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On the controversy over non-human culture: The reasons for disagreement and possible directions toward consensus Murillo Pagnotta ∗ Instituto de Psicologia, Universidade de São Paulo, Av. Prof. Mello Moraes 1721, São Paulo 05508-030, Brazil

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Article history: Available online xxx Keywords: Anthropology Culture Ethology Psychology

a b s t r a c t In recent decades, animal behaviorists have been using the term culture in relation to non-human animals, starting a controversy with social scientists that is still far from cooling down. I investigated the meanings of the term culture as used by social and cultural anthropologists, and also its recent use by ethologists, in order to better understand this controversy and identify possible paths that might lead to a consensus. I argue that disagreements in the level of theories involve definitions of culture and theories of behavioral development, while disagreements in the level of worldviews include the acceptance or rejection of the idea of a radical distinction between humans and other animals. Reaching a synthetic approach to (human and non-human) animal behavior depends on constructing a consensus in both levels. It is also necessary to discuss how to include symbolic communication in a comparative perspective. I conclude that this might lead to the abandonment or reconstruction of the related dichotomies of nature–culture, innate–acquired and gene–environment. This article is part of a Special Issue entitled: Neotropical Behaviour. © 2014 Elsevier B.V. All rights reserved.

1. Introduction In this article, I will look at the attribution of culture to non-human animals. My main objective is to clarify the differences between what anthropologists (specially social and cultural anthropologists) and animal behaviorists (including biologists and psychologists, regardless of their department association, henceforth simply ethologists) mean when they say culture, and indicate where a consensus might be constructed. First, I will stick to the academic distinction and overlook many differences between specific authors, so as to establish two deliberately course-grained statements about culture that will allow us to identify what is a central point of disagreement between ethologists and anthropologists – namely, the symbolic question. Next, I will analyze the controversy in terms of different levels of disagreement, and this analytical strategy will help us identify possible directions for collectively constructing a synthetic approach to behavior that includes humans and other animals. One of the epistemological foundations of Modernity (roughly, the historical period after the Middle Ages) has been the belief that

∗ Correspondence to: Centre for Social Learning & Cognitive Evolution, School of Biology, Harold Mitchell Building, University of St Andrews, St Andrews, Fife KY16 9TH, U.K. E-mail addresses: [email protected], [email protected]

the human condition corresponds to a radical separation from the rest of nature, often described as due to culture. The term culture itself has been used in a variety of different contexts and there is no consensual definition (e.g., compare Geertz, 1973; Ingold, 2000; Kroeber, 1917; Lévi-Strauss, 1963; Tylor, 1874). Nevertheless, it had generally been uncontentious that, whatever it is that makes humans unique, we could say that it belonged to the cultural (or social) domain. This is reflected in the historical separation between natural and social scientists. Natural and social scientists might agree that humans are animals, and that it is epistemologically legitimate to study human morphology, physiology, and even some aspects of human behavior (some might say instincts) using the same methods, concepts and philosophical commitments that have proven so useful in the study of other biological phenomena. But, according to the still dominant (but not unquestioned) view in the social sciences, there always remains something – namely culture – that, perhaps because of its emergent properties (Kroeber, 1917), or because of its symbolic foundation (White, 1949), escapes, in a sense, the biological realm. Hence, humans would not just be unique, as any other species is, but unique in a special way, something more than just another unique animal species. The difference lies, one could assume, in how our development is influenced by the social environment we grow into, and in the (exclusively human?) cognitive abilities that underlie this individual development as well as collective phenomena such as culture and history.

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However, it is now clear to ethologists that other animal species also exhibit behavioral diversity between populations that must be explained in terms of learning in a social context. For decades now, many ethologists have been using the word culture to describe this phenomenon (e.g. Boesch and Tomasello, 1998; Kawamura, 1959; Laland and Galef, 2009a; McGrew and Tutin, 1978; Nishida, 1987; Whiten et al., 1999). If we accept these claims, and culture is not a golden barrier any more (Boesch, 2003), human behavior would be even closer to the epistemological framework used to study animal behavior in general. And also, at least some nonhuman behavior would be closer to the epistemological framework used in the social sciences to study human behavior. But, if culture is used to describe the difference between humans and other animals, what could it possibly mean to say that chimpanzees (Whiten et al., 1999), bonobos (Hohmann and Fruth, 2003), orangutans (Van Schaik et al., 2003), whales (Rendell and Whitehead, 2001) or birds (Grant and Grant, 1996) have culture? How are we to distinguish human uniqueness, while acknowledging our animal condition? Should we deliberately abandon the concept of culture, or maybe reconstruct the old academic separation? Is it possible to find a common ground for the social and natural sciences? Instead of arguing that natural scientists should conquer the territory of the social sciences, or to insist on the absolute separation of humanity from animality, I consider that the dialog and the active search for consensus are more desirable and productive. According to this third-way view, both social and natural scientists might indeed have interesting and important contributions to make to the understanding of human behavior. A consensual framework would have to offer a solid base for investigation both of things of nature and of things of society and culture without even assuming such dichotomy as a given. Because of the historical division of labor between the natural and the social sciences, and the immense amount of cumulative theoretical elaborations within different disciplines and their multiple subfields, the search for a synthesis demands wide collaboration between scholars holding different and often incompatible theories and worldviews. In other words, this might as well demand a radical reconstruction of long established, individually and collectively held beliefs, commitments, prejudices and motivations. Here I review the meanings of the notion of culture for social/cultural anthropologists and for ethologists (as defined above). I had two starting points. On one hand I reviewed the works of classical and contemporary authors in the history of Anthropology that might help us understand the debate on which I am focusing. On the other hand I reviewed the history of the attribution of culture to non-human animals from pioneers such as Kinji Imanishi to the present. I will present my findings and then clarify the main reasons for disagreement between them and point to possible directions toward consensus.

2. The meanings of culture for anthropologists In 1896, Edward Burnett Tylor became the first Professor of Anthropology at Oxford University, in a time where evolutionary ideas still dominated the newborn discipline. Culture was used in the singular, indicating the télos of a universal development with its most advanced stage to be found in, not surprisingly, the Europeans (or maybe just the Victorian Englishman). In the beginning of his Primitive Culture, published in 1871, Tylor established the first formal or explicit anthropological definition of culture as “that complex whole which includes knowledge, belief, art, morals, law, custom, and any other capabilities and habits acquired by man as a member of society” (Tylor, 1874, p. 1). According to Washburn and Benedict (1979), the purpose of this kind of definition is not to propose a testable hypothesis (as ethologists would like to find),

but “to define a field for investigation by a positive listing of the behaviors that are of interest” (163). Franz Boas, intellectually raised in the German academic environment, developed on the idea of the humanist Johann Gottfried von Herder, who had used the term culture somewhat differently, to indicate the particular way of life of a people, including the spiritual values that unite them and distinguish them from other peoples. In this sense the term was to be used in the plural (the cultures of different people), in opposition to its use in the singular (the Culture), and with no claims of superiority. It is this use of the term that constitutes the foundation of its modern meaning in Anthropology. According to Boas, “Culture embraces all the manifestations of social habits of a community, the reactions of the individual as affected by the habits of the group in which he lives, and the products of human activities as determined by these habits.” (Boas, 1930, p. 79) In 1952, when Alfred L. Kroeber and Clyde Kluckhohn reviewed 164 definitions and hundreds of propositions about culture, they concluded that, even if anthropologists still had “no full theory of culture. . . [there was at least a] fairly-well delineated concept, and it is possible to enumerate conceptual elements embraced within that master concept” (Kroeber and Kluckhohn, 1952, p. 181). One of these conceptual elements was the idea that culture is exclusively human, and this privilege was said to be granted by our equally exclusive capacity to use symbols. Many anthropologists also supported the idea that cultural phenomena are emergent, thus corresponding to a level of analysis on their own right, beyond Biology and Psychology – e.g., that they are “superorganic” (Kroeber, 1917), or “exosomatic” (White, 1949). This was not supposed to mean that Biology and Psychology are not important in the study of human behavior. What was implied was the idea that human populations do not show fundamental biological or psychological differences, and that the (cultural) behavioral diversity must be explained in terms of learning and history. Another conceptual element in Kroeber and Kluckhohn’s (1952) review was the idea that culture is not individual behavior, but patterns of behavior abstracted from them. A culture could be understood in a descriptive sense (it is what people do), and also in a normative sense (it is what people are expected to do). It was manifested in material objects, ideas, attitudes and especially values, which “provide the only basis for the fully intelligible comprehension of culture, because the actual organization of all cultures is primarily in terms of their values” (Kroeber and Kluckhohn, 1952, p. 173). Despite its influence, from then on many anthropologists reacted to this culture paradigm, reevaluating the whole nature/culture distinction. In the 1950s, Claude Lévi-Strauss, the founder of Structural Anthropology, argued that, although the separation between nature and culture had had “the force almost of dogma” (Lévi-Strauss, 1992, p. 28) within Anthropology, culture should be taken as a heuristic tool and a relational concept – i.e., one that does not point to some concrete thing but to a differential relation between two ethnographic groups: What is called a ‘culture’ is a fragment of humanity which, from the point of view of the research at hand and of the scale on which the latter is carried out, presents significant discontinuities in relation to the rest of humanity. [. . .] Accordingly, the same set of individuals may be considered to be parts of many different cultural contexts: universal, continental, national, regional, local, etc., as well as familial, occupational, religious, political, etc. (Lévi-Strauss, 1963, p. 295) Another reaction in the 1950s was the rise of Symbolic Anthropology. Clifford Geertz, one of its exponents, argued against what he called a “’stratigraphic’ conception of the relations between biological, psychological, social and cultural factors”(Geertz, 1973, p. 37),

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each corresponding to a relatively independent level of analysis, in an explicit correspondence to established academic disciplines. Instead, he proposed a synthetic conception, in which those factors are treated as variables in the same analytical system. Abandoning the idea of a superorganic reality, of abstracted patterns of behavior, Geertz argued that culture was the context that allowed for a “thick description” (i.e., one that goes beyond the obvious and superficial relations) of events, behaviors, institutions and social processes. This context is symbolic systems, or historically created systems of meaning – in the plural, for there are incoherencies and incompatibilities within each culture, and not a harmonious complex whole. The majority of these symbols are already in place, publicly available, when a person is born, and take part in her development, as she uses them to convey meaning and orient her in her experiences. Since our nervous system grows – and, in a broader temporal perspective, evolved – in close interaction with these cultural systems, “it is incapable of directing our behavior or organizing our experience without the guidance provided by systems of significant symbols” (Geertz, 1973, p. 49). More recently, Tim Ingold suggested that instead of taking the idea of a mind (or the individual) separated from the world as our point of departure, we should take “the human condition to be that of a being immersed from the start, like other creatures, in an active, practical and perceptual engagement with constituents of the dwelt-in world” (Ingold, 2000, p. 43). In his perspective, “it makes no sense to speak of ‘culture’ as an independent body of context-free knowledge” (Ingold, 2000, p. 37), or “as an internalised system of rules and meanings as distinct from manifest behavior patterns and their artefactual products” (Ingold, 2000, p. 159, his emphasis). Instead, human (cultural) behavioral variation should be understood as variation in the skills embodied in the process of development of each human organism engaged with his environment. I will now take a step back and pay more attention to similarities than to differences between these authors. From this distance, I will formulate provisional statements that do not intend to make justice to the theoretical diversity, but that might be sufficiently precise to describe what anthropologists have in mind when they say culture. This will allow us to identify a central question in the controversy. In general terms, it might be said that among anthropologists the word culture evokes (1) the capacity to create, manipulate and share symbols – including linguistic symbols –, which supports (2) the existence, between individuals, of shared patterns of behavior, meanings and engagement with the constituents of their environment, and (3) that depend on learning in a social context to develop.

3. The attribution of culture to non-human animals With regard to ethologists, it is perhaps not surprising that the history of the recent attribution of culture to non-humans began far away from Europe, both geographically and philosophically. Raised in the general atmosphere of Buddhism, biologist Kinji Imanishi (1902–1992), the founder of Japanese primatology, did not believe in a radical separation between humans and other living things: “we cannot regard animals as mere complicated automata nor humans as special creations of an omnipotent and omniscient deity” (Imanishi, 2002, p. 6). Instead, he emphasized the unity and harmony of nature (humans included), the integrated and inseparable system formed by organism and its environment, and the need to recognize the simultaneous existence of similarities and differences among things. Accordingly, since there is no radical distinction, he saw no problem in considering humans as another animal species, and thus to consider human phenomena from a biological point of view: “This neither relegates humans to an animal

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level nor elevates animals to a human level, but allows us to discuss both on the same basis” (Imanishi, 2002, p. 4). In an essay written in 1952 (in Japanese and with no translation available), Imanishi (1952) and Nakamura and Nishida, 2006) argued that other animals might exhibit something like culture. But instead of using the word bunka, the usual Japanese translation to culture, he coined a neologism based on the sound of the English word: kaluchua. According to his students, “[Imanishi] said that kaluchua was much broader than bunka and was non-hereditary, acquired behavior that was acknowledged socially” (Nakamura and Nishida, 2006, p. 35). It was because of his conviction that the comparative study would help us trace the evolution of human society that Imanishi decided to study other species applying methods such as individual identification of animals and long-term observations of their behaviors and social relations. After World War II, and returning from Mongolia where he had been studying wild horses, Imanishi decided to study the Japanese monkeys (Macaca fuscata) in Koshima Island. Habituation with food provisioning began in 1952. In September 1953, the (now famous) young monkey Imo was seen washing a sweet potato in a nearby stream before eating it, and primatologists studied closely the spread of this novel behavior within the group. Following Imanishi, the first articles written in Japanese used kaluchua to refer to it (Nakamura and Nishida, 2006). The same precaution that had led Imanishi to coin a new word to distinguish between human and non-human behavior explains the use of prefixes in the early articles written in English, such as “subculture” (Kawamura, 1959) and “preculture” (Kawai, 1965). Later, Kawai explained that the prefixes were intended explicitly to take the differences into consideration, while defending an evolutionary approach: “We must not overestimate the situation and say that ‘monkeys have culture’, and then confuse it with human culture” (Hirata et al., 2001, p. 490). Field studies in Africa, starting in the sixties with Imanishi, Jane Goodall and others, revealed that the behavior of the chimpanzee, our most closely related species, was much more complex, diverse, and humanlike (or conversely, that human behavior was much more apelike) than ever realized. Ethologists gradually learned that chimpanzees live in social units (or communities) where they establish lifelong social relationships (Nishida, 1968); use various tools (Goodall, 1964), including hammers to open hard nuts (Boesch and Boesch, 1981, 1984); hunt cooperatively or collectively (Boesch and Boesch, 1989); and might even actively teach in some circumstances (Boesch, 1991). In 1975, William McGrew and Caroline Tutin observed a peculiar grooming behavior in the Mahale Mountains (study site coordinated by Itani, former student of Imanishi), that was absent in Gombe, a mere 50 km away. By that time, “tradition” was an established term in ethology that referred to behavioral differences (between members of a species) that could not be readily explained in terms of genetic or ecological differences (Galef, 2006). The best known examples at the time were behaviors explicitly directed to resources. Yet this behavior seemed like an arbitrary social tradition with no direct relation to resource exploitation, and the article that described it (McGrew and Tutin, 1978) was, according to Laland and Galef (2009b, p. 4), “the first in the modern era to directly address the question of the relationship between the traditions of animals and the culture of humans”. Following an argument by Galef (1976, p. 77) the authors suggested that there were three possible explanations for such a difference between the two populations: it could be due to (1) some genetic difference, or (2) to different individually learned responses to some different environmental condition, or (3) to the emergence of shared behavioral patterns through social learning. They favored the third hypothesis and asked: “Do such

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social traditions in animals satisfy accepted anthropological criteria so that they may be termed cultural?” (McGrew and Tutin, 1978). They defended the need for an operational definition of culture, one that would specify the necessary and sufficient conditions for a behavioral phenomenon to be considered cultural, and that would allow for a comparative approach. In their conclusion, the authors defended the legitimacy of the use of the (by them redefined) term in relation to chimpanzees. Anthropologists Washburn and Benedict (1979) replied that despite the similarities, articulate language makes human culture unique, and that the term should remain restricted to our species. Years later, anthropologist Tim Ingold would make a similar point, stating that “Socio-cultural anthropologists might be unwilling to attribute culture to chimpanzees until and unless the acts of the apes could be shown to have meaning for them.” (Tim Ingold, personal communication cited in McGrew, 1992, p. 86). Meaning, in this sense, exists only through discursive interactions, which allow for the entanglement of different aspects of human life activities in complex sets of associations. McGrew (1992, 2003, 2004) continued his discussion with Anthropology, arguing for the dissociation between culture and articulate language. The investigation of the (so-called) cultural behavioral phenomena in non-human animals such as primates, birds and cetaceans continued notwithstanding the lack of consensus about the use of the term to describe it (for different positions, see Laland and Galef, 2009a). If we pay more attention to similarities than to differences, it might be provisionally said that among ethologists the word culture evokes (1) the existence, between individuals, of shared patterns of behavior that (2) depend on learning in a social context to develop. Comparing my two provisional statements on the meaning of culture to both anthropologists and ethologists, it should be clear that the question of symbols, meanings and articulate language is central to the anthropological notion but is absent in the ethological notion. In order to render it equally applicable to humans and nonhumans, most ethologists deal with this question by unmaking – by definition – the relation between culture and language. My first conclusion is that any attempt to formulate a consensus will have to include a proposal for dealing with the symbolic question in a comparative, thus evolutionary way. At first sight, it might seem to be just a matter of definition. But the situation is more complex, as I argue next.

4. The different reasons for disagreement Anthropologists and ethologists agree or disagree with each other over many different issues. For example, two ethologists might debate whether they should name the phenomenon of interest as “culture” (McGrew, 2004) or “tradition” (Galef, 2009). But in this case, the two signifiers mean basically the same thing (namely, a shared pattern of behavior due to social learning). Also, since McGrew and Galef are both committed to Biology, they share many aspects of their epistemological and theoretical foundations; their disagreement is not so big. On the other hand, the debate between primatologists Boesch and Tomasello (1998) and anthropologist Ingold (1998) reveals disagreements of another order and must be understood from another perspective. Not only do they attribute different meanings to the shared signifier, but they also differ in how they deal with the distinction between nature and culture, and with the development of animal behavior. In a general way, authors might disagree over descriptions, explanations, theories and worldviews. These four levels of elaborations are absolutely not independent from each other. Every description and every explanation is formulated with terms that

refer to a specific theory. And every theory is supported by a specific epistemological foundation and expresses a particular worldview. 4.1. Descriptions The most basic level of disagreement is that in which the contentious elaborations are descriptions of phenomena made within a shared general theoretical context. If there is no reason to suppose fraud or methodological error, there will generally be no conflict at this level. I found no such accusations in my study, so that the controversy does not bring into question the precision of ethnographic descriptions made by anthropologists, or the behavioral descriptions made by ethologists. No one doubts that Malinowski really did see the exchange of bracelets and other items taking place in the Trobriand Islands in Papua New Guinea (Malinowski, 1932), or that the Japanese monkeys really do wash their sweet potatoes (Nishida, 1987), or that the chimpanzees in Mahale really do groom each others’ armpits (McGrew and Tutin, 1978). 4.2. Explanations In the second level of disagreement, the contentious elaborations are rival explanations of a phenomenon formulated within the same theoretical framework and with consensual description. In this case, scholars from the same research program debate over the best explanation for a certain empirical observation, but this does not jeopardize the “hard core” of their research program (Lakatos, 1970). It is possible that many such disagreements are resolved internally, even before the publication of articles. 4.3. Theories The third level of disagreement reveals conflicts between different, incommensurate theories that deal with the same phenomenon (more precisely, phenomena perceived as equivalent), each one with its own terminology, questions of interest, tools for investigation and criteria to decide which are the best explanations. With regard to the addressed controversy, some questions related to this level would be: What is culture? Does culture depend on articulate language? How do behaviors develop? Are “innate” and “acquired” reasonable explanatory categories or should they be abandoned? This level allows us to clarify some aspects on the controversy, since it puts into question whether the redefinition of the term culture by ethologists corresponds to an advance in the search for a synthetic theoretical approach to animal behavior that includes humans. I will develop this further before turning to the last level. McGrew and Tutin (1978) based their argument on a passage in which Kroeber (1928) points out what is missing in the behavior of the chimpanzees to be considered cultural. But contrary to what the authors suggest, Kroeber might not be implicitly defending a checklist-like, operational definition of culture. Instead, he was (1) explicitly affirming his belief in the gap that separates us even from our closest living relatives and (2) implicitly defending his view of culture as “superorganic” (Kroeber, 1917; Kroeber and Kluckhohn, 1952), dependent on human symbolic and linguistic skills, and thus defending the autonomy of Anthropology as an academic discipline. In their response to the young primatologists, Washburn and Benedict (1979) stated that the anthropological concept of culture was never intended to be an empirically testable hypothesis. That may be the reason why, after looking among anthropologists for a suitable working definition, McGrew states that “the ubiquity of frustratingly vague, untestable, all-inclusive, and anthropocentric definitions holds” (McGrew, 2004, p. 16), concluding that “students of cultural primatology need not bother about seeking consensus

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but, instead, scholars may set their own terms, so long as these are explicit, comprehensive, and clear” (McGrew, 2004, p. 52). Others decided to avoid the discussion with anthropologists by using the term tradition instead of culture (e.g. Fragaszy and Perry, 2003). Both strategies share the legitimate goal of focusing on the phenomenon they are interested in – namely, the existence of different behavioral patterns between different populations of a certain animal species – and might have indeed been a necessary step to justify and draw attention to the ethological study of this phenomenon and its resemblance to human (cultural) behavior. Its side effect, however, was that it tended to postpone the discussion with social sciences, especially with those contemporary authors that criticize the culture paradigm even in relation to humans (e.g. Descola and Pálsson, 1996; Latour, 1993, 2005; Ingold, 2000, 2004). Originally the term culture was used to defend the legitimacy of the anthropological discipline in its struggle for survival in the 19th century academic environment, amidst its competitors, Biology and Psychology. It expressed less a testable hypothesis than the belief in a radical distinction. But with our increasing knowledge of animal behavior, and the accumulated criticisms to western modern epistemology, it is not clear whether the belief in such a radical distinction is still defensible. This brings about the (philosophical) question of how to understand – and, possibly, reconstruct – the old, modern distinction between the social and the natural sciences. This discussion may be provisionally set aside, but it is inescapable. Thus, ethologists and anthropologists share the signifier, but not the signified, and the use of the term culture neither refers to the same concept nor does it imply the existence of shared, consensual theories of behavior. At least in this stage of the controversy, the redefinition of the term by ethologists rendered it fundamentally incompatible with the anthropological meaning and the more recent debates within Anthropology. Also, until we approach the symbolic question in a comparative perspective, natural and social scientists might not reach a theoretical consensus about (1) how to define culture; (2) whether its extension (that is, the number of species to which it may refer) should be expanded; (3) in so doing, how to explicate the similarities and differences between the culture of humans, chimpanzees, birds, rats and whatever other species; or even (4) if the term should be completely abandoned – perhaps even in relation to humans. 4.4. Worldviews The fourth level of disagreement indicates a conflict between worldviews (Weltanschauung, philosophical and epistemological foundations, metaphysics) to which different scholars are committed, and that might hinder mutual understanding and the making of a consensus. At this level, the relevant questions in the controversy are such as Do humans differ radically from non-human animals? How should we understand the relation between Anthropology and Biology? Should we commit ourselves to reach a consensus and a synthesis between the natural and social sciences? 5. Possible directions toward consensus Now I will once again take a step back and look from a broader perspective at the similarities and differences between different authors from both sides of the controversy, in terms of their theories and worldviews. In the level of worldviews, I have identified two main attitudes. One, which is becoming ever more obsolete, treats culture as part of a duality nature–culture, which expresses the idea that humans are fundamentally distinct from the rest of nature and that the cultural (human) phenomena are to be investigated by special disciplines (Anthropology, Sociology) that are relatively

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independent from Biology and Psychology. Most anthropologists, since the beginnings of the discipline in the 19th century until the sixties fit well this description, as does the widely cited review of Kroeber and Kluckhohn (1952). But several contemporary authors question the legitimacy of this duality as a foundation for Anthropology (e.g., see contributions in Descola and Pálsson, 1996). Most naturalists from Aristotle until the 19th century also fit well this description. It was only with the consolidation of the evolutionary ideas of Charles Darwin that the idea of the uniqueness of human beings began to be seriously questioned within natural sciences. The opposing attitude asserts that humans are not fundamentally distinct from other animals and that the study of human behavior is (logically) a part of the study of animal behavior. Thus, it would be legitimate to investigate human (cultural) phenomena from a comparative and evolutionary perspective, and founded in theories consistent with what is agreed upon in Biology. This is the first step to take in the direction toward a consensus. In the level of theories, the term culture refers to the way we describe, investigate and explain behaviors of humans and other animals. One theoretical orientation views ontogeny as a dualistic process: some aspects of the phenotype are said to be due to genes and others due to the environment. For some (e.g., Dawkins, 1976), culture would be the extreme example of environmentally determined behavior. Nowadays many authors seem to agree that this dualistic view of development is obsolete, and that genes and environment interact in such a way that renders it impossible to distinguish between two opposing and complementary causal pathways. Accordingly, criticisms and attempts to find alternatives have been accumulating for decades within Biology and Psychology (e.g. Bateson and Martin, 1999; Fox-Keller, 2010; Gibson, 1979; Gottlieb, 1992; Hogan, 2001; Ingold, 2004; Kuo, 1967; Lehrman, 1953; Lewontin, 1983; Oyama, 1985; Oyama et al., 2001; Schneirla, 1956). What all these authors have in common is (1) their emphasis in looking closely at development and (2) the explicit rejection of the distinction between innate and acquired, but this is not supposed to indicate theoretical homogeneity or a coherent and established school of thought. But since there is a growing consensus among both social and natural scientists, on the inadequacy of the dichotomies nature–culture, innate–acquired and gene–environment, it is probable that the consensus will be founded in behavioral theories that do not make use of these dualisms in the study of (human and non-human) behavior and its development.

6. Conclusions Do non-human animals have culture? Instead of trying to answer it with a yes or no, I have tried to better understand the very question and identify some of the reasons for disagreement. Assuming that (1) humans are animals; (2) since there is phylogenetic continuity, it is legitimate to do comparative, evolutionary studies; (3) we are willing to reconstruct the distinction between social and natural sciences; then we might agree that it would be desirable and coherent to search for a common terminology and behavioral theories that would allow for comparative research. I concluded that the controversy on the question of non-human culture is not only a matter of formulating an adequate definition. If my argument is valid, it follows that anthropologists interested in constructing this consensus will have to take seriously the possibility of investigating human behaviors in a comparative, thus evolutionary perspective. For asserting simultaneously that humans are animals and that the study of human behavior is independent from Biology is, to say the least, inconsistent. On the other hand, ethologists interested in constructing this consensus will have to consider explicitly both the similarities and

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Please cite this article in press as: Pagnotta, M., On the controversy over non-human culture: The reasons for disagreement and possible directions toward consensus. Behav. Process. (2014), http://dx.doi.org/10.1016/j.beproc.2014.04.008