also documented the loss and transformation of animal cultures themselves (Reinert, 2013; van Dooren, 2016). What has been less frequently explored is the way in which the loss or transformation of cultural traditions (human or other animal) might ripple out to impact not just on other species but on other cultural traditions. ³ In taking up this question we would be asking: how do these differing cultural forms emerge, endure, and expire in relationship with one another?

For example, consider the migration of many birds and whales, both of which might be classed as cultural behaviours that are maintained across generations through intimate, embodied processes of social learning. ⁴ As these animals move through different land- and sea-scapes, they come into contact with various human communities, and they are consequently woven into the cultural lives of these people. They might be observed, hunted, made the subject of stories, songs, dances, and more. ⁵ In these cases, if one cultural behaviour were to cease or dramatically change, the others would be impacted upon. This would be particularly apparent in a case in which it was the animal migratory behaviour that stopped or was significantly redirected, potentially leading to the breakdown of the human cultural practices associated with that now absent species. From one perspective, these are stories of the co-formation and co-loss of human and other animal cultures, two streams of cultural life that might otherwise be conceptualised in isolation. From another perspective, however, they are stories of multispecies cultural life and death. What does it mean to recognise these kinds of interactions as sites of shared multispecies culture?

In thinking through this question, the case of honeyguide-human cooperation offers an instructive contrast. In this now well-documented mutualistic association, Greater Honeyguide birds work with humans to gain access to bee hives in parts of Africa. Knowledgeable humans call out to the birds, who approach them and respond with a call of their own before, as their name implies, guiding the humans to the bees. Humans are provided with up-to-date information on the location of hives so they can harvest honey, while the birds are able to eat the discarded wax (the bees, on the other hand, have their hive and home destroyed).

We raise this example here because it is frequently offered as a readily identifiable example of a mutualistic, interspecies cultural practice: both species come together to co-produce a shared space of interaction that relies on both of their ongoing participation to endure (van der Wal et al., 2022). In contrast to the more recognisable honeyguide story, the migrating bird and whale examples offered above help us to see that a great many, less obvious, cultural practices are always already multispecies. In fact, there is no such thing as a single-species culture (Lestel, 2003). Importantly, these forms of multispecies cultural entanglement give rise to particular kinds of interdependence and fragility in which there is significant potential for the loss or transformation of one cultural practice to ripple outwards to impact on others in a range of ways. As Lestel notes, what this means in a time of extinctions is that what is often presented as a ‘loss of biodiversity’ is better conceived, in a fuller and more encompassing sense, as ‘the withering of shared life’ (Lestel, 2013).

Extinction studies has an important role to play in understanding and storying these processes of withering at the edge of extinction. The interdisciplinary, biocultural approach of extinction studies makes possible a more expansive articulation of the significance of multispecies cultures that encompasses not only their (often unacknowledged) widespread entanglements and mutualities, but also the (often invisible) rippling effects of cultural loss and unravelling.

Conclusion

Research on animal cultures in the natural sciences has developed significantly over the past several decades, opening up new opportunities to understand and appreciate the complex social dynamics of animal lives. This is, in many ways, a hopeful situation. But not in any straightforward way. This picture of animal life is emerging at a time when many species are disappearing along with the rich and intricate ways of life that they have developed in and through their ongoing evolutionary, developmental, experiential, and cultural interactions. Meanwhile, the ways of life of those species that survive – including many of those who are learning to thrive in the Anthropocene – are also undergoing significant transformations, for better and worse. While research on animal cultures offers important opportunities for deepening our appreciation of animal life as well as practical conservation interventions, it also carries with it the potential for new kinds of surveillance, control, harm and killing. Underlying this whole situation, we have argued, are problematic frameworks of understanding and studying animal cultures that tend to reduce it to specific, operationalisable, socially learned behaviours. In this context, there is a vital role for researchers in extinction studies and the broader environmental humanities to creatively and collaboratively work to thicken understandings of what (animal) culture is, why and how it matters, and what this recognition might demand of us all in a time of escalating loss.