The idea that culture is important for nonhumans, including whales, has a history of controversy. In the 1930s–40s, biology was given a strong theo­retical basis in the form of evolution through natural selection—natural selection as first suggested by Charles Darwin and Alfred Russel Wallace and then formalized with genes as the units of selection in the modern syn­thesis. The modern synthesis was not particularly about behavior, but be­havioral theorists in and around the 1970s realized it could be applied to behavior as well as morphological, physiological, or anatomical features. This new field was called behavioral ecology or, largely in the United States, sociobiology. Advocated comprehensively in E. O. Wilson’s book "Sociobiol­ogy" and summarized eloquently by Richard Dawkins in "The Selfish Gene," be­havioral ecology made a fine job of explaining why animals do what they do. Its application to human behavior was, and is, controversial. For the study of nonhuman behavior, however, behavioral ecology became a hugely successful scientific paradigm. From the 1980s onward, scientific papers describing the behavior of animals invariably started and ended with how the research was situated within the theory of behavioral ecology. We, and most of our scientific colleagues, found the theory very appealing and felt it well explained the behavior of animals. In the field of animal be­havior, behavioral ecology became “normal science,” in the terminology of the philosopher of science, Thomas Kuhn. Suggesting that culture could be a major driver of the behavior of nonhumans challenges this paradigm— making it “revolutionary science,” according to Kuhn—and, as with other challenges, was resisted. However, in contrast to the opposition facing most other scientific revolutions, the attacks are not coming from the stalwarts of “normal science.” Since the inception of their theory, behavioral ecolo­gists and sociobiologists have largely accepted the possibility that culture might have an important role in determining behavior, along with genes.

E. O. Wilson, for instance, cowrote "Genes, Minds and Culture: The Coevolu­tionary Process" and Richard Dawkins coined the term “meme,” the cultural analog of the gene.

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This is not to say that proposing culture as an explanation for animal behavior does not meet resistance from our closest colleagues, but it does so only from the angle of questioning what the evidence is that a particu­lar behavior results from some kind of social information. There are now, however, enough solidly demonstrated examples that this does in fact hap­pen in many species for the study of social learning to be accepted as a valid and growing field within mainstream animal behavior science. No, while behavioral ecologists may question the evidence and suggest alternative explanations, they are generally not appalled by the very notion of chim­panzee or whale culture. The fiercest critics come mostly from anthro­pology and psychology. Here, it is the very concept of animal culture that is anathema, not the nature of the evidence. It is part of the paradigm in most of the social sciences, insofar as the social sciences have paradigms, that humans are unique in having culture or, at least, in being overwhelmingly cultured. Culture in other species, if it exists, is an epiphenomenon, not ter­ribly important. It is the challenge to this paradigm that is being resisted.

Critics of all stripes argue against the evidence put forward. They pick away at the (necessarily) spotty evidence from field studies, suggesting that this or that pattern of behavior could have arisen genetically or through environmental correlations. In their laboratory experiments, chimpanzees don’t imitate and rats don’t teach—thus: no culture. On the other side of the debate, field scientists are convinced that culture is a major part of the lives of the animals that they study, but how can they show it?

People have thought about nonhumans having culture for a long time. Aristotle noted that at least some birdsong was learned. Darwin thought many animals possessed “inherited habits,” and although he did not know how inheritance worked, his conception of these inherited habits was very similar to what we now think of as culture. Following Darwin, many of those who studied the behavior of animals in the late nineteenth and early twentieth centuries believed that socially learned traditions were impor­tant in shaping the behavior of at least birds and mammals. However, once genes had become central to biology in the modern synthesis, thoughts that culture might have a role in animals other than humans faded. For a while it was nearly all about genes.

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After the end of the Second World War, ethologists began to study the be­havior of animals quite broadly and rigorously, and the trend against non­human culture reversed. These studies at first indicated and then showed that in two very different kinds of animals, social learning—and “culture” for those who wished to use that term—is an important determinant of what animals really do. The clearest case was birdsong. Birds were the model or­ganism for many, perhaps most, of the early ethologists, people like Konrad Lorenz and Niko Tinbergen. Many aspects of bird behavior are quite ame­nable to experimental study, and most especially their songs. It soon be­came clear that elements of the songs of many birds are socially learned, and social learning seemed the most plausible explanation for the spread of a technique by which blue tits opened the tops of British milk bottles. Birds seemed to have culture.

In the 1950s and 1960s, another group of animals also began to receive the culture label, the primates. This development occurred first in Japan, where, both in society generally and among scientists, the dichotomy be­tween humans and other primates is much less strict than in the European Christian tradition. Japanese scientists noted socially learned traditions in groups of Japanese macaques, most famously the spread of sweet potato washing among the monkeys on Koshima Island, where they dunked the tubers in the ocean to remove sand before eating them. The Japanese and other scientists were usually cautious in discussing these patterns, often referring to them as “precultural” behavior, “protoculture,” or “traditions” rather than unqualified “culture.”

The study of primate culture moved to another level in 1978 when William McGrew and Dorothy Tutin described variation in the grooming handclasp between groups of chimpanzees. Unlike the Japanese monkey traditions this was “arbitrary” behavior not involved in resource extraction, and McGrew and Tutin addressed the question of culture head on. They concluded that the evidence on grooming handclasps satisfied most but not all of eight conditions they listed for culture. Over the next few years pri­mate behavior was increasingly described as “culture,” without qualifiers. A highlight of this period was the publication of McGrew’s book "Chimpanzee Material Culture" in 1992. In this book and in his papers, McGrew showed the value of making systematic comparisons both across social groups and for a range of types of behavior, what he called an “ethnographic” approach. To virtually all field biologists studying chimpanzees, as well as orangutans, capuchin monkeys, and some other species, this approach made sense. It was clear that the animals that they had spent so much time with learned from each other, that they had culture, and that comparing what happened in different groups or at different times was a good way to look at this. How­ever, not everyone was happy with McGrew’s ethnographic approach.

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The promotion of culture as an important influence on behavior in chim­panzees and other nonhumans was attacked by two prominent psycholo­gists. Jeff Galef found little evidence that nonhumans either teach or imitate, and, given that he felt that culture propagated only through these processes, which he argued were different in humans compared to any other animal, he concluded that it is misleading to think of the evolution of culture in ani­mals. Galef also emphasized that culture in humans and what others call culture in animals were analogous—that is, evolved independently—rather than homologous—that is, similar through common descent. While the homology-analogy contrast is clearly a potentially important issue for those interested in chimpanzee culture, because of the recent common ancestors of humans and chimpanzees, from our cetological perspective this is an un­necessary controversy. The common ancestor of humans and whales was a small, probably fairly solitary, mammal, likely without much, if any, culture. The cultures of whales and those of humans, or chimpanzees, are analo­gous not homologous, and the fact that they evolved independently makes their similarities and differences particularly interesting. The second major attack on nonhuman culture came from Michael Tomasello. He reinforced Galef’s arguments, adding the potentially important point that of all the social learning processes only imitation and teaching can lead to cumula­tive cultures, which, as we have noted, are one of the most key attributes of human culture. Galef and Tomasello come from a null-hypothesis-testing, experimental psychology background. The null hypothesis is something like “chimpanzees do not possess culture,” with culture being defined by something like “traditional behavior transmitted by imitation or teaching.” They could not show in their own or others’ experimental studies that captive chimpanzees could imitate or teach, so did not reject the null hypothe­sis. No culture.

The “chimpanzee culture wars” were on. The field scientists responded, most prominently in a remarkable paper published in the journal Nature in 1999. The study, headed by psychologist Andrew Whiten, charted the inci­dence of thirty-nine chimpanzee cultural behaviors at seven study sites and concluded that “the combined repertoire of these behavior patterns in each chimpanzee community is itself highly distinctive, a phenomenon charac­teristic of human cultures.” Their basis for saying that these behaviors were cultural was that they did not seem obviously linked to some variation in ecology (that might guide individual learning along different paths in dif­ferent places) nor could it be explained by obvious genetic differences, an argument termed the “method of exclusion.” Similar studies of orangutans and capuchin monkeys followed.

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Unlike some anthropologists, the psychologists Galef and Tomasello had been specific in their criticism of animal culture. They set out argu­ments, hypotheses really, that could be falsified. For instance, their claim that chimpanzees did not imitate could be refuted. It subsequently was refuted—chimpanzees do imitate—in several studies, including some by Tomasello and his colleagues. However, Galef and Tomasello believe that other criticisms stand. For instance, Tomasello still maintains that there has been no “convincing demonstration of the ratchet effect or any other form of cumulative cultural evolution for chimpanzees or any other non­human animal.”

While this culture war was ostensibly about nonhuman cultures in gen­eral, it was, and is, highly focused on chimpanzees and, to a lesser extent, on other primates. All the evidence on the culture of birds, especially the detailed studies of birdsong, was largely and rather unfairly put aside, per­haps on the basis that birds are perceived as “one-trick ponies” with just one cultural behavior—song—both by the proponents and opponents of nonhuman culture. Whales were also initially outside the discussion.

In 2001, though, we wrote a review titled “Culture in Whales and Dol­phins,” and cetaceans joined the fray. Our paper attracted commentaries from those on both sides of the chimpanzee culture wars. The proponents of chimpanzee culture were generally positive about whale culture. For in­stance, Christophe Boesch wrote: “The sacrilegious proposition of the exis­tence of cultures in whales and dolphins should open the discussion of cultures in other animals, allowing us to find what is unique in human cul­tures.” And Andrew Whiten, who was the primary author of the famous 1999 chimpanzee culture paper, believed that we actually underestimated the imitative ability of at least some cetaceans.

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In contrast, those who did not like chimpanzee culture were far from convinced by our arguments for whale culture. The anthropologist, Tim Ingold wrote: “It is sad to see such rich empirical material, about such won­derful creatures, harnessed to such an impoverished theoretical agenda,” by which he specifically referred to our use of the word “ethnography” but, more broadly, seemed to dislike the dual inheritance perspective that we embraced. Jeff Galef, following his critiques of chimpanzee culture, went on about analogs and homologues of human culture, as well as a lack of conclusive evidence for imitation or teaching. We pushed back, titling our response to the commentaries, with tongues somewhat in cheeks, “Ceta­cean Culture: Still Afloat after the First Naval Engagement of the Culture Wars.”

An important criticism made by culture skeptics concerns how difficult it is to really know what causes behavior to develop in wild animals. Cul­tural transmission can cause variation in behavior within a species. It is not enough, though, simply to demonstrate that animals in one place, or one group, behave differently to those in other places or groups. Environ­mental variation can also produce differences in behavior between com­munities in several ways. For instance, if animals can learn a behavior with no social input, then in places with a particular food or tool present these may get used, whereas their use will be absent from communities where the necessary food or tool does not occur. This behavioral variation is not cul­ture; it results from individuals learning in different ways on their own. Ge­netic variation can also lead to differences in behavior. For an example we’ll take gibbon romance. Gibbon couples across Southeast Asia sing duets together in the mornings. The songs of these apes vary systematically be­tween populations. However, this is not the result of cultural transmission. There are strong correlations between genetic and vocal differences be­tween populations. Most crucially, when gibbons of different species mate, the songs their offspring produce are intermediate hybrids of their parents’ songs. Thus there is little role for cultural transmission in their develop­ment. All of this makes the method of exclusion a tricky beast, which for intellectual health and safety must always be labeled “Handle with care!”

There have been several attempts at peacekeeping in the culture wars. Kevin Laland and his colleagues have probably been most effective. They have criticized as well as commended those on both sides of the issue. Laland was the primary editor of the 2009 book "The Question of Animal Cul­ture," which brought the views of the pro- and anti-animal culture scien­tists together (although birds were almost completely ignored). They rec­ommend a broad and simple definition of culture—basically the one that we have adopted—as being most useful. Such a definition shifts attention from whether a particular behavior type fulfills all the conditions for “cul­ture” to the types of cultural behavior that seem to be present. However, Laland and his colleagues also criticized some of the inferences that pro-culture scientists have made from their data, suggesting alternative noncul­tural explanations. For instance they propose that some of the variation in the thirty-nine “cultural” chimpanzee behavior patterns might be, at least partially, ecologically or genetically determined, through processes such as we’ve just discussed. As an alternative, they argue strongly that we must develop better methods of studying culture in both wild and captive ani­mals. We must, they rightly caution, be wary of slipping into simplistic thinking about potential causes for a given behavior. If we force ourselves to decide that it’s either ecology, or genes, or culture, then we are doomed to fail in understanding behavioral development. For example, one of the most powerful benefits we get from our culture is an ability to adapt flexibly to the prevailing ecological conditions, be it in the Arctic or the jungle. We should expect in some cases a relationship between behavior and ecology even if the behavior is culturally transmitted. We therefore need to get away from the “either culture or genes or environment” trichotomy and toward asking, “How important are culture, genes, and the environment” in the development of behavior? This is, however, rather challenging if you are talking about large, wild animals that you can only partially observe part of the time, such as most cetaceans. Laland and his colleagues have them­selves worked to develop methods to overcome this challenge, as we have ourselves, and we’ll meet some of these techniques later on.

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We have got ourselves an idea of what we mean by “culture.” We have seen how controversial the notion that nonhumans might have culture is in some quarters. We have also seen how quickly things are changing in the way we understand these issues. We have a way to go before we have a good methodology for identifying the importance of culture to wild animals, but over the past decade the picture has changed in favor of nonhumans of some species possessing culture. Without the information they learn from each other, their behavior would be very different. We hope that scholars with as much insight as Robert Boyd and Joe Henrich would not write today, “Unlike other animal species, much of the variation among human groups is cultural,” as they did in 1998.

There has been a lot to chew over in this chapter. Let’s hope, though, we’ve now built ourselves a not-too-leaky vessel of ideas to carry us out to where the fun really starts—at sea, with the dolphins and whales.

Excerpted from “The Cultural Lives of Whales and Dolphins” by Hal Whitehead and Luke Rendell. Copyright © 2014 by Hal Whitehead and Luke Rendell. Reprinted by arrangement with University Of Chicago Press. All rights reserved.