Cetacean Culture Research Articles

The Racialization of Killer Whales: An Application of Gene-Culture Coevolutionary Theory

Abstract An expanding body of research aims to identify culture in cetaceans, often positing killer whales as an exemplar species. To this end, gene-culture coevolutionary theory provides a conceptual language with which whales are discussed in raciological terms. It renders killer whale ecotypes as discrete cultures that are intrinsically xenophobic and evolutionarily divergent. Such research on whale culture intends to substantiate theories of divergent natural selection between human cultures as well. This effort furthers the essentialism, simultaneously biological and cultural, that has long impelled colonial technoscientific projects of racial typologization. The exchange between the raciologies of humans and killer whales is facilitated by the infrahumanizing frameworks of gene-culture coevolutionary theory that locate whale populations alongside indigenous peoples on a spectrum of cultural simplicity and complexity. At this convergence of the animal and the human, gene-culture coevolutionary theory posits indigenous peoples and killer whales as evidence for social Darwinist theories that dehistoricize and naturalize intercultural conflict and ethnicization. The extension of raciology beyond the human species signals an urgent need to decolonize the biological sciences.

Negotiating the BoundaryThe Question of Animal Culture. Edited by Kevin N. Laland and Bennett G. Galef. Cambridge, MA: Harvard University Press, 2009.

Negotiating the Boundary<i>The Question of Animal Culture</i>. Edited by Kevin N. Laland and Bennett G. Galef. Cambridge, MA: Harvard University Press, 2009.

CALF INSPECTIONS OF FISH CATCHES IN BOTTLENOSE DOLPHINS (TURSIOPS SP.): OPPORTUNITIES FOR OBLIQUE SOCIAL LEARNING?

In recent years, the study of social learning and culture in wild cetaceans has visibly increased (Rendell and Whitehead 2001, Mann and Sargeant 2003, Krutzen et al. 2005). Some of the best evidence for some type of social learning in wild cetaceans comes from studies of vocal traditions in killer whales (Deecke et al. 2000, Yurk et al. 2002), song change in humpback whales (Noad et al. 2000), coda dialects in sperm whales (Rendell and Whitehead 2005), whistle convergence in bottlenose dolphins (Smolker and Pepper 1999, Watwood et al. 2004), and foraging behaviors in bottlenose dolphins (Mann and Sargeant 2003, Sargeant et al. 2005). The mechanisms (e.g., local enhancement, imitation) of social transmission have received very little attention, largely because this requires well-controlled experiments, compounded further by the difficulties in studying marine mammals. In a broader sense, determining whether transmission is vertical (parent to offspring), oblique (older to younger), or horizontal (laterally to peers or similar age classes), even without precise determination of mechanisms, offers insights into the ecological and social demands of a species. For example, horizontal transmission of song types in humpback whales is consistent with the proposed competitive function of song (Noad et al. 2000, Cerchio et al. 2001), and the context of pod-specific calls in killer whales suggests that calls promote cohesiveness (Miller et al. 2004). To date, our work on Shark Bay bottlenose dolphins (Tursiops sp.) suggests that vertical transmission of foraging types from mother to calf are commonplace (Mann and Sargeant 2003), but other forms of transmission may also occur (Sargeant et al. 2005). We argue that habitat heterogeneity and bisexual philopatry favors vertical transmission, especially for daughters who, more than sons, maintain a range similar to the mother’s (Krutzen et al. 2004). For

Cetacean Culture?

Cetacean Culture?

Generation and partial characterization of a transformed cetacean cell line

A primary epithelial cell line, DK1, established from renal tissue of a spontaneously aborted female Atlantic bottlenose dolphin was transfected with linearized pSV3.neo, an SV40 virus-derived plasmid encoding large tumor antigen (Tag). Transfected cells were grown in cetacean culture medium supplemented with 400 μg/ml geneticin (G418), and individual clones were selected using cloning rings. DKN1 was the first clone to be evaluated for future research use, and has been continuously cultured for 8 years. Intracellular cytokeratin and the expression of Tag were determined in DKN1, and cell growth was evaluated under different concentrations of l-glutamine, glutathione, and N-acetylcysteine. DKN1 cells did not require high levels of l-glutamine as previously reported for cetacean cells, and addition of antioxidants at the concentrations used in this study (2.0 mM) decreased the rate of cell division. These data suggest strongly that these immortalized bottlenose dolphin epithelial cells have different levels of, and requirements for, glutathione than would be considered normal for terrestrial mammalian cells, do not require high levels of l-glutamine as previously suggested for dolphin cells, and exhibit decreased levels of cell growth and viability in high levels of the antioxidant GSH and its precursor, NAC.

Cetacean culture: Still afloat after the first naval engagement of the culture wars

Although the majority of commentators implicitly or explicitly accept that field data allow us to ascribe culture to whales, dolphins, and other nonhumans, there is no consensus. While we define culture as information or behaviour shared by a population or subpopulation which is acquired from conspecifics through some form of social learning, some commentators suggest restricting this by requiring imitation/teaching, human analogy, adaptiveness, stability across generations, progressive evolution (ratchetting), or specific functions. Such restrictions fall down because they either preclude the attribution of culture to nonhumans using currently available methods, or exclude parts of human culture. The evidence for cetacean culture is strong in some cases, but weak in others. The commentaries provide important information on the social learning abilities of bottlenose dolphins and some interesting speculation about the evolution of cetacean cultures and differences between the cultures of different taxa. We maintain that some attributes of cetacean culture are currently unknown outside humans. While experimental studies, both in the laboratory and in the wild, have an important role in the study of culture in whales and dolphins (for instance in determining whether dolphins have a Theory-of-Mind), the real treasures will be uncovered by long-term observational studies at sea using new approaches and technologies.

Remarks on whale cultures from a complex systems perspective

The target article provides stimulating evidence for culture in cetaceans but does not provide a coherent theoretical framework. We argue that a complex, adaptive systems approach not only can provide such a framework but also can contribute advanced data analysis and simulation methods. For humpback whale songs, we suggest the framework of “small-world networks” to model the observed spatio-temporal dynamics.

Cetacean culture: Slippery when wet

Cetaceans are likely candidates for social learning and culture. Meager experimental evidence suggests that some cetaceans possess the requisite cognitive skills for social learning. Equally sparse ethnographic data provide clues about possible outcomes of social learning. Although the available evidence is consistent with the notion of culture in cetaceans, caution is warranted due to the many gaps in the data.

Culture in cetaceans: Why put the cart before the horse?

Twisting definitions of a concept to serve one's purposes is rarely a fruitful exercise and the demonstration of culture is, necessarily, a two-step procedure: (1) documenting a behavioral difference between populations, (2) demonstrating that this difference has spread through the group by means of social learning. This will avoid putting the cart before the horse.

Getting at animal culture: The interface of experimental and ethnographic evidence in dolphins

While supporting the claim for culture in cetaceans, I suggest that Rendell and Whitehead's argument is potentially incomplete if based solely on ethnographic evidence. The notion of cetacean culture can also be explored experimentally. I hope to complement the authors' assertion by discussing dolphin brain and behavioural research findings, thus contributing to a more holistic argument for culture in dolphins.

Cultural transmission of behavior in animals: How a modern training technology uses spontaneous social imitation in cetaceans and facilitates social imitation in horses and dogs

Social learning and imitation is central to culture in cetaceans. The training technology used with cetaceans facilitates reinforcing imitation of one dolphin's behavior by another; the same technology, now widely used by pet owners, can lead to imitative learning in such unlikely species as dogs and horses. A capacity for imitation, and thus for cultural learning, may exist in many species.

On not drawing the line about culture: Inconsistencies in interpretation of nonhuman cultures

Defining culture as social learning means that culture is present in many birds and mammals, suggesting that cetacean culture is not so special and does not require special explanation. Contrary to their own claims, Rendell and Whitehead present culture as having variant forms in different species, and these forms seem inconsistently applied and compared across species.

Is cetacean social learning unique?

Studies on captive dolphins have shown that they are capable of social learning. However, ethnographic data are less conclusive and many examples given for social learning can be explained in other ways. Before we can claim that cetacean culture is unique we need more rigorous studies which are fortunately not as difficult as Rendell and Whitehead seem to think.

Cetacean culture: Humans of the sea?

Rendell and Whitehead adopt a weak definition of culture to allow low standards of evidence for marine mammals, but they do not adequately rule out genetic factors or individual versus social learning. They then use these low standards to argue that some whales have unique cultures only matched by humans. It would have been more helpful to specify data gaps and suggest critical tests.

Cetacean culture: Resisting myths and addressing lacunae

Assessments of cetacean behavior has been hampered by popular misconceptions and mythic imagery. Rendell and Whitehead argue persuasively for accepting the idea of cetacean culture. Approaches to “culture,” however, must resist positivist approaches that reaffirm the observable. Culture is also comprised of “lacunae” when organisms choose to resist or avoid behavioral patterns.

Cetacean culture: Definitions and evidence

Rendell and Whitehead have drawn attention to some striking cetacean behaviour patterns. However, the claims for “culture” are premature. Weak examples of cetacean social learning do not, in sum, provide strong evidence for culture. Other terms, such as social learning, vocal learning, imitation, and tradition may be applied in some cases without resorting to more complex and controversial terms.

A sound approach to the study of culture

Rendell and Whitehead's thorough review dispels notions that culture is an exclusive faculty of humans and higher primates. We applaud the authors, but differ with them regarding the evolution of cetacean culture, which we argue resulted from the availability of abundant but spatially and temporally patchy prey such as schooling fish. We propose two examples of gene-culture coevolution: (1) acoustic abilities and acoustic traditions, and (2) transmission of environmental information and longevity.

Individual foraging specializations in marine mammals: Culture and ecology

Rendell and Whitehead argue persuasively that individual foraging specializations, if socially learned, are examples of cetacean culture. However, they discount ecological variation experienced by individuals within a population as a factor in such behavior. I suggest that ecological variation may play an important role in individual foraging specializations and describe several ecological parameters that may help us understand the high frequency of this interesting behavior in the marine habitat.

The mimetic dolphin

Rendell and Whitehead note the necessary, complementary relationship between field and laboratory studies in other species, but conclude their article by de-emphasizing the role of laboratory findings in cetacean research. The ambiguity in field studies of cetaceans should argue for greater reliance on the laboratory, which has provided much of the available research supporting the hypothesis of cetacean culture.

Culture in whales and dolphins.

Studies of animal culture have not normally included a consideration of cetaceans. However, with several long-term field studies now maturing, this situation should change. Animal culture is generally studied by either investigating transmission mechanisms experimentally, or observing patterns of behavioural variation in wild populations that cannot be explained by either genetic or environmental factors. Taking this second, ethnographic, approach, there is good evidence for cultural transmission in several cetacean species. However, only the bottlenose dolphin (Tursiops) has been shown experimentally to possess sophisticated social learning abilities, including vocal and motor imitation; other species have not been studied. There is observational evidence for imitation and teaching in killer whales. For cetaceans and other large, wide-ranging animals, excessive reliance on experimental data for evidence of culture is not productive; we favour the ethnographic approach. The complex and stable vocal and behavioural cultures of sympatric groups of killer whales (Orcinus orca) appear to have no parallel outside humans, and represent an independent evolution of cultural faculties. The wide movements of cetaceans, the greater variability of the marine environment over large temporal scales relative to that on land, and the stable matrilineal social groups of some species are potentially important factors in the evolution of cetacean culture. There have been suggestions of gene-culture coevolution in cetaceans, and culture may be implicated in some unusual behavioural and life-history traits of whales and dolphins. We hope to stimulate discussion and research on culture in these animals.